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</ul> </li> <li id="toc-Publication" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Publication"> <div class="vector-toc-text"> 3 Publication </div> </a> <button aria-controls="toc-Publication-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Publication subsection </button> <ul id="toc-Publication-sublist" class="vector-toc-list"> <li id="toc-Time_taken_to_publish" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Time_taken_to_publish"> <div class="vector-toc-text"> 3.1 Time taken to publish </div> </a> <ul id="toc-Time_taken_to_publish-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Events_leading_to_publication:_"big_book"_manuscript" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Events_leading_to_publication:_"big_book"_manuscript"> <div class="vector-toc-text"> 3.2 Events leading to publication: "big book" manuscript </div> </a> <ul id="toc-Events_leading_to_publication:_"big_book"_manuscript-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Joint_publication_of_papers_by_Wallace_and_Darwin" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Joint_publication_of_papers_by_Wallace_and_Darwin"> <div class="vector-toc-text"> 3.3 Joint publication of papers by Wallace and Darwin </div> </a> <ul id="toc-Joint_publication_of_papers_by_Wallace_and_Darwin-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Abstract_of_Species_book" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Abstract_of_Species_book"> <div class="vector-toc-text"> 3.4 Abstract of Species book </div> </a> <ul id="toc-Abstract_of_Species_book-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Murray_as_publisher;_choice_of_title" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Murray_as_publisher;_choice_of_title"> <div class="vector-toc-text"> 3.5 Murray as publisher; choice of title </div> </a> <ul id="toc-Murray_as_publisher;_choice_of_title-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Publication_and_subsequent_editions" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Publication_and_subsequent_editions"> <div class="vector-toc-text"> 3.6 Publication and subsequent editions </div> </a> <ul id="toc-Publication_and_subsequent_editions-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Publication_outside_Great_Britain" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Publication_outside_Great_Britain"> <div class="vector-toc-text"> 3.7 Publication outside Great Britain </div> </a> <ul id="toc-Publication_outside_Great_Britain-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Content" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Content"> <div class="vector-toc-text"> 4 Content </div> </a> <button aria-controls="toc-Content-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Content subsection </button> <ul id="toc-Content-sublist" class="vector-toc-list"> <li id="toc-Title_pages_and_introduction" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Title_pages_and_introduction"> <div class="vector-toc-text"> 4.1 Title pages and introduction </div> </a> <ul id="toc-Title_pages_and_introduction-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Variation_under_domestication_and_under_nature" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Variation_under_domestication_and_under_nature"> <div class="vector-toc-text"> 4.2 Variation under domestication and under nature </div> </a> <ul id="toc-Variation_under_domestication_and_under_nature-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Struggle_for_existence,_natural_selection,_and_divergence" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Struggle_for_existence,_natural_selection,_and_divergence"> <div class="vector-toc-text"> 4.3 Struggle for existence, natural selection, and divergence </div> </a> <ul id="toc-Struggle_for_existence,_natural_selection,_and_divergence-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Variation_and_heredity" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Variation_and_heredity"> <div class="vector-toc-text"> 4.4 Variation and heredity </div> </a> <ul id="toc-Variation_and_heredity-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Difficulties_for_the_theory" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Difficulties_for_the_theory"> <div class="vector-toc-text"> 4.5 Difficulties for the theory </div> </a> <ul id="toc-Difficulties_for_the_theory-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Geological_record" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Geological_record"> <div class="vector-toc-text"> 4.6 Geological record </div> </a> <ul id="toc-Geological_record-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Geographic_distribution" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Geographic_distribution"> <div class="vector-toc-text"> 4.7 Geographic distribution </div> </a> <ul id="toc-Geographic_distribution-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Classification,_morphology,_embryology,_rudimentary_organs" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Classification,_morphology,_embryology,_rudimentary_organs"> <div class="vector-toc-text"> 4.8 Classification, morphology, embryology, rudimentary organs </div> </a> <ul id="toc-Classification,_morphology,_embryology,_rudimentary_organs-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Concluding_remarks" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Concluding_remarks"> <div class="vector-toc-text"> 4.9 Concluding remarks </div> </a> <ul id="toc-Concluding_remarks-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Structure,_style,_and_themes" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Structure,_style,_and_themes"> <div class="vector-toc-text"> 5 Structure, style, and themes </div> </a> <button aria-controls="toc-Structure,_style,_and_themes-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Structure, style, and themes subsection </button> <ul id="toc-Structure,_style,_and_themes-sublist" class="vector-toc-list"> <li id="toc-Nature_and_structure_of_Darwin's_argument" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Nature_and_structure_of_Darwin's_argument"> <div class="vector-toc-text"> 5.1 Nature and structure of Darwin's argument </div> </a> <ul id="toc-Nature_and_structure_of_Darwin's_argument-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Literary_style" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Literary_style"> <div class="vector-toc-text"> 5.2 Literary style </div> </a> <ul id="toc-Literary_style-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Human_evolution" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Human_evolution"> <div class="vector-toc-text"> 5.3 Human evolution </div> </a> <ul id="toc-Human_evolution-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Reception" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Reception"> <div class="vector-toc-text"> 6 Reception </div> </a> <button aria-controls="toc-Reception-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Reception subsection </button> <ul id="toc-Reception-sublist" class="vector-toc-list"> <li id="toc-Impact_on_the_scientific_community" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Impact_on_the_scientific_community"> <div class="vector-toc-text"> 6.1 Impact on the scientific community </div> </a> <ul id="toc-Impact_on_the_scientific_community-sublist" class="vector-toc-list"> <li id="toc-Impact_outside_Great_Britain" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Impact_outside_Great_Britain"> <div class="vector-toc-text"> 6.1.1 Impact outside Great Britain </div> </a> <ul id="toc-Impact_outside_Great_Britain-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Challenges_to_natural_selection" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Challenges_to_natural_selection"> <div class="vector-toc-text"> 6.1.2 Challenges to natural selection </div> </a> <ul id="toc-Challenges_to_natural_selection-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Impact_on_economic_and_political_debates" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Impact_on_economic_and_political_debates"> <div class="vector-toc-text"> 6.2 Impact on economic and political debates </div> </a> <ul id="toc-Impact_on_economic_and_political_debates-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Religious_attitudes" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Religious_attitudes"> <div class="vector-toc-text"> 6.3 Religious attitudes </div> </a> <ul id="toc-Religious_attitudes-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Modern_influence" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Modern_influence"> <div class="vector-toc-text"> 7 Modern influence </div> </a> <ul id="toc-Modern_influence-sublist" class="vector-toc-list"> </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"> 8 See also </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"> 9 References </div> </a> <button aria-controls="toc-References-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle References subsection </button> <ul id="toc-References-sublist" class="vector-toc-list"> <li id="toc-Works_cited" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Works_cited"> <div class="vector-toc-text"> 9.1 Works cited </div> </a> <ul id="toc-Works_cited-sublist" class="vector-toc-list"> </ul> </li> </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"> 10 Further reading </div> </a> <button aria-controls="toc-Further_reading-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Further reading subsection </button> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> <li id="toc-Contemporary_reviews" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Contemporary_reviews"> <div class="vector-toc-text"> 10.1 Contemporary reviews </div> </a> <ul id="toc-Contemporary_reviews-sublist" class="vector-toc-list"> </ul> </li> </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"> 11 External links </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" title="Table of Contents" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" > Toggle the table of contents </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">On the Origin of Species</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 76 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-76" aria-hidden="true" > 76 languages </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/On_the_Origin_of_Species" title="On the Origin of Species – Afrikaans" lang="af" hreflang="af" data-title="On the Origin of Species" data-language-autonym="Afrikaans" data-language-local-name="Afrikaans" class="interlanguage-link-target">Afrikaans</a></li><li class="interlanguage-link interwiki-ar badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://ar.wikipedia.org/wiki/%D8%A3%D8%B5%D9%84_%D8%A7%D9%84%D8%A3%D9%86%D9%88%D8%A7%D8%B9" title="أصل الأنواع – Arabic" lang="ar" hreflang="ar" data-title="أصل الأنواع" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target">العربية</a></li><li class="interlanguage-link interwiki-ast mw-list-item"><a href="https://ast.wikipedia.org/wiki/On_the_Origin_of_Species" title="On the Origin of Species – Asturian" lang="ast" hreflang="ast" data-title="On the Origin of Species" data-language-autonym="Asturianu" data-language-local-name="Asturian" class="interlanguage-link-target">Asturianu</a></li><li class="interlanguage-link interwiki-az mw-list-item"><a href="https://az.wikipedia.org/wiki/N%C3%B6vl%C9%99rin_m%C9%99n%C5%9F%C9%99yi" title="Növlərin mənşəyi – Azerbaijani" lang="az" hreflang="az" data-title="Növlərin mənşəyi" data-language-autonym="Azərbaycanca" data-language-local-name="Azerbaijani" class="interlanguage-link-target">Azərbaycanca</a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%85%E0%A6%A8_%E0%A6%A6%E0%A7%8D%E0%A6%AF_%E0%A6%85%E0%A6%B0%E0%A6%BF%E0%A6%9C%E0%A6%BF%E0%A6%A8_%E0%A6%85%E0%A6%AC_%E0%A6%B8%E0%A7%8D%E0%A6%AA%E0%A6%BF%E0%A6%B8%E0%A6%BF%E0%A6%B8" title="অন দ্য অরিজিন অব স্পিসিস – Bangla" lang="bn" hreflang="bn" data-title="অন দ্য অরিজিন অব স্পিসিস" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target">বাংলা</a></li><li class="interlanguage-link interwiki-zh-min-nan mw-list-item"><a href="https://zh-min-nan.wikipedia.org/wiki/L%C5%ABn_Ch%C3%A9ng_%C3%AA_Kh%C3%AD-go%C3%A2n" title="Lūn Chéng ê Khí-goân – Minnan" lang="nan" hreflang="nan" data-title="Lūn Chéng ê Khí-goân" data-language-autonym="閩南語 / Bân-lâm-gú" data-language-local-name="Minnan" class="interlanguage-link-target">閩南語 / Bân-lâm-gú</a></li><li class="interlanguage-link interwiki-be mw-list-item"><a href="https://be.wikipedia.org/wiki/%D0%9F%D0%B0%D1%85%D0%BE%D0%B4%D0%B6%D0%B0%D0%BD%D0%BD%D0%B5_%D0%B2%D1%96%D0%B4%D0%B0%D1%9E" title="Паходжанне відаў – Belarusian" lang="be" hreflang="be" data-title="Паходжанне відаў" data-language-autonym="Беларуская" data-language-local-name="Belarusian" class="interlanguage-link-target">Беларуская</a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%9F%D1%80%D0%BE%D0%B8%D0%B7%D1%85%D0%BE%D0%B4_%D0%BD%D0%B0_%D0%B2%D0%B8%D0%B4%D0%BE%D0%B2%D0%B5%D1%82%D0%B5" title="Произход на видовете – Bulgarian" lang="bg" hreflang="bg" data-title="Произход на видовете" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target">Български</a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/L%27origen_de_les_esp%C3%A8cies" title="L'origen de les espècies – Catalan" lang="ca" hreflang="ca" data-title="L'origen de les espècies" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target">Català</a></li><li class="interlanguage-link interwiki-cs badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://cs.wikipedia.org/wiki/O_p%C5%AFvodu_druh%C5%AF" title="O původu druhů – Czech" lang="cs" hreflang="cs" data-title="O původu druhů" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target">Čeština</a></li><li class="interlanguage-link interwiki-cy mw-list-item"><a href="https://cy.wikipedia.org/wiki/On_the_Origin_of_Species" title="On the Origin of Species – Welsh" lang="cy" hreflang="cy" data-title="On the Origin of Species" data-language-autonym="Cymraeg" data-language-local-name="Welsh" class="interlanguage-link-target">Cymraeg</a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Arternes_Oprindelse" title="Arternes Oprindelse – Danish" lang="da" hreflang="da" data-title="Arternes Oprindelse" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target">Dansk</a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/%C3%9Cber_die_Entstehung_der_Arten" title="Über die Entstehung der Arten – German" lang="de" hreflang="de" data-title="Über die Entstehung der Arten" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target">Deutsch</a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/Liikide_tekkimine" title="Liikide tekkimine – Estonian" lang="et" hreflang="et" data-title="Liikide tekkimine" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target">Eesti</a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%97_%CE%BA%CE%B1%CF%84%CE%B1%CE%B3%CF%89%CE%B3%CE%AE_%CF%84%CF%89%CE%BD_%CE%B5%CE%B9%CE%B4%CF%8E%CE%BD" title="Η καταγωγή των ειδών – Greek" lang="el" hreflang="el" data-title="Η καταγωγή των ειδών" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target">Ελληνικά</a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/El_origen_de_las_especies" title="El origen de las especies – Spanish" lang="es" hreflang="es" data-title="El origen de las especies" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target">Español</a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/La_origino_de_specioj" title="La origino de specioj – Esperanto" lang="eo" hreflang="eo" data-title="La origino de specioj" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target">Esperanto</a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Espezieen_jatorriaz" title="Espezieen jatorriaz – Basque" lang="eu" hreflang="eu" data-title="Espezieen jatorriaz" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target">Euskara</a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%AE%D8%A7%D8%B3%D8%AA%DA%AF%D8%A7%D9%87_%DA%AF%D9%88%D9%86%D9%87%E2%80%8C%D9%87%D8%A7" title="خاستگاه گونه‌ها – Persian" lang="fa" hreflang="fa" data-title="خاستگاه گونه‌ها" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target">فارسی</a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/L%27Origine_des_esp%C3%A8ces" title="L'Origine des espèces – French" lang="fr" hreflang="fr" data-title="L'Origine des espèces" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target">Français</a></li><li class="interlanguage-link interwiki-fy mw-list-item"><a href="https://fy.wikipedia.org/wiki/Oer_de_oarsprong_fan_de_soarten" title="Oer de oarsprong fan de soarten – Western Frisian" lang="fy" hreflang="fy" data-title="Oer de oarsprong fan de soarten" data-language-autonym="Frysk" data-language-local-name="Western Frisian" class="interlanguage-link-target">Frysk</a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/A_orixe_das_especies" title="A orixe das especies – Galician" lang="gl" hreflang="gl" data-title="A orixe das especies" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target">Galego</a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EC%A2%85%EC%9D%98_%EA%B8%B0%EC%9B%90" title="종의 기원 – Korean" lang="ko" hreflang="ko" data-title="종의 기원" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target">한국어</a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D5%8F%D5%A5%D5%BD%D5%A1%D5%AF%D5%B6%D5%A5%D6%80%D5%AB_%D5%AE%D5%A1%D5%A3%D5%B8%D6%82%D5%B4%D5%A8" title="Տեսակների ծագումը – Armenian" lang="hy" hreflang="hy" data-title="Տեսակների ծագումը" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target">Հայերեն</a></li><li class="interlanguage-link interwiki-hi mw-list-item"><a href="https://hi.wikipedia.org/wiki/%E0%A4%9C%E0%A5%80%E0%A4%B5%E0%A4%9C%E0%A4%BE%E0%A4%A4%E0%A4%BF_%E0%A4%95%E0%A4%BE_%E0%A4%89%E0%A4%A6%E0%A5%8D%E0%A4%AD%E0%A4%B5" title="जीवजाति का उद्भव – Hindi" lang="hi" hreflang="hi" data-title="जीवजाति का उद्भव" data-language-autonym="हिन्दी" data-language-local-name="Hindi" class="interlanguage-link-target">हिन्दी</a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Podrijetlo_vrsta" title="Podrijetlo vrsta – Croatian" lang="hr" hreflang="hr" data-title="Podrijetlo vrsta" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target">Hrvatski</a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Asal-usul_Spesies" title="Asal-usul Spesies – Indonesian" lang="id" hreflang="id" data-title="Asal-usul Spesies" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target">Bahasa Indonesia</a></li><li class="interlanguage-link interwiki-is mw-list-item"><a href="https://is.wikipedia.org/wiki/Uppruni_tegundanna" title="Uppruni tegundanna – Icelandic" lang="is" hreflang="is" data-title="Uppruni tegundanna" data-language-autonym="Íslenska" data-language-local-name="Icelandic" class="interlanguage-link-target">Íslenska</a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/L%27origine_delle_specie" title="L'origine delle specie – Italian" lang="it" hreflang="it" data-title="L'origine delle specie" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target">Italiano</a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%9E%D7%95%D7%A6%D7%90_%D7%94%D7%9E%D7%99%D7%A0%D7%99%D7%9D" title="מוצא המינים – Hebrew" lang="he" hreflang="he" data-title="מוצא המינים" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target">עברית</a></li><li class="interlanguage-link interwiki-jv mw-list-item"><a href="https://jv.wikipedia.org/wiki/On_the_Origin_of_Species" title="On the Origin of Species – Javanese" lang="jv" hreflang="jv" data-title="On the Origin of Species" data-language-autonym="Jawa" data-language-local-name="Javanese" class="interlanguage-link-target">Jawa</a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/On_the_Origin_of_Species" title="On the Origin of Species – Latin" lang="la" hreflang="la" data-title="On the Origin of Species" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target">Latina</a></li><li class="interlanguage-link interwiki-lv mw-list-item"><a href="https://lv.wikipedia.org/wiki/Sugu_izcel%C5%A1an%C4%81s" title="Sugu izcelšanās – Latvian" lang="lv" hreflang="lv" data-title="Sugu izcelšanās" data-language-autonym="Latviešu" data-language-local-name="Latvian" class="interlanguage-link-target">Latviešu</a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/R%C5%AB%C5%A1i%C5%B3_atsiradimas" title="Rūšių atsiradimas – Lithuanian" lang="lt" hreflang="lt" data-title="Rūšių atsiradimas" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target">Lietuvių</a></li><li class="interlanguage-link interwiki-nia mw-list-item"><a href="https://nia.wikipedia.org/wiki/On_the_Origin_of_Species" title="On the Origin of Species – Nias" lang="nia" hreflang="nia" data-title="On the Origin of Species" data-language-autonym="Li Niha" data-language-local-name="Nias" class="interlanguage-link-target">Li Niha</a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/A_fajok_eredete" title="A fajok eredete – Hungarian" lang="hu" hreflang="hu" data-title="A fajok eredete" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target">Magyar</a></li><li class="interlanguage-link interwiki-mk mw-list-item"><a href="https://mk.wikipedia.org/wiki/%D0%97%D0%B0_%D0%BF%D0%BE%D1%82%D0%B5%D0%BA%D0%BB%D0%BE%D1%82%D0%BE_%D0%BD%D0%B0_%D0%B2%D0%B8%D0%B4%D0%BE%D0%B2%D0%B8%D1%82%D0%B5" title="За потеклото на видовите – Macedonian" lang="mk" hreflang="mk" data-title="За потеклото на видовите" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target">Македонски</a></li><li class="interlanguage-link interwiki-ml mw-list-item"><a href="https://ml.wikipedia.org/wiki/%E0%B4%92%E0%B4%B1%E0%B4%BF%E0%B4%9C%E0%B4%BF%E0%B5%BB_%E0%B4%93%E0%B4%AB%E0%B5%8D_%E0%B4%B8%E0%B5%8D%E0%B4%AA%E0%B5%80%E0%B4%B7%E0%B5%80%E0%B4%B8%E0%B5%8D" title="ഒറിജിൻ ഓഫ് സ്പീഷീസ് – Malayalam" lang="ml" hreflang="ml" data-title="ഒറിജിൻ ഓഫ് സ്പീഷീസ്" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target">മലയാളം</a></li><li class="interlanguage-link interwiki-mr mw-list-item"><a href="https://mr.wikipedia.org/wiki/%E0%A4%9C%E0%A4%BE%E0%A4%A4%E0%A5%80%E0%A4%82%E0%A4%9A%E0%A5%8D%E0%A4%AF%E0%A4%BE_%E0%A4%89%E0%A4%A6%E0%A5%8D%E0%A4%97%E0%A4%AE%E0%A4%B5%E0%A4%B0" title="जातींच्या उद्गमवर – Marathi" lang="mr" hreflang="mr" data-title="जातींच्या उद्गमवर" data-language-autonym="मराठी" data-language-local-name="Marathi" class="interlanguage-link-target">मराठी</a></li><li class="interlanguage-link interwiki-arz mw-list-item"><a href="https://arz.wikipedia.org/wiki/%D8%A7%D8%B5%D9%84_%D8%A7%D9%84%D8%A7%D9%86%D9%88%D8%A7%D8%B9" title="اصل الانواع – Egyptian Arabic" lang="arz" hreflang="arz" data-title="اصل الانواع" data-language-autonym="مصرى" data-language-local-name="Egyptian Arabic" class="interlanguage-link-target">مصرى</a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Asal_Usul_Spesies" title="Asal Usul Spesies – Malay" lang="ms" hreflang="ms" data-title="Asal Usul Spesies" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target">Bahasa Melayu</a></li><li class="interlanguage-link interwiki-mwl mw-list-item"><a href="https://mwl.wikipedia.org/wiki/La_Ourige_de_las_Speces" title="La Ourige de las Speces – Mirandese" lang="mwl" hreflang="mwl" data-title="La Ourige de las Speces" data-language-autonym="Mirandés" data-language-local-name="Mirandese" class="interlanguage-link-target">Mirandés</a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/De_oorsprong_der_soorten" title="De oorsprong der soorten – Dutch" lang="nl" hreflang="nl" data-title="De oorsprong der soorten" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target">Nederlands</a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E7%A8%AE%E3%81%AE%E8%B5%B7%E6%BA%90" title="種の起源 – Japanese" lang="ja" hreflang="ja" data-title="種の起源" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target">日本語</a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Artenes_opprinnelse" title="Artenes opprinnelse – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Artenes opprinnelse" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target">Norsk bokmål</a></li><li class="interlanguage-link interwiki-nn mw-list-item"><a href="https://nn.wikipedia.org/wiki/On_the_Origin_of_Species" title="On the Origin of Species – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="On the Origin of Species" data-language-autonym="Norsk nynorsk" data-language-local-name="Norwegian Nynorsk" class="interlanguage-link-target">Norsk nynorsk</a></li><li class="interlanguage-link interwiki-oc mw-list-item"><a href="https://oc.wikipedia.org/wiki/L%27origina_de_las_esp%C3%A8cias" title="L'origina de las espècias – Occitan" lang="oc" hreflang="oc" data-title="L'origina de las espècias" data-language-autonym="Occitan" data-language-local-name="Occitan" class="interlanguage-link-target">Occitan</a></li><li class="interlanguage-link interwiki-pa mw-list-item"><a href="https://pa.wikipedia.org/wiki/%E0%A8%9C%E0%A9%80%E0%A8%B5_%E0%A8%AA%E0%A9%8D%E0%A8%B0%E0%A8%9C%E0%A8%BE%E0%A8%A4%E0%A9%80%E0%A8%86%E0%A8%82_%E0%A8%A6%E0%A9%80_%E0%A8%89%E0%A8%A4%E0%A8%AA%E0%A8%A4%E0%A9%80" title="ਜੀਵ ਪ੍ਰਜਾਤੀਆਂ ਦੀ ਉਤਪਤੀ – Punjabi" lang="pa" hreflang="pa" data-title="ਜੀਵ ਪ੍ਰਜਾਤੀਆਂ ਦੀ ਉਤਪਤੀ" data-language-autonym="ਪੰਜਾਬੀ" data-language-local-name="Punjabi" class="interlanguage-link-target">ਪੰਜਾਬੀ</a></li><li class="interlanguage-link interwiki-pnb mw-list-item"><a href="https://pnb.wikipedia.org/wiki/%D8%A7%D9%86%D9%88%D8%A7%D8%AD_%D8%AF%DB%8C_%D8%A7%D8%B5%D9%84" title="انواح دی اصل – Western Punjabi" lang="pnb" hreflang="pnb" data-title="انواح دی اصل" data-language-autonym="پنجابی" data-language-local-name="Western Punjabi" class="interlanguage-link-target">پنجابی</a></li><li class="interlanguage-link interwiki-ps mw-list-item"><a href="https://ps.wikipedia.org/wiki/%D8%AF_%DA%89%D9%88%D9%84%D9%88%D9%86%D9%88_%D8%A7%D8%B5%D9%84" title="د ډولونو اصل – Pashto" lang="ps" hreflang="ps" data-title="د ډولونو اصل" data-language-autonym="پښتو" data-language-local-name="Pashto" class="interlanguage-link-target">پښتو</a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/O_powstawaniu_gatunk%C3%B3w" title="O powstawaniu gatunków – Polish" lang="pl" hreflang="pl" data-title="O powstawaniu gatunków" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target">Polski</a></li><li class="interlanguage-link interwiki-pt badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://pt.wikipedia.org/wiki/A_Origem_das_Esp%C3%A9cies" title="A Origem das Espécies – Portuguese" lang="pt" hreflang="pt" data-title="A Origem das Espécies" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target">Português</a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/Originea_speciilor" title="Originea speciilor – Romanian" lang="ro" hreflang="ro" data-title="Originea speciilor" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target">Română</a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%9F%D1%80%D0%BE%D0%B8%D1%81%D1%85%D0%BE%D0%B6%D0%B4%D0%B5%D0%BD%D0%B8%D0%B5_%D0%B2%D0%B8%D0%B4%D0%BE%D0%B2" title="Происхождение видов – Russian" lang="ru" hreflang="ru" data-title="Происхождение видов" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target">Русский</a></li><li class="interlanguage-link interwiki-si mw-list-item"><a href="https://si.wikipedia.org/wiki/%E0%B6%94%E0%B6%B1%E0%B7%8A_%E0%B6%AF_%E0%B6%94%E0%B6%BB%E0%B7%92%E0%B6%A2%E0%B7%92%E0%B6%B1%E0%B7%8A_%E0%B6%94%E0%B7%86%E0%B7%8A_%E0%B7%83%E0%B7%8A%E0%B6%B4%E0%B7%93%E0%B7%82%E0%B7%93%E0%B7%83%E0%B7%8A" title="ඔන් ද ඔරිජින් ඔෆ් ස්පීෂීස් – Sinhala" lang="si" hreflang="si" data-title="ඔන් ද ඔරිජින් ඔෆ් ස්පීෂීස්" data-language-autonym="සිංහල" data-language-local-name="Sinhala" class="interlanguage-link-target">සිංහල</a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/On_the_Origin_of_Species" title="On the Origin of Species – Simple English" lang="en-simple" hreflang="en-simple" data-title="On the Origin of Species" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target">Simple English</a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/O_p%C3%B4vode_druhov" title="O pôvode druhov – Slovak" lang="sk" hreflang="sk" data-title="O pôvode druhov" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target">Slovenčina</a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/O_nastanku_vrst" title="O nastanku vrst – Slovenian" lang="sl" hreflang="sl" data-title="O nastanku vrst" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target">Slovenščina</a></li><li class="interlanguage-link interwiki-ckb mw-list-item"><a href="https://ckb.wikipedia.org/wiki/%D8%AF%DB%95%D8%B1%D8%A8%D8%A7%D8%B1%DB%95%DB%8C_%D8%B3%DB%95%D8%B1%DA%86%D8%A7%D9%88%DB%95%DB%8C_%D8%AC%DB%86%D8%B1%DB%95%DA%A9%D8%A7%D9%86" title="دەربارەی سەرچاوەی جۆرەکان – Central Kurdish" lang="ckb" hreflang="ckb" data-title="دەربارەی سەرچاوەی جۆرەکان" data-language-autonym="کوردی" data-language-local-name="Central Kurdish" class="interlanguage-link-target">کوردی</a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%9E_%D0%BF%D0%BE%D1%80%D0%B5%D0%BA%D0%BB%D1%83_%D0%B2%D1%80%D1%81%D1%82%D0%B0" title="О пореклу врста – Serbian" lang="sr" hreflang="sr" data-title="О пореклу врста" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target">Српски / srpski</a></li><li class="interlanguage-link interwiki-sh mw-list-item"><a href="https://sh.wikipedia.org/wiki/On_the_Origin_of_Species" title="On the Origin of Species – Serbo-Croatian" lang="sh" hreflang="sh" data-title="On the Origin of Species" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target">Srpskohrvatski / српскохрватски</a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Lajien_synty" title="Lajien synty – Finnish" lang="fi" hreflang="fi" data-title="Lajien synty" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target">Suomi</a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Om_arternas_uppkomst" title="Om arternas uppkomst – Swedish" lang="sv" hreflang="sv" data-title="Om arternas uppkomst" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target">Svenska</a></li><li class="interlanguage-link interwiki-tl mw-list-item"><a href="https://tl.wikipedia.org/wiki/On_the_Origin_of_Species" title="On the Origin of Species – Tagalog" lang="tl" hreflang="tl" data-title="On the Origin of Species" data-language-autonym="Tagalog" data-language-local-name="Tagalog" class="interlanguage-link-target">Tagalog</a></li><li class="interlanguage-link interwiki-ta mw-list-item"><a href="https://ta.wikipedia.org/wiki/%E0%AE%89%E0%AE%AF%E0%AE%BF%E0%AE%B0%E0%AE%BF%E0%AE%A9%E0%AE%99%E0%AF%8D%E0%AE%95%E0%AE%B3%E0%AE%BF%E0%AE%A9%E0%AF%8D_%E0%AE%A4%E0%AF%8B%E0%AE%B1%E0%AF%8D%E0%AE%B1%E0%AE%AE%E0%AF%8D_(%E0%AE%A8%E0%AF%82%E0%AE%B2%E0%AF%8D)" title="உயிரினங்களின் தோற்றம் (நூல்) – Tamil" lang="ta" hreflang="ta" data-title="உயிரினங்களின் தோற்றம் (நூல்)" data-language-autonym="தமிழ்" data-language-local-name="Tamil" class="interlanguage-link-target">தமிழ்</a></li><li class="interlanguage-link interwiki-kab mw-list-item"><a href="https://kab.wikipedia.org/wiki/I%E1%BA%93uran_n_telmas" title="Iẓuran n telmas – Kabyle" lang="kab" hreflang="kab" data-title="Iẓuran n telmas" data-language-autonym="Taqbaylit" data-language-local-name="Kabyle" class="interlanguage-link-target">Taqbaylit</a></li><li class="interlanguage-link interwiki-th mw-list-item"><a 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Click here for more information."><img alt="Featured article" src="//upload.wikimedia.org/wikipedia/en/thumb/e/e7/Cscr-featured.svg/20px-Cscr-featured.svg.png" decoding="async" width="20" height="19" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/e/e7/Cscr-featured.svg/30px-Cscr-featured.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/e/e7/Cscr-featured.svg/40px-Cscr-featured.svg.png 2x" data-file-width="466" data-file-height="443" /></a></div></div> </div> <div id="siteSub" class="noprint">From Wikipedia, the free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">1859 book on evolution by Charles Darwin</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">"Origin of Species" and "Origin of the species" redirect here. For other uses, see <a href="/wiki/Origin_of_Species_(disambiguation)" class="mw-disambig" title="Origin of Species (disambiguation)">Origin of Species (disambiguation)</a>.</div> <style data-mw-deduplicate="TemplateStyles:r1257001546">.mw-parser-output .infobox-subbox{padding:0;border:none;margin:-3px;width:auto;min-width:100%;font-size:100%;clear:none;float:none;background-color:transparent}.mw-parser-output .infobox-3cols-child{margin:auto}.mw-parser-output .infobox .navbar{font-size:100%}@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox-full-data:not(.notheme)>div:not(.notheme)[style]{background:#1f1f23!important;color:#f8f9fa}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox-full-data:not(.notheme) div:not(.notheme){background:#1f1f23!important;color:#f8f9fa}}@media(min-width:640px){body.skin--responsive .mw-parser-output .infobox-table{display:table!important}body.skin--responsive .mw-parser-output .infobox-table>caption{display:table-caption!important}body.skin--responsive .mw-parser-output .infobox-table>tbody{display:table-row-group}body.skin--responsive .mw-parser-output .infobox-table tr{display:table-row!important}body.skin--responsive .mw-parser-output .infobox-table th,body.skin--responsive .mw-parser-output .infobox-table td{padding-left:inherit;padding-right:inherit}}</style><table class="infobox vcard"><caption class="infobox-title" style="font-size:125%; font-style:italic; padding-bottom:0.2em;">On the Origin of Species <span class="Z3988" title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=On+the+Origin+of+Species&rft.author=%5B%5BCharles+Darwin%5D%5D&rft.place=%5B%5BUnited+Kingdom+of+Great+Britain+and+Ireland%5D%5D&rft.pages=502&rft_id=info:oclcnum/352242"></caption><tbody><tr><td colspan="2" class="infobox-image"><a href="/wiki/File:Origin_of_Species_title_page.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/cd/Origin_of_Species_title_page.jpg/220px-Origin_of_Species_title_page.jpg" decoding="async" width="220" height="353" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/cd/Origin_of_Species_title_page.jpg/330px-Origin_of_Species_title_page.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/cd/Origin_of_Species_title_page.jpg/440px-Origin_of_Species_title_page.jpg 2x" data-file-width="2888" data-file-height="4636" /></a><div class="infobox-caption">Title page of the 1859 edition<a href="#cite_note-1">[1]</a></div></td></tr><tr><th scope="row" class="infobox-label">Author</th><td class="infobox-data"><a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a></td></tr><tr><th scope="row" class="infobox-label">Language</th><td class="infobox-data">English</td></tr><tr><th scope="row" class="infobox-label">Subject</th><td class="infobox-data"><a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a> <a href="/wiki/Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></td></tr><tr><th scope="row" class="infobox-label">Published</th><td class="infobox-data">24 November 1859<a href="#cite_note-Freeman_1977-2">[2]</a> (<a href="/wiki/John_Murray_(publishing_house)" title="John Murray (publishing house)">John Murray</a>)</td></tr><tr><th scope="row" class="infobox-label">Publication place</th><td class="infobox-data"><a href="/wiki/United_Kingdom_of_Great_Britain_and_Ireland" title="United Kingdom of Great Britain and Ireland">United Kingdom of Great Britain and Ireland</a></td></tr><tr><th scope="row" class="infobox-label">Media type</th><td class="infobox-data">Print (<a href="/wiki/Hardcover" title="Hardcover">hardback</a>)</td></tr><tr><th scope="row" class="infobox-label">Pages</th><td class="infobox-data">502</td></tr><tr><th scope="row" class="infobox-label"><a href="/wiki/OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)"><abbr title="Online Computer Library Center number">OCLC</abbr></a></th><td class="infobox-data"><a rel="nofollow" class="external text" href="https://www.worldcat.org/oclc/352242">352242</a></td></tr><tr><th scope="row" class="infobox-label">Preceded by</th><td class="infobox-data"><a href="/wiki/On_the_Tendency_of_Species_to_form_Varieties;_and_on_the_Perpetuation_of_Varieties_and_Species_by_Natural_Means_of_Selection" title="On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection">On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection</a> </td></tr><tr><th scope="row" class="infobox-label">Followed by</th><td class="infobox-data"><a href="/wiki/Fertilisation_of_Orchids" title="Fertilisation of Orchids">Fertilisation of Orchids</a> </td></tr><tr><th scope="row" class="infobox-label">Text</th><td class="infobox-data"><a href="https://en.wikisource.org/wiki/On_the_Origin_of_Species" class="extiw" title="s:On the Origin of Species">On the Origin of Species</a> at <a href="/wiki/Wikisource" title="Wikisource">Wikisource</a></td></tr></tbody></table> <style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline ol,.mw-parser-output .hlist.inline ul,.mw-parser-output .hlist dl dl,.mw-parser-output .hlist dl ol,.mw-parser-output .hlist dl ul,.mw-parser-output .hlist ol dl,.mw-parser-output .hlist ol ol,.mw-parser-output .hlist ol ul,.mw-parser-output .hlist ul dl,.mw-parser-output .hlist ul ol,.mw-parser-output .hlist ul ul{display:inline}.mw-parser-output .hlist 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sidebar-collapse nomobile nowraplinks" style="border:2px solid #90ee90"><tbody><tr><td class="sidebar-pretitle">Part of a series on</td></tr><tr><th class="sidebar-title-with-pretitle" 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"><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><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 mw-collapsible-content plainlist"><div class="hlist"> <ul><li><a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">Overview</a></li> <li><a href="/wiki/Evolutionary_ideas_of_the_Renaissance_and_Enlightenment" title="Evolutionary ideas of the Renaissance and Enlightenment">Renaissance</a></li> <li><a href="/wiki/Transmutation_of_species" title="Transmutation of species">Before Darwin</a></li> <li><a href="/wiki/Charles_Darwin" title="Charles Darwin">Darwin</a></li> <li><a class="mw-selflink selflink">Origin of Species</a></li> <li><a href="/wiki/The_eclipse_of_Darwinism" title="The eclipse of Darwinism">Before synthesis</a></li> <li><a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis</a></li> <li><a href="/wiki/History_of_molecular_evolution" title="History of molecular evolution">Molecular evolution</a></li> <li><a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evo-devo</a></li> <li><a href="/wiki/Current_research_in_evolutionary_biology" class="mw-redirect" title="Current research in evolutionary biology">Current research</a></li> <li><a href="/wiki/History_of_speciation" title="History of speciation">History of speciation</a></li> <li><a href="/wiki/History_of_paleontology" title="History of paleontology">History of paleontology</a> (<a href="/wiki/Timeline_of_paleontology" title="Timeline of paleontology">timeline</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)">Fields and applications</div><div class="sidebar-list-content mw-collapsible-content plainlist"> <ul><li><a href="/wiki/Applications_of_evolution" title="Applications of evolution">Applications of evolution</a></li> <li><a href="/wiki/Biosocial_criminology" title="Biosocial criminology">Biosocial criminology</a></li> <li><a href="/wiki/Ecological_genetics" title="Ecological genetics">Ecological genetics</a></li> <li><a href="/wiki/Evolutionary_aesthetics" title="Evolutionary aesthetics">Evolutionary aesthetics</a></li> <li><a href="/wiki/Evolutionary_anthropology" title="Evolutionary anthropology">Evolutionary anthropology</a></li> <li><a href="/wiki/Evolutionary_computation" title="Evolutionary computation">Evolutionary computation</a></li> <li><a href="/wiki/Evolutionary_ecology" title="Evolutionary ecology">Evolutionary ecology</a></li> <li><a href="/wiki/Evolutionary_economics" title="Evolutionary economics">Evolutionary economics</a></li> <li><a href="/wiki/Evolutionary_epistemology" title="Evolutionary epistemology">Evolutionary epistemology</a></li> <li><a href="/wiki/Evolutionary_ethics" title="Evolutionary ethics">Evolutionary ethics</a></li> <li><a href="/wiki/Evolutionary_game_theory" title="Evolutionary game theory">Evolutionary game theory</a></li> <li><a href="/wiki/Evolutionary_linguistics" title="Evolutionary linguistics">Evolutionary linguistics</a></li> <li><a href="/wiki/Evolutionary_medicine" title="Evolutionary medicine">Evolutionary medicine</a></li> <li><a href="/wiki/Evolutionary_neuroscience" title="Evolutionary neuroscience">Evolutionary neuroscience</a></li> <li><a href="/wiki/Evolutionary_physiology" title="Evolutionary physiology">Evolutionary physiology</a></li> <li><a href="/wiki/Evolutionary_psychology" title="Evolutionary psychology">Evolutionary psychology</a></li> <li><a href="/wiki/Experimental_evolution" title="Experimental evolution">Experimental evolution</a></li> <li><a href="/wiki/Invasion_genetics" title="Invasion genetics">Invasion genetics</a></li> <li><a href="/wiki/Island_biogeography" class="mw-redirect" title="Island biogeography">Island biogeography</a></li> <li><a href="/wiki/Phylogenetics" title="Phylogenetics">Phylogenetics</a></li> <li><a href="/wiki/Paleontology" title="Paleontology">Paleontology</a></li> <li><a href="/wiki/Selective_breeding" title="Selective breeding">Selective breeding</a></li> <li><a href="/wiki/Laboratory_experiments_of_speciation" title="Laboratory experiments of speciation">Speciation experiments</a></li> <li><a href="/wiki/Sociobiology" title="Sociobiology">Sociobiology</a></li> <li><a href="/wiki/Systematics" title="Systematics">Systematics</a></li> <li><a href="/wiki/Universal_Darwinism" title="Universal Darwinism">Universal Darwinism</a></li></ul></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)">Social implications</div><div class="sidebar-list-content mw-collapsible-content plainlist"> <ul><li><a href="/wiki/Eugenics" title="Eugenics">Eugenics</a></li> <li><a href="/wiki/Evolution_as_fact_and_theory" title="Evolution as fact and theory">Evolution as fact and theory</a></li> <li><a href="/wiki/Dysgenics" title="Dysgenics">Dysgenics</a></li> <li><a href="/wiki/Social_effects_of_evolutionary_theory" title="Social effects of evolutionary theory">Social effects</a></li> <li><a href="/wiki/Rejection_of_evolution_by_religious_groups" title="Rejection of evolution by religious groups">Creation–evolution controversy</a></li> <li><a href="/wiki/Theistic_evolution" title="Theistic evolution">Theistic evolution</a></li> <li><a href="/wiki/Objections_to_evolution" title="Objections to evolution">Objections to evolution</a></li> <li><a href="/wiki/Level_of_support_for_evolution" title="Level of support for evolution">Level of support</a></li> <li><a href="/wiki/Nature_versus_nurture" title="Nature versus nurture">Nature-nurture controversy</a></li></ul></div></div></td> </tr><tr><td class="sidebar-below hlist"> <ul><li><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> <a href="/wiki/Portal:Evolutionary_biology" title="Portal:Evolutionary biology">Evolutionary biology portal</a></li> <li><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" /> <a href="/wiki/Category:Evolutionary_biology" title="Category:Evolutionary biology">Category</a></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> On the Origin of Species (or, more completely, On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life)<a href="#cite_note-titles-3">[3]</a> is a work of <a href="/wiki/Scientific_literature" title="Scientific literature">scientific literature</a> by <a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a> that is considered to be the foundation of <a href="/wiki/Evolutionary_biology" title="Evolutionary biology">evolutionary biology</a>. It was published on 24 November 1859.<a href="#cite_note-FOOTNOTEDesmondMoore1991477-4">[4]</a> Darwin's book introduced the <a href="/wiki/Scientific_theory" title="Scientific theory">scientific theory</a> that populations <a href="/wiki/Evolution" title="Evolution">evolve</a> over the course of generations through a process of <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a>, although <a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a> was also included as a mechanism of lesser importance. The book presented a body of evidence that <a href="/wiki/Biodiversity" title="Biodiversity">the diversity of life</a> arose by <a href="/wiki/Common_descent" title="Common descent">common descent</a> through a <a href="/wiki/Tree_of_life_(science)" class="mw-redirect" title="Tree of life (science)">branching pattern of evolution</a>. Darwin included evidence that he had collected on <a href="/wiki/Second_voyage_of_HMS_Beagle" title="Second voyage of HMS Beagle">the Beagle expedition</a> in the 1830s and his subsequent findings from research, correspondence, and experimentation.<a href="#cite_note-5">[5]</a> <a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">Various evolutionary ideas</a> had already been proposed to explain <a href="/wiki/History_of_biology" title="History of biology">new findings in biology</a>. There was growing support for such ideas among dissident anatomists and the general public, but during the first half of the 19th century the English scientific establishment was closely tied to the <a href="/wiki/Church_of_England" title="Church of England">Church of England</a>, while science was part of <a href="/wiki/Natural_theology" title="Natural theology">natural theology</a>. Ideas about the <a href="/wiki/Transmutation_of_species" title="Transmutation of species">transmutation of species</a> were controversial as they conflicted with the beliefs that species were unchanging parts of a designed hierarchy and that humans were unique, unrelated to other animals. The political and theological implications were intensely debated, but transmutation was not accepted by the scientific mainstream. The book was written for non-specialist readers and attracted widespread interest upon its publication. Darwin was already highly regarded as a scientist, so his findings were taken seriously and the evidence he presented generated scientific, philosophical, and religious discussion. The debate over the book contributed to the campaign by <a href="/wiki/Thomas_Henry_Huxley" title="Thomas Henry Huxley">T. H. Huxley</a> and his fellow members of the <a href="/wiki/X_Club" title="X Club">X Club</a> to <a href="/wiki/Secularization" title="Secularization">secularise</a> science by promoting <a href="/wiki/Naturalism_(philosophy)" title="Naturalism (philosophy)">scientific naturalism</a>. Within two decades, there was widespread scientific agreement that evolution, with a branching pattern of common descent, had occurred, but scientists were slow to give natural selection the significance that Darwin thought appropriate. During "<a href="/wiki/The_eclipse_of_Darwinism" title="The eclipse of Darwinism">the eclipse of Darwinism</a>" from the 1880s to the 1930s, various other mechanisms of evolution were given more credit. With the development of the <a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">modern evolutionary synthesis</a> in the 1930s and 1940s, Darwin's concept of evolutionary <a href="/wiki/Adaptation" title="Adaptation">adaptation</a> through natural selection became central to modern evolutionary theory, and it has now become the unifying concept of the <a href="/wiki/Life_science" class="mw-redirect" title="Life science">life sciences</a>. <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Summary_of_Darwin's_theory">Summary of Darwin's theory</h2>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=1" title="Edit section: Summary of Darwin's theory">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Charles_Darwin_seated.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/86/Charles_Darwin_seated.jpg/170px-Charles_Darwin_seated.jpg" decoding="async" width="170" height="240" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/86/Charles_Darwin_seated.jpg/255px-Charles_Darwin_seated.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/86/Charles_Darwin_seated.jpg/340px-Charles_Darwin_seated.jpg 2x" data-file-width="2448" data-file-height="3456" /></a><figcaption>Darwin pictured shortly before publication</figcaption></figure> Darwin's theory of evolution is based on key <a href="/wiki/Fact" title="Fact">facts</a> and the <a href="/wiki/Inference" title="Inference">inferences</a> drawn from them, which biologist <a href="/wiki/Ernst_Mayr" title="Ernst Mayr">Ernst Mayr</a> summarised as follows:<a href="#cite_note-6">[6]</a> <dl><dd><ul><li>Every <a href="/wiki/Species" title="Species">species</a> is fertile enough that if all offspring survived to reproduce, the population would grow (fact).</li> <li>Despite periodic fluctuations, populations remain roughly the same size (fact).</li> <li>Resources such as food are limited and are relatively stable over time (fact).</li> <li>A <a href="/wiki/Struggle_for_existence" title="Struggle for existence">struggle for survival</a> ensues (inference).</li> <li>Individuals in a population vary significantly from one another (fact).</li> <li>Much of this variation is <a href="/wiki/Heredity" title="Heredity">heritable</a> (fact).</li> <li>Individuals less suited to the environment are less likely to survive and less likely to reproduce; individuals more suited to the environment are more likely to survive and more likely to reproduce and leave their heritable traits to future generations, which produces the process of <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a> (fact).</li> <li>This slowly effected process results in populations changing to adapt to their environments, and ultimately, these variations accumulate over time to form new species (inference).</li></ul></dd></dl> <div class="mw-heading mw-heading2"><h2 id="Background">Background</h2>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=2" title="Edit section: Background">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">History of evolutionary thought</a> and <a href="/wiki/History_of_biology" title="History of biology">History of biology</a></div> <div class="mw-heading mw-heading3"><h3 id="Developments_before_Darwin's_theory">Developments before Darwin's theory</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=3" title="Edit section: Developments before Darwin's theory">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Georges_Cuvier_-_Elephant_jaw.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d2/Georges_Cuvier_-_Elephant_jaw.png/170px-Georges_Cuvier_-_Elephant_jaw.png" decoding="async" width="170" height="300" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d2/Georges_Cuvier_-_Elephant_jaw.png/255px-Georges_Cuvier_-_Elephant_jaw.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d2/Georges_Cuvier_-_Elephant_jaw.png/340px-Georges_Cuvier_-_Elephant_jaw.png 2x" data-file-width="600" data-file-height="1060" /></a><figcaption>Cuvier's 1799 paper on living and fossil elephants helped establish the reality of <a href="/wiki/Extinction" title="Extinction">extinction</a>.</figcaption></figure> In later editions of the book, Darwin traced evolutionary ideas as far back as <a href="/wiki/Aristotle" title="Aristotle">Aristotle</a>;<a href="#cite_note-7">[7]</a> the text he cites is a summary by Aristotle of the ideas of the earlier Greek philosopher <a href="/wiki/Empedocles" title="Empedocles">Empedocles</a>.<a href="#cite_note-8">[8]</a> Early <a href="/wiki/Christianity" title="Christianity">Christian</a> <a href="/wiki/Church_Fathers" title="Church Fathers">Church Fathers</a> and <a href="/wiki/Medieval_Europe" class="mw-redirect" title="Medieval Europe">Medieval European</a> scholars interpreted the <a href="/wiki/Genesis_creation_narrative" title="Genesis creation narrative">Genesis creation narrative</a> allegorically rather than as a literal historical account;<a href="#cite_note-9">[9]</a> organisms were described by their mythological and heraldic significance as well as by their physical form. Nature was widely believed to be unstable and capricious, with monstrous births from union between species, and <a href="/wiki/Spontaneous_generation" title="Spontaneous generation">spontaneous generation</a> of life.<a href="#cite_note-10">[10]</a> The <a href="/wiki/Protestant_Reformation" class="mw-redirect" title="Protestant Reformation">Protestant Reformation</a> inspired <a href="/wiki/Biblical_literalism" title="Biblical literalism">a literal interpretation of the Bible</a>, with concepts of creation that conflicted with the findings of an <a href="/wiki/Scientific_Revolution" title="Scientific Revolution">emerging science</a> seeking explanations congruent with the <a href="/wiki/Mechanism_(philosophy)" title="Mechanism (philosophy)">mechanical philosophy</a> of <a href="/wiki/Ren%C3%A9_Descartes" title="René Descartes">René Descartes</a> and the <a href="/wiki/Empiricism" title="Empiricism">empiricism</a> of the <a href="/wiki/Baconian_method" title="Baconian method">Baconian method</a>. After the turmoil of the <a href="/wiki/English_Civil_War" title="English Civil War">English Civil War</a>, the <a href="/wiki/Royal_Society" title="Royal Society">Royal Society</a> wanted to show that science did not threaten religious and political stability. <a href="/wiki/John_Ray" title="John Ray">John Ray</a> developed an influential <a href="/wiki/Natural_theology" title="Natural theology">natural theology</a> of rational order; in his <a href="/wiki/Taxonomy_(biology)" title="Taxonomy (biology)">taxonomy</a>, <a href="/wiki/Species" title="Species">species</a> were static and fixed, their adaptation and complexity designed by God, and <a href="/wiki/Subspecies" title="Subspecies">varieties</a> showed minor differences caused by local conditions. In God's benevolent design, carnivores caused mercifully swift death, but the suffering caused by <a href="/wiki/Parasitism" title="Parasitism">parasitism</a> was a <a href="/wiki/Problem_of_evil" title="Problem of evil">puzzling problem</a>. The <a href="/wiki/Biological_classification" class="mw-redirect" title="Biological classification">biological classification</a> introduced by <a href="/wiki/Carl_Linnaeus" title="Carl Linnaeus">Carl Linnaeus</a> in 1735 also viewed species as fixed according to the divine plan, but did recognize the hierarchical nature of different taxa. In 1766, <a href="/wiki/Georges_Buffon" class="mw-redirect" title="Georges Buffon">Georges Buffon</a> suggested that some similar species, such as horses and asses, or lions, tigers, and leopards, might be varieties descended from a common ancestor. The <a href="/wiki/Ussher_chronology" title="Ussher chronology">Ussher chronology</a> of the 1650s had calculated creation at 4004 BC, but by the 1780s geologists assumed a much older world. <a href="/wiki/Abraham_Gottlob_Werner" title="Abraham Gottlob Werner">Wernerians</a> thought <a href="/wiki/Stratum" title="Stratum">strata</a> were <a href="/wiki/Neptunism" title="Neptunism">deposits from shrinking seas</a>, but <a href="/wiki/James_Hutton" title="James Hutton">James Hutton</a> proposed a self-maintaining infinite cycle, anticipating <a href="/wiki/Uniformitarianism_(science)" class="mw-redirect" title="Uniformitarianism (science)">uniformitarianism</a>.<a href="#cite_note-11">[11]</a> Charles Darwin's grandfather <a href="/wiki/Erasmus_Darwin" title="Erasmus Darwin">Erasmus Darwin</a> outlined a hypothesis of <a href="/wiki/Transmutation_of_species" title="Transmutation of species">transmutation of species</a> in the 1790s, and French naturalist <a href="/wiki/Jean-Baptiste_Lamarck" title="Jean-Baptiste Lamarck">Jean-Baptiste Lamarck</a> published a more developed theory in 1809. Both envisaged that spontaneous generation produced simple forms of life that progressively developed greater complexity, adapting to the environment by inheriting changes in adults caused by use or disuse. This process was later called <a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a>. Lamarck thought there was an inherent progressive tendency driving organisms continuously towards greater complexity, in parallel but separate lineages with no perceptible extinction.<a href="#cite_note-12">[12]</a> <a href="/wiki/%C3%89tienne_Geoffroy_Saint-Hilaire" title="Étienne Geoffroy Saint-Hilaire">Geoffroy</a> contended that <a href="/wiki/Recapitulation_theory" title="Recapitulation theory">embryonic development recapitulated</a> transformations of organisms in past <a href="/wiki/Era" title="Era">eras</a> when the environment acted on embryos, and that animal structures were determined by a constant plan as demonstrated by <a href="/wiki/Homology_(biology)" title="Homology (biology)">homologies</a>. <a href="/wiki/Georges_Cuvier" title="Georges Cuvier">Georges Cuvier</a> strongly disputed such ideas, holding that unrelated, fixed species showed similarities that reflected a design for functional needs.<a href="#cite_note-13">[13]</a> His <a href="/wiki/History_of_paleontology" title="History of paleontology">palæontological work</a> in the 1790s had established the reality of extinction, which he explained by local <a href="/wiki/Catastrophism" title="Catastrophism">catastrophes</a>, followed by repopulation of the affected areas by other species.<a href="#cite_note-14">[14]</a> In Britain, <a href="/wiki/William_Paley" title="William Paley">William Paley</a>'s Natural Theology saw adaptation as evidence of beneficial <a href="/wiki/Teleological_argument" title="Teleological argument">"design" by the Creator</a> acting through natural laws. All naturalists in the two English universities (<a href="/wiki/University_of_Oxford" title="University of Oxford">Oxford</a> and <a href="/wiki/University_of_Cambridge" title="University of Cambridge">Cambridge</a>) were <a href="/wiki/Church_of_England" title="Church of England">Church of England</a> clergymen, and science became a search for these laws.<a href="#cite_note-15">[15]</a> Geologists adapted catastrophism to show repeated worldwide annihilation and creation of new fixed species adapted to a changed environment, initially identifying the most recent catastrophe as the <a href="/wiki/Deluge_(mythology)" class="mw-redirect" title="Deluge (mythology)">biblical flood</a>.<a href="#cite_note-16">[16]</a> Some anatomists such as <a href="/wiki/Robert_Edmond_Grant" title="Robert Edmond Grant">Robert Grant</a> were influenced by Lamarck and Geoffroy, but most naturalists regarded their ideas of transmutation as a threat to divinely appointed social order.<a href="#cite_note-17">[17]</a> <div class="mw-heading mw-heading3"><h3 id="Inception_of_Darwin's_theory">Inception of Darwin's theory</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=4" title="Edit section: Inception of Darwin's theory">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Charles_Darwin%27s_education" title="Charles Darwin's education">Charles Darwin's education</a> and <a href="/wiki/Inception_of_Darwin%27s_theory" title="Inception of Darwin's theory">Inception of Darwin's theory</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Darwin_tree.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/58/Darwin_tree.png/170px-Darwin_tree.png" decoding="async" width="170" height="289" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/58/Darwin_tree.png/255px-Darwin_tree.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/58/Darwin_tree.png/340px-Darwin_tree.png 2x" data-file-width="695" data-file-height="1181" /></a><figcaption>In mid-July 1837 Darwin started his "B" notebook on Transmutation of Species, and on page 36 wrote "I think" above his first <a href="/wiki/Tree_of_life_(science)" class="mw-redirect" title="Tree of life (science)">evolutionary tree</a>.</figcaption></figure> Darwin went to <a href="/wiki/University_of_Edinburgh" title="University of Edinburgh">Edinburgh University</a> in 1825 to study medicine. In his second year he neglected his medical studies for <a href="/wiki/Natural_history" title="Natural history">natural history</a> and spent four months helping <a href="/wiki/Robert_Edmund_Grant" class="mw-redirect" title="Robert Edmund Grant">Robert Grant</a>'s research into <a href="/wiki/Marine_invertebrate" class="mw-redirect" title="Marine invertebrate">marine invertebrates</a>. Grant revealed his enthusiasm for the transmutation of species, but Darwin rejected it.<a href="#cite_note-18">[18]</a> Starting in 1827, at <a href="/wiki/University_of_Cambridge" title="University of Cambridge">Cambridge University</a>, Darwin learnt science as <a href="/wiki/Natural_theology" title="Natural theology">natural theology</a> from botanist <a href="/wiki/John_Stevens_Henslow" title="John Stevens Henslow">John Stevens Henslow</a>, and read <a href="/wiki/William_Paley" title="William Paley">Paley</a>, <a href="/wiki/John_Herschel" title="John Herschel">John Herschel</a> and <a href="/wiki/Alexander_von_Humboldt" title="Alexander von Humboldt">Alexander von Humboldt</a>. Filled with zeal for science, he studied <a href="/wiki/Catastrophism" title="Catastrophism">catastrophist</a> <a href="/wiki/Geology" title="Geology">geology</a> with <a href="/wiki/Adam_Sedgwick" title="Adam Sedgwick">Adam Sedgwick</a>.<a href="#cite_note-19">[19]</a><a href="#cite_note-20">[20]</a> In December 1831, he joined the <a href="/wiki/Second_voyage_of_HMS_Beagle" title="Second voyage of HMS Beagle">Beagle expedition</a> as a gentleman naturalist and geologist. He read <a href="/wiki/Charles_Lyell" title="Charles Lyell">Charles Lyell</a>'s <a href="/wiki/Principles_of_Geology" title="Principles of Geology">Principles of Geology</a> and from the first stop ashore, at <a href="/wiki/Santiago,_Cape_Verde" title="Santiago, Cape Verde">St. Jago</a>, found <a href="/wiki/Uniformitarianism_(science)" class="mw-redirect" title="Uniformitarianism (science)">Lyell's uniformitarianism</a> a key to the geological history of landscapes. Darwin discovered fossils resembling <a href="/wiki/Glyptodon" title="Glyptodon">huge armadillos</a>, and noted the geographical distribution of modern species in hope of finding their "centre of creation".<a href="#cite_note-21">[21]</a> The three <a href="/wiki/Fuegians" title="Fuegians">Fuegian</a> missionaries the expedition returned to <a href="/wiki/Tierra_del_Fuego" title="Tierra del Fuego">Tierra del Fuego</a> were friendly and civilised, yet to Darwin their relatives on the island seemed "miserable, degraded savages",<a href="#cite_note-22">[22]</a> and he no longer saw an unbridgeable gap between humans and animals.<a href="#cite_note-23">[23]</a> As the Beagle neared England in 1836, he noted that species might not be fixed.<a href="#cite_note-xix-24">[24]</a><a href="#cite_note-25">[25]</a> <a href="/wiki/Richard_Owen" title="Richard Owen">Richard Owen</a> showed that fossils of extinct species Darwin found in South America were allied to living species on the same continent. In March 1837, ornithologist <a href="/wiki/John_Gould" title="John Gould">John Gould</a> announced that <a href="/wiki/Darwin%27s_rhea" title="Darwin's rhea">Darwin's rhea</a> was a separate species from the previously described <a href="/wiki/Rhea_(bird)" title="Rhea (bird)">rhea</a> (though their territories overlapped), that <a href="/wiki/Mockingbird" title="Mockingbird">mockingbirds</a> collected on the <a href="/wiki/Gal%C3%A1pagos_Islands" title="Galápagos Islands">Galápagos Islands</a> represented three separate species each <a href="/wiki/Endemism" title="Endemism">unique to</a> a particular island, and that several distinct birds from those islands were all classified as <a href="/wiki/Darwin%27s_finch" class="mw-redirect" title="Darwin's finch">finches</a>.<a href="#cite_note-26">[26]</a> Darwin began speculating, in a series of notebooks, on the possibility that "one species does change into another" to explain these findings, and around July sketched a <a href="/wiki/Genealogy" title="Genealogy">genealogical</a> branching of a single <a href="/wiki/Tree_of_life_(science)" class="mw-redirect" title="Tree of life (science)">evolutionary tree</a>, discarding Lamarck's independent <a href="/wiki/Lineage_(evolution)" title="Lineage (evolution)">lineages</a> progressing to higher forms.<a href="#cite_note-27">[27]</a><a href="#cite_note-28">[28]</a><a href="#cite_note-29">[29]</a> Unconventionally, Darwin asked questions of <a href="/wiki/Fancy_pigeon" title="Fancy pigeon">fancy pigeon</a> and <a href="/wiki/Animal_husbandry" title="Animal husbandry">animal breeders</a> as well as established scientists. At the zoo he had his first sight of an ape, and was profoundly impressed by how human the <a href="/wiki/Orangutan" title="Orangutan">orangutan</a> seemed.<a href="#cite_note-30">[30]</a> In late September 1838, he started reading <a href="/wiki/Thomas_Malthus" class="mw-redirect" title="Thomas Malthus">Thomas Malthus</a>'s <a href="/wiki/An_Essay_on_the_Principle_of_Population" title="An Essay on the Principle of Population">An Essay on the Principle of Population</a> with <a href="/wiki/Malthusian_trap" class="mw-redirect" title="Malthusian trap">its statistical argument</a> that human populations, if unrestrained, breed beyond their means and <a href="/wiki/Struggle_for_existence" title="Struggle for existence">struggle to survive</a>. Darwin related this to the struggle for existence among wildlife and botanist <a href="/wiki/Augustin_Pyramus_de_Candolle" title="Augustin Pyramus de Candolle">de Candolle's</a> "warring of the species" in plants; he immediately envisioned "a force like a hundred thousand wedges" pushing well-adapted variations into "gaps in the economy of nature", so that the survivors would pass on their form and abilities, and unfavourable variations would be destroyed.<a href="#cite_note-wyhe-31">[31]</a><a href="#cite_note-32">[32]</a><a href="#cite_note-33">[33]</a> By December 1838, he had noted a similarity between the act of breeders selecting traits and a Malthusian Nature selecting among variants thrown up by "chance" so that "every part of newly acquired structure is fully practical and perfected".<a href="#cite_note-34">[34]</a> Darwin now had the basic framework of his theory of natural selection, but he was fully occupied with his career as a geologist and held back from compiling it until his book on <a href="/wiki/The_Structure_and_Distribution_of_Coral_Reefs" title="The Structure and Distribution of Coral Reefs">The Structure and Distribution of Coral Reefs</a> was completed.<a href="#cite_note-vw186-35">[35]</a><a href="#cite_note-36">[36]</a> As he recalled in his autobiography, he had "at last got a theory by which to work", but it was only in June 1842 that he allowed himself "the satisfaction of writing a very brief abstract of my theory in pencil".<a href="#cite_note-AB_120-37">[37]</a> <div class="mw-heading mw-heading3"><h3 id="Further_development">Further development</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=5" title="Edit section: Further development">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Development_of_Darwin%27s_theory" title="Development of Darwin's theory">Development of Darwin's theory</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Charles_Darwin_-_Pigeon_skulls.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c6/Charles_Darwin_-_Pigeon_skulls.png/220px-Charles_Darwin_-_Pigeon_skulls.png" decoding="async" width="220" height="196" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c6/Charles_Darwin_-_Pigeon_skulls.png/330px-Charles_Darwin_-_Pigeon_skulls.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c6/Charles_Darwin_-_Pigeon_skulls.png/440px-Charles_Darwin_-_Pigeon_skulls.png 2x" data-file-width="1800" data-file-height="1600" /></a><figcaption>Darwin researched how the skulls of different pigeon breeds varied, as shown in his Variation of Plants and Animals Under Domestication of 1868.</figcaption></figure> Darwin continued to research and extensively revise his theory while focusing on his main work of publishing the scientific results of the Beagle voyage.<a href="#cite_note-vw186-35">[35]</a> He tentatively wrote of his ideas to Lyell in January 1842;<a href="#cite_note-38">[38]</a> then in June he roughed out a 35-page "Pencil Sketch" of his theory.<a href="#cite_note-39">[39]</a> Darwin began correspondence about his theorising with the botanist <a href="/wiki/Joseph_Dalton_Hooker" title="Joseph Dalton Hooker">Joseph Dalton Hooker</a> in January 1844, and by July had rounded out his "sketch" into a 230-page "Essay", to be expanded with his research results and published if he died prematurely.<a href="#cite_note-40">[40]</a> In November 1844, the anonymously published <a href="/wiki/Popular_science" title="Popular science">popular science</a> book <a href="/wiki/Vestiges_of_the_Natural_History_of_Creation" title="Vestiges of the Natural History of Creation">Vestiges of the Natural History of Creation</a>, written by Scottish journalist <a href="/wiki/Robert_Chambers_(journalist)" class="mw-redirect" title="Robert Chambers (journalist)">Robert Chambers</a>, widened public interest in the concept of transmutation of species. Vestiges used evidence from the fossil record and embryology to support the claim that living things had progressed from the simple to the more complex over time. But it proposed a linear progression rather than the branching common descent theory behind Darwin's work in progress, and it ignored <a href="/wiki/Adaptation" title="Adaptation">adaptation</a>. Darwin read it soon after publication, and scorned its amateurish geology and zoology,<a href="#cite_note-41">[41]</a> but he carefully reviewed his own arguments after leading scientists, including Adam Sedgwick, attacked its morality and scientific errors.<a href="#cite_note-42">[42]</a> Vestiges had significant influence on public opinion, and the intense debate helped to pave the way for the acceptance of the more scientifically sophisticated Origin by moving evolutionary speculation into the mainstream. While few naturalists were willing to consider transmutation, <a href="/wiki/Herbert_Spencer" title="Herbert Spencer">Herbert Spencer</a> became an active proponent of Lamarckism and progressive development in the 1850s.<a href="#cite_note-43">[43]</a> Hooker was persuaded to take away a copy of the "Essay" in January 1847, and eventually sent a page of notes giving Darwin much-needed feedback. Reminded of his lack of expertise in <a href="/wiki/Taxonomy_(biology)" title="Taxonomy (biology)">taxonomy</a>, Darwin began an eight-year study of <a href="/wiki/Barnacle" title="Barnacle">barnacles</a>, becoming the leading expert on their classification. Using his theory, he discovered <a href="/wiki/Homology_(biology)" title="Homology (biology)">homologies</a> showing that slightly changed body parts served different functions to meet new conditions, and he found an <a href="/wiki/Androdioecy" title="Androdioecy">intermediate stage</a> in the evolution of <a href="/wiki/Gonochorism" title="Gonochorism">distinct sexes</a>.<a href="#cite_note-bowl169-44">[44]</a><a href="#cite_note-45">[45]</a> Darwin's barnacle studies convinced him that variation arose constantly and not just in response to changed circumstances. In 1854, he completed the last part of his Beagle-related writing and began working full-time on evolution. He now realised that the branching pattern of <a href="/wiki/Genetic_divergence" title="Genetic divergence">evolutionary divergence</a> was explained by natural selection working constantly to improve adaptation. His thinking changed from the view that <a href="/wiki/Allopatric_speciation" title="Allopatric speciation">species formed in isolated populations only</a>, as on islands, to an emphasis on <a href="/wiki/Sympatric_speciation" title="Sympatric speciation">speciation without isolation</a>; that is, he saw increasing specialisation within large stable populations as continuously exploiting new <a href="/wiki/Ecological_niche" title="Ecological niche">ecological niches</a>. He conducted empirical research focusing on difficulties with his theory. He studied the developmental and anatomical differences between different breeds of many domestic animals, became actively involved in <a href="/wiki/Fancy_pigeon" title="Fancy pigeon">fancy pigeon</a> breeding, and experimented (with the help of his young son <a href="/wiki/Francis_Darwin" title="Francis Darwin">Francis</a>) on ways that plant seeds and animals might disperse across oceans to colonise distant islands. By 1856, his theory was much more sophisticated, with a mass of supporting evidence.<a href="#cite_note-bowl169-44">[44]</a><a href="#cite_note-46">[46]</a> <div class="mw-heading mw-heading2"><h2 id="Publication">Publication</h2>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=6" title="Edit section: Publication">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Publication_of_Darwin%27s_theory" title="Publication of Darwin's theory">Publication of Darwin's theory</a></div> <div class="mw-heading mw-heading3"><h3 id="Time_taken_to_publish">Time taken to publish</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=7" title="Edit section: Time taken to publish">edit</a>]</div> In his autobiography, Darwin said he had "gained much by my delay in publishing from about 1839, when the theory was clearly conceived, to 1859; and I lost nothing by it".<a href="#cite_note-47">[47]</a> On the first page of his 1859 book he noted that, having begun work on the topic in 1837, he had drawn up "some short notes" after five years, had enlarged these into a sketch in 1844, and "from that period to the present day I have steadily pursued the same object."<a href="#cite_note-vW_time_to_publish-48">[48]</a><a href="#cite_note-FOOTNOTEDarwin1859[httpdarwin-onlineorgukcontentframesetpageseq16itemIDF373viewtypetext_1]-49">[49]</a> Various biographers have proposed that Darwin avoided or delayed making his ideas public for personal reasons. Reasons suggested have included fear of religious persecution or social disgrace if his views were revealed, and concern about upsetting his clergymen naturalist friends or his pious wife Emma. <a href="/wiki/Charles_Darwin%27s_illness" class="mw-redirect" title="Charles Darwin's illness">Charles Darwin's illness</a> caused repeated delays. His paper on <a href="/wiki/Glen_Roy" title="Glen Roy">Glen Roy</a> had proved embarrassingly wrong, and he may have wanted to be sure he was correct. <a href="/wiki/David_Quammen" title="David Quammen">David Quammen</a> has suggested all these factors may have contributed, and notes Darwin's large output of books and busy family life during that time.<a href="#cite_note-50">[50]</a> A more recent study by science historian <a href="/wiki/John_van_Wyhe" title="John van Wyhe">John van Wyhe</a> has determined that the idea that Darwin delayed publication only dates back to the 1940s, and Darwin's contemporaries thought the time he took was reasonable. Darwin always finished one book before starting another. While he was researching, he told many people about his interest in transmutation without causing outrage. He firmly intended to publish, but it was not until September 1854 that he could work on it full-time. His 1846 estimate that writing his "big book" would take five years proved optimistic.<a href="#cite_note-vW_time_to_publish-48">[48]</a> <div class="mw-heading mw-heading3"><h3 id="Events_leading_to_publication:_"big_book"_manuscript">Events leading to publication: "big book" manuscript</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=8" title="Edit section: Events leading to publication: "big book" manuscript">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Alfred_Russel_Wallace_1862_-_Project_Gutenberg_eText_15997.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b2/Alfred_Russel_Wallace_1862_-_Project_Gutenberg_eText_15997.png/170px-Alfred_Russel_Wallace_1862_-_Project_Gutenberg_eText_15997.png" decoding="async" width="170" height="324" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b2/Alfred_Russel_Wallace_1862_-_Project_Gutenberg_eText_15997.png/255px-Alfred_Russel_Wallace_1862_-_Project_Gutenberg_eText_15997.png 1.5x, //upload.wikimedia.org/wikipedia/commons/b/b2/Alfred_Russel_Wallace_1862_-_Project_Gutenberg_eText_15997.png 2x" data-file-width="304" data-file-height="580" /></a><figcaption>A photograph of <a href="/wiki/Alfred_Russel_Wallace" title="Alfred Russel Wallace">Alfred Russel Wallace</a> (1823–1913) taken in <a href="/wiki/Singapore" title="Singapore">Singapore</a> in 1862</figcaption></figure> An 1855 paper on the "introduction" of species, written by <a href="/wiki/Alfred_Russel_Wallace" title="Alfred Russel Wallace">Alfred Russel Wallace</a>, claimed that patterns in the geographical distribution of living and fossil species could be explained if every new species always came into existence near an already existing, closely related species.<a href="#cite_note-51">[51]</a> <a href="/wiki/Charles_Lyell" title="Charles Lyell">Charles Lyell</a> recognised the implications of Wallace's paper and its possible connection to Darwin's work, although Darwin did not, and in a letter written on 1–2 May 1856 Lyell urged Darwin to publish his theory to establish priority. Darwin was torn between the desire to set out a full and convincing account and the pressure to quickly produce a short paper. He met Lyell, and in correspondence with <a href="/wiki/Joseph_Dalton_Hooker" title="Joseph Dalton Hooker">Joseph Dalton Hooker</a> affirmed that he did not want to expose his ideas to review by an editor as would have been required to publish in an academic journal. He began a "sketch" account on 14 May 1856, and by July had decided to produce a full technical treatise on species as his "big book" on <a href="/wiki/Natural_Selection_(manuscript)" title="Natural Selection (manuscript)">Natural Selection</a>. His theory including the principle of <a href="/wiki/Genetic_divergence" title="Genetic divergence">divergence</a> was complete by 5 September 1857 when he sent <a href="/wiki/Asa_Gray" title="Asa Gray">Asa Gray</a> a brief but detailed abstract of his ideas.<a href="#cite_note-Quammen135-158-52">[52]</a><a href="#cite_note-letters:_Big_Book-53">[53]</a> <div class="mw-heading mw-heading3"><h3 id="Joint_publication_of_papers_by_Wallace_and_Darwin">Joint publication of papers by Wallace and Darwin</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=9" title="Edit section: Joint publication of papers by Wallace and Darwin">edit</a>]</div> Darwin was hard at work on the manuscript for his "big book" on <a href="/wiki/Natural_Selection_(manuscript)" title="Natural Selection (manuscript)">Natural Selection</a>, when on 18 June 1858 he received a parcel from Wallace, who stayed on the <a href="/wiki/Maluku_Islands" title="Maluku Islands">Maluku Islands</a> (<a href="/wiki/Ternate" title="Ternate">Ternate</a> and Gilolo). It enclosed twenty pages describing an evolutionary mechanism, a response to Darwin's recent encouragement, with a request to send it on to Lyell if Darwin thought it worthwhile. The mechanism was similar to Darwin's own theory.<a href="#cite_note-Quammen135-158-52">[52]</a> Darwin wrote to Lyell that "your words have come true with a vengeance, ... forestalled" and he would "of course, at once write and offer to send [it] to any journal" that Wallace chose, adding that "all my originality, whatever it may amount to, will be smashed".<a href="#cite_note-54">[54]</a> Lyell and Hooker agreed that a joint publication putting together Wallace's pages with extracts from Darwin's 1844 Essay and his 1857 letter to Gray should be presented at the <a href="/wiki/Linnean_Society_of_London" title="Linnean Society of London">Linnean Society</a>, and on 1 July 1858, the papers entitled <a href="/wiki/On_the_Tendency_of_Species_to_form_Varieties;_and_on_the_Perpetuation_of_Varieties_and_Species_by_Natural_Means_of_Selection" title="On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection">On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection</a>, by Wallace and Darwin respectively, were read out but drew little reaction. While Darwin considered Wallace's idea to be identical to his concept of natural selection, historians have pointed out differences. Darwin described natural selection as being analogous to the <a href="/wiki/Artificial_selection" class="mw-redirect" title="Artificial selection">artificial selection</a> practised by animal breeders, and emphasised competition between individuals; Wallace drew no comparison to <a href="/wiki/Selective_breeding" title="Selective breeding">selective breeding</a>, and focused on ecological pressures that kept different varieties adapted to local conditions.<a href="#cite_note-55">[55]</a><a href="#cite_note-56">[56]</a><a href="#cite_note-57">[57]</a> Some historians have suggested that Wallace was actually discussing <a href="/wiki/Group_selection" title="Group selection">group selection</a> rather than selection acting on individual variation.<a href="#cite_note-58">[58]</a> <div class="mw-heading mw-heading3"><h3 id="Abstract_of_Species_book">Abstract of Species book</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=10" title="Edit section: Abstract of Species book">edit</a>]</div> Soon after the meeting, Darwin decided to write "an abstract of my whole work" in the form of one or more papers to be published by the <a href="/wiki/Linnean_Society_of_London" title="Linnean Society of London">Linnean Society</a>, but was concerned about "how it can be made scientific for a Journal, without giving facts, which would be impossible." He asked Hooker how many pages would be available, but "If the Referees were to reject it as not strictly scientific I would, perhaps publish it as pamphlet."<a href="#cite_note-Correspondence_1858-1859:_Origin-59">[59]</a><a href="#cite_note-Letter_2303-60">[60]</a> He began his "abstract of Species book" on 20 July 1858, while on holiday at <a href="/wiki/Sandown" title="Sandown">Sandown</a>,<a href="#cite_note-Journal_36v-61">[61]</a> and wrote parts of it from memory, while sending the manuscripts to his friends for checking.<a href="#cite_note-Letter_2432-62">[62]</a> By early October, he began to "expect my abstract will run into a small volume, which will have to be published separately."<a href="#cite_note-Letter_2339-63">[63]</a> Over the same period, he continued to collect information and write large fully detailed sections of the manuscript for his "big book" on Species, <a href="/wiki/Natural_Selection_(manuscript)" title="Natural Selection (manuscript)">Natural Selection</a>.<a href="#cite_note-Correspondence_1858-1859:_Origin-59">[59]</a> <div class="mw-heading mw-heading3"><h3 id="Murray_as_publisher;_choice_of_title">Murray as publisher; choice of title</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=11" title="Edit section: Murray as publisher; choice of title">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:On_the_origin_of_species_by_means_of_natural_selection_OR_the_preservation_of_favoured_races_in_the_struggle_for_life,_2nd_edition._By_Chalres_Darwin,_John_Murray,_London,_1860._National_Museum_of_Scotland.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fb/On_the_origin_of_species_by_means_of_natural_selection_OR_the_preservation_of_favoured_races_in_the_struggle_for_life%2C_2nd_edition._By_Chalres_Darwin%2C_John_Murray%2C_London%2C_1860._National_Museum_of_Scotland.jpg/290px-thumbnail.jpg" decoding="async" width="290" height="247" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fb/On_the_origin_of_species_by_means_of_natural_selection_OR_the_preservation_of_favoured_races_in_the_struggle_for_life%2C_2nd_edition._By_Chalres_Darwin%2C_John_Murray%2C_London%2C_1860._National_Museum_of_Scotland.jpg/435px-thumbnail.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fb/On_the_origin_of_species_by_means_of_natural_selection_OR_the_preservation_of_favoured_races_in_the_struggle_for_life%2C_2nd_edition._By_Chalres_Darwin%2C_John_Murray%2C_London%2C_1860._National_Museum_of_Scotland.jpg/580px-thumbnail.jpg 2x" data-file-width="2571" data-file-height="2191" /></a><figcaption>On the Origin of Species by means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life, 2nd edition. By Charles Darwin, John Murray, London, 1860. National Museum of Scotland.</figcaption></figure> By mid-March 1859 Darwin's abstract had reached the stage where he was thinking of early publication; Lyell suggested the publisher <a href="/wiki/John_Murray_III" title="John Murray III">John Murray</a>, and met with him to find if he would be willing to publish. On 28 March Darwin wrote to Lyell asking about progress, and offering to give Murray assurances "that my Book is not more un-orthodox, than the subject makes inevitable." He enclosed a draft title sheet proposing An abstract of an Essay on the Origin of Species and Varieties Through natural selection, with the year shown as "1859".<a href="#cite_note-Letter_2437-64">[64]</a><a href="#cite_note-FOOTNOTEDesmondMoore2009306-65">[65]</a> Murray's response was favourable, and a very pleased Darwin told Lyell on 30 March that he would "send shortly a large bundle of M.S. but unfortunately I cannot for a week, as the three first chapters are in three copyists' hands". He bowed to Murray's objection to "abstract" in the title, though he felt it excused the lack of references, but wanted to keep "natural selection" which was "constantly used in all works on Breeding", and hoped "to retain it with Explanation, somewhat as thus",— Through Natural Selection or the preservation of favoured races.<a href="#cite_note-FOOTNOTEDesmondMoore2009306-65">[65]</a><a href="#cite_note-Letter_2439-66">[66]</a> On 31 March Darwin wrote to Murray in confirmation, and listed headings of the 12 chapters in progress: he had drafted all except "XII. Recapitulation & Conclusion".<a href="#cite_note-Letter_2441-67">[67]</a> Murray responded immediately with an agreement to publish the book on the same terms as he published Lyell, without even seeing the manuscript: he offered Darwin ⅔ of the profits.<a href="#cite_note-Letter_2443-68">[68]</a> Darwin promptly accepted with pleasure, insisting that Murray would be free to withdraw the offer if, having read the chapter manuscripts, he felt the book would not sell well<a href="#cite_note-Letter_2445-69">[69]</a> (eventually Murray paid £180 to Darwin for the first edition and by Darwin's death in 1882 the book was in its sixth edition, earning Darwin nearly £3000<a href="#cite_note-Darwin_and_his_publisher-70">[70]</a>). On 5 April, Darwin sent Murray the first three chapters, and a proposal for the book's title.<a href="#cite_note-Letter_2447-71">[71]</a> An early draft title page suggests On the Mutability of Species.<a href="#cite_note-72">[72]</a> Murray cautiously asked <a href="/wiki/Whitwell_Elwin" title="Whitwell Elwin">Whitwell Elwin</a> to review the chapters.<a href="#cite_note-Correspondence_1858-1859:_Origin-59">[59]</a> At Lyell's suggestion, Elwin recommended that, rather than "put forth the theory without the evidence", the book should focus on observations upon <a href="/wiki/Pigeon" class="mw-redirect" title="Pigeon">pigeons</a>, briefly stating how these illustrated Darwin's general principles and preparing the way for the larger work expected shortly: "Every body is interested in pigeons."<a href="#cite_note-Letter_2457A-73">[73]</a> Darwin responded that this was impractical: he had only the last chapter still to write.<a href="#cite_note-Letter_2459-74">[74]</a> In September the main title still included "An essay on the origin of species and varieties", but Darwin now proposed dropping "varieties".<a href="#cite_note-Letter_2448-75">[75]</a> With Murray's persuasion, the title was eventually agreed as On the Origin of Species, with the title page adding by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life.<a href="#cite_note-titles-3">[3]</a> In this extended title (and elsewhere in the book) Darwin used the biological term "<a href="/wiki/Race_(biology)" title="Race (biology)">races</a>" interchangeably with "<a href="/wiki/Variety_(botany)" title="Variety (botany)">varieties</a>", meaning varieties within a <a href="/wiki/Species" title="Species">species</a>.<a href="#cite_note-NCSE_define_evo-76">[76]</a><a href="#cite_note-BernasconiLott2000-77">[77]</a> He used the term broadly,<a href="#cite_note-Sober45-78">[78]</a> and as well as discussions of "the several races, for instance, of the cabbage" and "the hereditary varieties or races of our domestic animals and plants",<a href="#cite_note-79">[79]</a> there are three instances in the book where the phrase "races of man" is used, referring to <a href="/wiki/Race_(human_classification)" class="mw-redirect" title="Race (human classification)">races</a> of humans.<a href="#cite_note-80">[80]</a> <div class="mw-heading mw-heading3"><h3 id="Publication_and_subsequent_editions">Publication and subsequent editions</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=12" title="Edit section: Publication and subsequent editions">edit</a>]</div> On the Origin of Species was first published on Thursday 24 November 1859, priced at fifteen <a href="/wiki/Shilling" title="Shilling">shillings</a> with a first printing of 1250 copies.<a href="#cite_note-dupree-81">[81]</a> The book had been offered to booksellers at Murray's autumn sale on Tuesday 22 November, and all available copies had been taken up immediately. In total, 1,250 copies were printed but after deducting presentation and review copies, and five for <a href="/wiki/Stationers%27_Hall" class="mw-redirect" title="Stationers' Hall">Stationers' Hall</a> copyright, around 1,170 copies were available for sale.<a href="#cite_note-Freeman_1977-2">[2]</a> Significantly, 500 were taken by <a href="/wiki/Charles_Edward_Mudie" title="Charles Edward Mudie">Mudie's Library</a>, ensuring that the book promptly reached a large number of subscribers to the library.<a href="#cite_note-82">[82]</a> The second edition of 3,000 copies was quickly brought out on 7 January 1860,<a href="#cite_note-83">[83]</a> and incorporated numerous corrections as well as a response to religious objections by the addition of a new epigraph on page ii, a quotation from <a href="/wiki/Charles_Kingsley" title="Charles Kingsley">Charles Kingsley</a>, and the phrase "by the Creator" added to the closing sentence.<a href="#cite_note-b95-84">[84]</a> During Darwin's lifetime the book went through six editions, with cumulative changes and revisions to deal with counter-arguments raised. The third edition came out in 1861, with a number of sentences rewritten or added and an introductory appendix, An Historical Sketch of the Recent Progress of Opinion on the Origin of Species.<a href="#cite_note-85">[85]</a> In response to objections that the <a href="/wiki/Abiogenesis" title="Abiogenesis">origin of life</a> was unexplained, Darwin pointed to acceptance of <a href="/wiki/Newton%27s_law_of_universal_gravitation" title="Newton's law of universal gravitation">Newton's law</a> even though the cause of gravity was unknown, and <a href="/wiki/Gottfried_Wilhelm_Leibniz" title="Gottfried Wilhelm Leibniz">Leibnitz</a> had accused Newton of introducing "occult qualities & miracles".<a href="#cite_note-old_Darwin_manuscript_2022-86">[86]</a><a href="#cite_note-87">[87]</a> The fourth edition in 1866 had further revisions. The fifth edition, published on 10 February 1869, incorporated more changes and for the first time included the phrase "<a href="/wiki/Survival_of_the_fittest" title="Survival of the fittest">survival of the fittest</a>", which had been coined by the philosopher <a href="/wiki/Herbert_Spencer" title="Herbert Spencer">Herbert Spencer</a> in his Principles of Biology (1864).<a href="#cite_note-sotf-88">[88]</a> In January 1871, <a href="/wiki/George_Jackson_Mivart" class="mw-redirect" title="George Jackson Mivart">George Jackson Mivart</a>'s On the Genesis of Species listed detailed arguments against natural selection, and claimed it included false <a href="/wiki/Metaphysics" title="Metaphysics">metaphysics</a>.<a href="#cite_note-miv-89">[89]</a> Darwin made extensive revisions to the sixth edition of the Origin (this was the first edition in which he used the word "<a href="/wiki/Evolution" title="Evolution">evolution</a>" which had commonly been associated with <a href="/wiki/Prenatal_development_(biology)" class="mw-redirect" title="Prenatal development (biology)">embryological development</a>, though all editions concluded with the word "evolved"<a href="#cite_note-90">[90]</a><a href="#cite_note-91">[91]</a>), and added a new chapter VII, Miscellaneous objections, to address Mivart's arguments.<a href="#cite_note-Freeman_1977-2">[2]</a><a href="#cite_note-6th-92">[92]</a> The sixth edition was published by Murray on 19 February 1872 as The Origin of Species, with "On" dropped from the title. Darwin had told Murray of working men in <a href="/wiki/Lancashire" title="Lancashire">Lancashire</a> clubbing together to buy the fifth edition at 15 shillings and wanted it made more widely available; the price was halved to 7<a href="/wiki/Shilling" title="Shilling">s</a> 6<a href="/wiki/Old_pence" class="mw-redirect" title="Old pence">d</a> by printing in a smaller <a href="/wiki/Font" title="Font">font</a>. It includes a glossary compiled by W.S. Dallas. Book sales increased from 60 to 250 per month.<a href="#cite_note-titles-3">[3]</a><a href="#cite_note-6th-92">[92]</a> <div class="mw-heading mw-heading3"><h3 id="Publication_outside_Great_Britain">Publication outside Great Britain</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=13" title="Edit section: Publication outside Great Britain">edit</a>]</div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Asa_Gray_(1867).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/40/Asa_Gray_%281867%29.jpg/170px-Asa_Gray_%281867%29.jpg" decoding="async" width="170" height="185" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/40/Asa_Gray_%281867%29.jpg/255px-Asa_Gray_%281867%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/40/Asa_Gray_%281867%29.jpg/340px-Asa_Gray_%281867%29.jpg 2x" data-file-width="368" data-file-height="400" /></a><figcaption>American botanist Asa Gray (1810–1888)</figcaption></figure> In the United States, botanist <a href="/wiki/Asa_Gray" title="Asa Gray">Asa Gray</a>, an American colleague of Darwin, negotiated with a Boston publisher for publication of an authorised American version, but learnt that two New York publishing firms were already planning to exploit the absence of international <a href="/wiki/Copyright" title="Copyright">copyright</a> to print Origin.<a href="#cite_note-Letter2592-93">[93]</a> Darwin was delighted by the popularity of the book, and asked Gray to keep any profits.<a href="#cite_note-Letter2665-94">[94]</a> Gray managed to negotiate a 5% royalty with <a href="/wiki/D._Appleton_%26_Company" title="D. Appleton & Company">Appleton's</a> of New York,<a href="#cite_note-Letter2706-95">[95]</a> who got their edition out in mid-January 1860, and the other two withdrew. In a May letter, Darwin mentioned a print run of 2,500 copies, but it is not clear if this referred to the first printing only, as there were four that year.<a href="#cite_note-Freeman_1977-2">[2]</a><a href="#cite_note-96">[96]</a> The book was widely translated in Darwin's lifetime, but problems arose with translating concepts and metaphors, and some translations were biased by the translator's own agenda.<a href="#cite_note-browne256-97">[97]</a> Darwin distributed presentation copies in France and Germany, hoping that suitable applicants would come forward, as translators were expected to make their own arrangements with a local publisher. He welcomed the distinguished elderly naturalist and geologist <a href="/wiki/Heinrich_Georg_Bronn" title="Heinrich Georg Bronn">Heinrich Georg Bronn</a>, but the German translation published in 1860 imposed Bronn's own ideas, adding controversial themes that Darwin had deliberately omitted. Bronn translated "favoured races" as "perfected races", and added essays on issues including the origin of life, as well as a final chapter on religious implications partly inspired by Bronn's adherence to <a href="/wiki/Naturphilosophie" title="Naturphilosophie">Naturphilosophie</a>.<a href="#cite_note-browne140-98">[98]</a> In 1862, Bronn produced a second edition based on the third English edition and Darwin's suggested additions, but then died of a heart attack.<a href="#cite_note-correspondence1862-99">[99]</a> Darwin corresponded closely with <a href="/wiki/Julius_Victor_Carus" title="Julius Victor Carus">Julius Victor Carus</a>, who published an improved translation in 1867.<a href="#cite_note-correspondence1866-100">[100]</a> Darwin's attempts to find a translator in France fell through, and the translation by <a href="/wiki/Cl%C3%A9mence_Royer" title="Clémence Royer">Clémence Royer</a> published in 1862 added an introduction praising Darwin's ideas as an alternative to religious revelation and promoting ideas anticipating <a href="/wiki/Social_Darwinism" title="Social Darwinism">social Darwinism</a> and <a href="/wiki/Eugenics" title="Eugenics">eugenics</a>, as well as numerous explanatory notes giving her own answers to doubts that Darwin expressed. Darwin corresponded with Royer about a second edition published in 1866 and a third in 1870, but he had difficulty getting her to remove her notes and was troubled by these editions.<a href="#cite_note-correspondence1862-99">[99]</a><a href="#cite_note-browne142-101">[101]</a> He remained unsatisfied until a translation by Edmond Barbier was published in 1876.<a href="#cite_note-Freeman_1977-2">[2]</a> A Dutch translation by <a href="/wiki/Tiberius_Cornelis_Winkler" title="Tiberius Cornelis Winkler">Tiberius Cornelis Winkler</a> was published in 1860.<a href="#cite_note-102">[102]</a> By 1864, additional translations had appeared in Italian and Russian.<a href="#cite_note-browne256-97">[97]</a> In Darwin's lifetime, Origin was published in Swedish in 1871,<a href="#cite_note-103">[103]</a> Danish in 1872, Polish in 1873, Hungarian in 1873–1874, Spanish in 1877 and Serbian in 1878. As of 1977, Origin had appeared in an additional 18 languages,<a href="#cite_note-free83-104">[104]</a> including Chinese by <a href="/wiki/Ma_Junwu" title="Ma Junwu">Ma Chün-wu</a> who added non-Darwinian ideas; he published the preliminaries and chapters 1–5 in 1902–1904, and his complete translation in 1920.<a href="#cite_note-free101-105">[105]</a><a href="#cite_note-Jin_2018-106">[106]</a> <div class="mw-heading mw-heading2"><h2 id="Content">Content</h2>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=14" title="Edit section: Content">edit</a>]</div> <div class="mw-heading mw-heading3"><h3 id="Title_pages_and_introduction">Title pages and introduction</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=15" title="Edit section: Title pages and introduction">edit</a>]</div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Image-Rhea_Darwinii1.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/92/Image-Rhea_Darwinii1.jpg/220px-Image-Rhea_Darwinii1.jpg" decoding="async" width="220" height="281" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/92/Image-Rhea_Darwinii1.jpg/330px-Image-Rhea_Darwinii1.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/92/Image-Rhea_Darwinii1.jpg/440px-Image-Rhea_Darwinii1.jpg 2x" data-file-width="601" data-file-height="768" /></a><figcaption><a href="/wiki/John_Gould" title="John Gould">John Gould</a>'s illustration of <a href="/wiki/Darwin%27s_rhea" title="Darwin's rhea">Darwin's rhea</a> was published in 1841. The existence of two rhea species with overlapping ranges influenced Darwin.</figcaption></figure> Page ii contains quotations by <a href="/wiki/William_Whewell" title="William Whewell">William Whewell</a> and <a href="/wiki/Francis_Bacon" title="Francis Bacon">Francis Bacon</a> on the theology of <a href="/wiki/Physical_law" class="mw-redirect" title="Physical law">natural laws</a>,<a href="#cite_note-107">[107]</a> harmonising science and religion in accordance with <a href="/wiki/Isaac_Newton" title="Isaac Newton">Isaac Newton</a>'s belief in a rational God who established a law-abiding cosmos.<a href="#cite_note-scicreate-108">[108]</a> In the second edition, Darwin added an epigraph from <a href="/wiki/Joseph_Butler" title="Joseph Butler">Joseph Butler</a> affirming that God could work through scientific laws as much as through <a href="/wiki/Miracles" class="mw-redirect" title="Miracles">miracles</a>, in a nod to the religious concerns of his oldest friends.<a href="#cite_note-b95-84">[84]</a> The Introduction establishes Darwin's credentials as a naturalist and author,<a href="#cite_note-109">[109]</a> then refers to <a href="/wiki/John_Herschel" title="John Herschel">John Herschel</a>'s letter suggesting that the origin of species "would be found to be a natural in contradistinction to a miraculous process":<a href="#cite_note-110">[110]</a> <blockquote>WHEN on board HMS Beagle, as naturalist, I was much struck with certain facts in the distribution of the inhabitants of South America, and in the geological relations of the present to the past inhabitants of that continent. These facts seemed to me to throw some light on the origin of species—that mystery of mysteries, as it has been called by one of our greatest philosophers.<a href="#cite_note-111">[111]</a></blockquote> Darwin refers specifically to the distribution of the species <a href="/wiki/Rheas" class="mw-redirect" title="Rheas">rheas</a>, and to that of the <a href="/wiki/Gal%C3%A1pagos_tortoise" title="Galápagos tortoise">Galápagos tortoises</a> and <a href="/wiki/Mockingbird" title="Mockingbird">mockingbirds</a>. He mentions his years of work on his theory, and the arrival of Wallace at the same conclusion, which led him to "publish this Abstract" of his incomplete work. He outlines his ideas, and sets out the essence of his theory: <blockquote>As many more individuals of each species are born than can possibly survive; and as, consequently, there is a frequently recurring struggle for existence, it follows that any being, if it vary however slightly in any manner profitable to itself, under the complex and sometimes varying conditions of life, will have a better chance of surviving, and thus be naturally selected. From the strong principle of inheritance, any selected variety will tend to propagate its new and modified form.<a href="#cite_note-112">[112]</a></blockquote> Starting with the third edition, Darwin prefaced the introduction with a sketch of the historical development of evolutionary ideas.<a href="#cite_note-113">[113]</a> In that sketch he acknowledged that <a href="/wiki/Patrick_Matthew" title="Patrick Matthew">Patrick Matthew</a> had, unknown to Wallace or himself, anticipated the concept of natural selection in an appendix to a book published in 1831;<a href="#cite_note-114">[114]</a> in the fourth edition he mentioned that <a href="/wiki/William_Charles_Wells" title="William Charles Wells">William Charles Wells</a> had done so as early as 1813.<a href="#cite_note-115">[115]</a> <div class="mw-heading mw-heading3"><h3 id="Variation_under_domestication_and_under_nature">Variation under domestication and under nature</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=16" title="Edit section: Variation under domestication and under nature">edit</a>]</div> Chapter I covers <a href="/wiki/Animal_husbandry" title="Animal husbandry">animal husbandry</a> and <a href="/wiki/Plant_breeding" title="Plant breeding">plant breeding</a>, going back to <a href="/wiki/Ancient_Egypt" title="Ancient Egypt">ancient Egypt</a>. Darwin discusses contemporary opinions on the origins of different breeds under cultivation to argue that many have been produced from common ancestors by <a href="/wiki/Selective_breeding" title="Selective breeding">selective breeding</a>.<a href="#cite_note-116">[116]</a> As an illustration of <a href="/wiki/Artificial_selection" class="mw-redirect" title="Artificial selection">artificial selection</a>, he describes <a href="/wiki/Fancy_pigeon" title="Fancy pigeon">fancy pigeon</a> breeding,<a href="#cite_note-q184-117">[117]</a> noting that "[t]he diversity of the breeds is something astonishing", yet all were descended from one species, the <a href="/wiki/Rock_dove" title="Rock dove">rock dove</a>.<a href="#cite_note-118">[118]</a> Darwin saw two distinct kinds of variation: (1) rare abrupt changes he called "sports" or "monstrosities" (example: <a href="/wiki/Ancon_sheep" title="Ancon sheep">Ancon sheep</a> with short legs), and (2) ubiquitous small differences (example: slightly shorter or longer bill of pigeons).<a href="#cite_note-rez-119">[119]</a> Both types of hereditary changes can be used by breeders. However, for Darwin the small changes were most important in evolution. In this chapter Darwin expresses his erroneous belief that environmental change is necessary to generate variation.<a href="#cite_note-120">[120]</a> The opening two sentences of On The Origin demonstrate this point and also show that Darwin did not see the role of <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a> in <a href="/wiki/Stabilizing_selection" title="Stabilizing selection">stabilizing selection</a>: When we look to the individuals of the same variety or sub-variety of our older cultivated plants and animals, one of the first points which strikes us, is, that they generally differ much more from each other, than do the individuals of any one species or variety in a state of nature. When we reflect on the vast diversity of the plants and animals which have been cultivated, and which have varied during all ages under the most different climates and treatment, I think we are driven to conclude that this greater variability is simply due to our domestic productions having been raised under conditions of life not so uniform as, and somewhat different from, those to which the parent-species have been exposed under nature. It can be seen here that Darwin attributes the greater variation amongst individuals of domestic varieties compared to their progenitor populations in nature as being due to their "conditions of life" (environment) being "not so uniform as", and "somewhat different from" those of the parent species". He later erroneously elaborates that changed "conditions of life" act on the reproductive organs to generate greater variability in the progeny. Even after 1860 when Darwin read the correct reason for greater variation in domestic varieties in <a href="/wiki/Patrick_Matthew" title="Patrick Matthew">Patrick Matthew</a>'s book, <a href="/wiki/On_Naval_Timber_and_Arboriculture" title="On Naval Timber and Arboriculture">On Naval Timber and Arboriculture</a> published in 1831, future editions of On the Origin of Species contained these first two sentences unchanged and continued to omit the correct explanation of <a href="/wiki/Stabilizing_selection" title="Stabilizing selection">stabilizing selection</a>. In Chapter II, Darwin specifies that the distinction between <a href="/wiki/Species" title="Species">species</a> and <a href="/wiki/Variety_(biology)" class="mw-redirect" title="Variety (biology)">varieties</a> is arbitrary, with experts disagreeing and changing their decisions when new forms were found. He concludes that "a well-marked variety may be justly called an incipient species" and that "species are only strongly marked and permanent varieties".<a href="#cite_note-121">[121]</a> He argues for the ubiquity of variation in nature.<a href="#cite_note-122">[122]</a> Historians have noted that naturalists had long been aware that the individuals of a species differed from one another, but had generally considered such variations to be limited and unimportant deviations from the <a href="/wiki/Archetype" title="Archetype">archetype</a> of each species, that archetype being a fixed ideal in the mind of God. Darwin and Wallace made variation among individuals of the same species central to understanding the natural world.<a href="#cite_note-q184-117">[117]</a> <div class="mw-heading mw-heading3"><h3 id="Struggle_for_existence,_natural_selection,_and_divergence">Struggle for existence, natural selection, and divergence</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=17" title="Edit section: Struggle for existence, natural selection, and divergence">edit</a>]</div> In Chapter III, Darwin asks how varieties "which I have called incipient species" become distinct species, and in answer introduces the key concept he calls "<a href="/wiki/Natural_selection" title="Natural selection">natural selection</a>";<a href="#cite_note-dar5960-123">[123]</a> in the fifth edition he adds, "But the expression often used by Mr. <a href="/wiki/Herbert_Spencer" title="Herbert Spencer">Herbert Spencer</a>, of the <a href="/wiki/Survival_of_the_Fittest" class="mw-redirect" title="Survival of the Fittest">Survival of the Fittest</a>, is more accurate, and is sometimes equally convenient."<a href="#cite_note-124">[124]</a> <blockquote>Owing to this struggle for life, any variation, however slight and from whatever cause proceeding, if it be in any degree profitable to an individual of any species, in its infinitely complex relations to other organic beings and to external nature, will tend to the preservation of that individual, and will generally be inherited by its offspring ... I have called this principle, by which each slight variation, if useful, is preserved, by the term of Natural Selection, in order to mark its relation to man's power of selection.<a href="#cite_note-dar5960-123">[123]</a></blockquote> He notes that both <a href="/wiki/Augustin_Pyramus_de_Candolle" title="Augustin Pyramus de Candolle">A. P. de Candolle</a> and <a href="/wiki/Charles_Lyell" title="Charles Lyell">Charles Lyell</a> had stated that all organisms are exposed to severe competition. Darwin emphasizes that he used the phrase "<a href="/wiki/The_struggle_for_existence" class="mw-redirect" title="The struggle for existence">struggle for existence</a>" in "a large and metaphorical sense, including dependence of one being on another"; he gives examples ranging from plants struggling against drought to plants competing for birds to eat their fruit and disseminate their seeds. He describes the struggle resulting from population growth: "It is the doctrine of Malthus applied with manifold force to the whole animal and vegetable kingdoms." He discusses checks to such increase including complex <a href="/wiki/Ecology" title="Ecology">ecological</a> interdependencies, and notes that competition is most severe between closely related forms "which fill nearly the same place in the economy of nature".<a href="#cite_note-125">[125]</a> Chapter IV details natural selection under the "infinitely complex and close-fitting ... mutual relations of all organic beings to each other and to their physical conditions of life".<a href="#cite_note-126">[126]</a> Darwin takes as an example a country where a change in conditions led to extinction of some species, immigration of others and, where suitable variations occurred, descendants of some species became adapted to new conditions. He remarks that the artificial selection practised by animal breeders frequently produced sharp divergence in character between breeds, and suggests that natural selection might do the same, saying: <blockquote>But how, it may be asked, can any analogous principle apply in nature? I believe it can and does apply most efficiently, from the simple circumstance that the more diversified the descendants from any one species become in structure, constitution, and habits, by so much will they be better enabled to seize on many and widely diversified places in the polity of nature, and so be enabled to increase in numbers.<a href="#cite_note-127">[127]</a></blockquote> Historians have remarked that here Darwin anticipated the modern concept of an <a href="/wiki/Ecological_niche" title="Ecological niche">ecological niche</a>.<a href="#cite_note-128">[128]</a> He did not suggest that every favourable variation must be selected, nor that the favoured animals were better or higher, but merely more adapted to their surroundings. <figure typeof="mw:File/Thumb"><a href="/wiki/File:Darwin_divergence.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/45/Darwin_divergence.jpg/350px-Darwin_divergence.jpg" decoding="async" width="350" height="250" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/45/Darwin_divergence.jpg/525px-Darwin_divergence.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/45/Darwin_divergence.jpg/700px-Darwin_divergence.jpg 2x" data-file-width="1400" data-file-height="1000" /></a><figcaption>This tree diagram, used to show the divergence of species, is the only illustration in the Origin of Species.</figcaption></figure> Darwin proposes <a href="/wiki/Sexual_selection" title="Sexual selection">sexual selection</a>, driven by competition between males for mates, to explain <a href="/wiki/Sexual_dimorphism" title="Sexual dimorphism">sexually dimorphic</a> features such as lion manes, deer antlers, peacock tails, bird songs, and the bright plumage of some male birds.<a href="#cite_note-129">[129]</a> He analysed sexual selection more fully in <a href="/wiki/The_Descent_of_Man,_and_Selection_in_Relation_to_Sex" title="The Descent of Man, and Selection in Relation to Sex">The Descent of Man, and Selection in Relation to Sex</a> (1871). Natural selection was expected to work very slowly in forming new species, but given the effectiveness of artificial selection, he could "see no limit to the amount of change, to the beauty and infinite complexity of the coadaptations between all organic beings, one with another and with their physical conditions of life, which may be effected in the long course of time by nature's power of selection". Using a <a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">tree diagram</a> and calculations, he indicates the "divergence of character" from original species into new species and genera. He describes branches falling off as extinction occurred, while new branches formed in "the great <a href="/wiki/Tree_of_life_(science)" class="mw-redirect" title="Tree of life (science)">Tree of life</a> ... with its ever branching and beautiful ramifications".<a href="#cite_note-130">[130]</a> <div class="mw-heading mw-heading3"><h3 id="Variation_and_heredity">Variation and heredity</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=18" title="Edit section: Variation and heredity">edit</a>]</div> In Darwin's time there was no agreed-upon model of <a href="/wiki/Heredity" title="Heredity">heredity</a>;<a href="#cite_note-131">[131]</a> in Chapter I Darwin admitted, "The laws governing inheritance are quite unknown."<a href="#cite_note-132">[132]</a> He accepted a version of the inheritance of acquired characteristics (which after Darwin's death came to be called <a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a>), and Chapter V discusses what he called the effects of use and disuse; he wrote that he thought "there can be little doubt that use in our domestic animals strengthens and enlarges certain parts, and disuse diminishes them; and that such modifications are inherited", and that this also applied in nature.<a href="#cite_note-133">[133]</a> Darwin stated that some changes that were commonly attributed to use and disuse, such as the loss of functional wings in some island-dwelling insects, might be produced by natural selection. In later editions of Origin, Darwin expanded the role attributed to the inheritance of acquired characteristics. Darwin also admitted ignorance of the source of inheritable variations, but speculated they might be produced by environmental factors.<a href="#cite_note-134">[134]</a><a href="#cite_note-135">[135]</a> However, one thing was clear: whatever the exact nature and causes of new variations, Darwin knew from observation and experiment that breeders were able to select such variations and produce huge differences in many generations of selection.<a href="#cite_note-rez-119">[119]</a> The observation that selection works in domestic animals is not destroyed by lack of understanding of the underlying hereditary mechanism. Breeding of animals and plants showed related varieties varying in similar ways, or tending to revert to an ancestral form, and similar patterns of variation in distinct species were explained by Darwin as demonstrating <a href="/wiki/Common_descent" title="Common descent">common descent</a>. He recounted how <a href="/wiki/Lord_Morton%27s_mare" title="Lord Morton's mare">Lord Morton's mare</a> apparently demonstrated <a href="/wiki/Telegony_(pregnancy)" class="mw-redirect" title="Telegony (pregnancy)">telegony</a>, offspring inheriting characteristics of a previous mate of the female parent, and accepted this process as increasing the variation available for natural selection.<a href="#cite_note-136">[136]</a><a href="#cite_note-137">[137]</a> More detail was given in Darwin's 1868 book on <a href="/wiki/The_Variation_of_Animals_and_Plants_Under_Domestication" title="The Variation of Animals and Plants Under Domestication">The Variation of Animals and Plants Under Domestication</a>, which tried to explain heredity through his hypothesis of <a href="/wiki/Pangenesis" title="Pangenesis">pangenesis</a>. Although Darwin had privately questioned <a href="/wiki/Blending_inheritance" title="Blending inheritance">blending inheritance</a>, he struggled with the theoretical difficulty that novel individual variations would tend to blend into a population. However, inherited variation could be seen,<a href="#cite_note-flowering_of_genetics-138">[138]</a> and Darwin's concept of selection working on a population with a range of small variations was workable.<a href="#cite_note-Bowler_200-139">[139]</a> It was not until the <a href="/wiki/Extended_evolutionary_synthesis" title="Extended evolutionary synthesis">modern evolutionary synthesis</a> in the 1930s and 1940s that a model of heredity became completely integrated with a model of variation.<a href="#cite_note-140">[140]</a> This modern evolutionary synthesis had been dubbed Neo Darwinian Evolution because it encompasses <a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a>'s theories of evolution with <a href="/wiki/Gregor_Mendel" title="Gregor Mendel">Gregor Mendel</a>'s theories of genetic inheritance.<a href="#cite_note-141">[141]</a> <div class="mw-heading mw-heading3"><h3 id="Difficulties_for_the_theory">Difficulties for the theory</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=19" title="Edit section: Difficulties for the theory">edit</a>]</div> Chapter VI begins by saying the next three chapters will address possible objections to the theory, the first being that often no intermediate forms between closely related species are found, though the theory implies such forms must have existed. As Darwin noted, "Firstly, why, if species have descended from other species by insensibly fine gradations, do we not everywhere see innumerable transitional forms? Why is not all nature in confusion, instead of the species being, as we see them, well defined?"<a href="#cite_note-142">[142]</a> Darwin attributed this to the competition between different forms, combined with the small number of individuals of intermediate forms, often leading to extinction of such forms.<a href="#cite_note-143">[143]</a> Another difficulty, related to the first one, is the absence or rarity of transitional varieties in time. Darwin commented that by the theory of natural selection "innumerable transitional forms must have existed," and wondered "why do we not find them embedded in countless numbers in the crust of the earth?"<a href="#cite_note-144">[144]</a> (For further discussion of these difficulties, see <a href="/wiki/Speciation#Darwin's_dilemma:_why_species_exist" title="Speciation">Speciation#Darwin's dilemma: Why do species exist?</a> and Bernstein et al.<a href="#cite_note-145">[145]</a> and Michod.<a href="#cite_note-146">[146]</a>) The chapter then deals with whether natural selection could produce complex specialised structures, and the behaviours to use them, when it would be difficult to imagine how intermediate forms could be functional. Darwin said: <blockquote>Secondly, is it possible that an animal having, for instance, the structure and habits of a bat, could have been formed by the modification of some animal with wholly different habits? Can we believe that natural selection could produce, on the one hand, organs of trifling importance, such as the tail of a giraffe, which serves as a fly-flapper, and, on the other hand, organs of such wonderful structure, as the eye, of which we hardly as yet fully understand the inimitable perfection?<a href="#cite_note-147">[147]</a></blockquote> His answer was that in many cases animals exist with intermediate structures that are functional. He presented <a href="/wiki/Flying_squirrels" class="mw-redirect" title="Flying squirrels">flying squirrels</a>, and <a href="/wiki/Flying_lemurs" class="mw-redirect" title="Flying lemurs">flying lemurs</a> as examples of how bats might have evolved from non-flying ancestors.<a href="#cite_note-148">[148]</a> He discussed various simple eyes found in invertebrates, starting with nothing more than an optic nerve coated with pigment, as examples of how the <a href="/wiki/Evolution_of_the_eye" title="Evolution of the eye">vertebrate eye could have evolved</a>. Darwin concludes: "If it could be demonstrated that any complex organ existed, which could not possibly have been formed by numerous, successive, slight modifications, my theory would absolutely break down. But I can find out no such case."<a href="#cite_note-149">[149]</a> In a section on "organs of little apparent importance", Darwin discusses the difficulty of explaining various seemingly trivial traits with no evident adaptive function, and outlines some possibilities such as correlation with useful features. He accepts that we "are profoundly ignorant of the causes producing slight and unimportant variations" which distinguish <a href="/wiki/Domestication" title="Domestication">domesticated</a> breeds of animals,<a href="#cite_note-150">[150]</a> and <a href="/wiki/Race_(human_categorization)" title="Race (human categorization)">human races</a>. He suggests that <a href="/wiki/Sexual_selection" title="Sexual selection">sexual selection</a> might explain these variations:<a href="#cite_note-151">[151]</a><a href="#cite_note-FOOTNOTEDesmondMoore2009310-152">[152]</a> <blockquote>I might have adduced for this same purpose the differences between the races of man, which are so strongly marked; I may add that some little light can apparently be thrown on the origin of these differences, chiefly through sexual selection of a particular kind, but without here entering on copious details my reasoning would appear frivolous.<a href="#cite_note-OTOoS_199-153">[153]</a></blockquote> Chapter VII (of the first edition) addresses the evolution of instincts. His examples included two he had investigated experimentally: <a href="/wiki/Slave-making_ant" title="Slave-making ant">slave-making ants</a> and the construction of hexagonal cells by honey bees. Darwin noted that some species of slave-making ants were more dependent on slaves than others, and he observed that many ant species will collect and store the pupae of other species as food. He thought it reasonable that species with an extreme dependency on slave workers had evolved in incremental steps. He suggested that bees that make hexagonal cells evolved in steps from bees that made round cells, under pressure from natural selection to economise wax. Darwin concluded: <blockquote>Finally, it may not be a logical deduction, but to my imagination it is far more satisfactory to look at such instincts as the young cuckoo ejecting its foster-brothers, —ants making slaves, —the larvæ of ichneumonidæ feeding within the live bodies of caterpillars, —not as specially endowed or created instincts, but as small consequences of one general law, leading to the advancement of all organic beings, namely, multiply, vary, let the strongest live and the weakest die.<a href="#cite_note-154">[154]</a></blockquote> Chapter VIII addresses the idea that species had special characteristics that prevented hybrids from being fertile in order to preserve separately created species. Darwin said that, far from being constant, the difficulty in producing hybrids of related species, and the viability and fertility of the hybrids, varied greatly, especially among plants. Sometimes what were widely considered to be separate species produced fertile hybrid offspring freely, and in other cases what were considered to be mere varieties of the same species could only be crossed with difficulty. Darwin concluded: "Finally, then, the facts briefly given in this chapter do not seem to me opposed to, but even rather to support the view, that there is no fundamental distinction between species and varieties."<a href="#cite_note-155">[155]</a> In the sixth edition Darwin inserted a new chapter VII (renumbering the subsequent chapters) to respond to criticisms of earlier editions, including the objection that many features of organisms were not adaptive and could not have been produced by natural selection. He said some such features could have been by-products of adaptive changes to other features, and that often features seemed non-adaptive because their function was unknown, as shown by his book on <a href="/wiki/Fertilisation_of_Orchids" title="Fertilisation of Orchids">Fertilisation of Orchids</a> that explained how their elaborate structures facilitated pollination by insects. Much of the chapter responds to <a href="/wiki/George_Jackson_Mivart" class="mw-redirect" title="George Jackson Mivart">George Jackson Mivart</a>'s criticisms, including his claim that features such as <a href="/wiki/Baleen" title="Baleen">baleen</a> filters in whales, <a href="/wiki/Flatfish" title="Flatfish">flatfish</a> with both eyes on one side and the camouflage of <a href="/wiki/Phasmatodea" title="Phasmatodea">stick insects</a> could not have evolved through natural selection because intermediate stages would not have been adaptive. Darwin proposed scenarios for the incremental evolution of each feature.<a href="#cite_note-156">[156]</a> <div class="mw-heading mw-heading3"><h3 id="Geological_record">Geological record</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=20" title="Edit section: Geological record">edit</a>]</div> Chapter IX deals with the fact that the <a href="/wiki/Geological_record" class="mw-redirect" title="Geological record">geological record</a> appears to show forms of life suddenly arising, without the innumerable <a href="/wiki/Transitional_fossil" title="Transitional fossil">transitional fossils</a> expected from gradual changes. Darwin borrowed <a href="/wiki/Charles_Lyell" title="Charles Lyell">Charles Lyell</a>'s argument in <a href="/wiki/Principles_of_Geology" title="Principles of Geology">Principles of Geology</a> that the record is extremely imperfect as <a href="/wiki/Fossil" title="Fossil">fossilisation</a> is a very rare occurrence, spread over vast periods of time; since few areas had been geologically explored, there could only be fragmentary knowledge of <a href="/wiki/Formation_(stratigraphy)" class="mw-redirect" title="Formation (stratigraphy)">geological formations</a>, and fossil collections were very poor. Evolved local varieties which migrated into a wider area would seem to be the sudden appearance of a new species. Darwin did not expect to be able to reconstruct evolutionary history, but continuing discoveries gave him well-founded hope that new finds would occasionally reveal transitional forms.<a href="#cite_note-bowl182-157">[157]</a><a href="#cite_note-punkeek-158">[158]</a> To show that there had been enough time for natural selection to work slowly, he cited the example of <a href="/wiki/Weald_Basin" title="Weald Basin">The Weald</a> as discussed in Principles of Geology together with other observations from <a href="/wiki/Hugh_Miller" title="Hugh Miller">Hugh Miller</a>, <a href="/wiki/James_Smith_of_Jordanhill" title="James Smith of Jordanhill">James Smith of Jordanhill</a> and <a href="/wiki/Andrew_Ramsay_(geologist)" title="Andrew Ramsay (geologist)">Andrew Ramsay</a>. Combining this with an estimate of recent rates of sedimentation and erosion, Darwin calculated that erosion of The Weald had taken around 300 million years.<a href="#cite_note-159">[159]</a> The initial appearance of entire groups of well-developed organisms in the oldest fossil-bearing layers, now known as the <a href="/wiki/Cambrian_explosion" title="Cambrian explosion">Cambrian explosion</a>, posed a problem. Darwin had no doubt that earlier seas had swarmed with living creatures, but stated that he had no satisfactory explanation for the lack of fossils.<a href="#cite_note-160">[160]</a> Fossil evidence of <a href="/wiki/Pre-Cambrian" class="mw-redirect" title="Pre-Cambrian">pre-Cambrian</a> life has since been found, extending the history of life back for billions of years.<a href="#cite_note-161">[161]</a> Chapter X examines whether patterns in the fossil record are better explained by common descent and branching evolution through natural selection, than by the individual creation of fixed species. Darwin expected species to change slowly, but not at <a href="/wiki/Phyletic_gradualism" title="Phyletic gradualism">the same rate</a> – some organisms such as <a href="/wiki/Lingula_(genus)" class="mw-redirect" title="Lingula (genus)">Lingula</a> were unchanged since the earliest fossils. The pace of natural selection would depend on variability and change in the environment.<a href="#cite_note-darX-162">[162]</a> This distanced his theory from <a href="/wiki/Lamarckism" title="Lamarckism">Lamarckian</a> laws of inevitable progress.<a href="#cite_note-bowl182-157">[157]</a> It has been argued that this anticipated the <a href="/wiki/Punctuated_equilibrium" title="Punctuated equilibrium">punctuated equilibrium</a> hypothesis,<a href="#cite_note-punkeek-158">[158]</a><a href="#cite_note-163">[163]</a> but other scholars have preferred to emphasise Darwin's commitment to gradualism.<a href="#cite_note-164">[164]</a> He cited <a href="/wiki/Richard_Owen" title="Richard Owen">Richard Owen</a>'s findings that the earliest members of a class were a few simple and generalised species with characteristics intermediate between modern forms, and were followed by increasingly diverse and specialised forms, matching the branching of common descent from an ancestor.<a href="#cite_note-bowl182-157">[157]</a> Patterns of extinction matched his theory, with related groups of species having a continued existence until extinction, then not reappearing. Recently extinct species were more similar to living species than those from earlier eras, and as he had seen in South America, and <a href="/wiki/William_Clift" title="William Clift">William Clift</a> had shown in Australia, fossils from recent geological periods resembled species still living in the same area.<a href="#cite_note-darX-162">[162]</a> <div class="mw-heading mw-heading3"><h3 id="Geographic_distribution">Geographic distribution</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=21" title="Edit section: Geographic distribution">edit</a>]</div> Chapter XI deals with evidence from <a href="/wiki/Biogeography" title="Biogeography">biogeography</a>, starting with the observation that differences in flora and fauna from separate regions cannot be explained by environmental differences alone; South America, Africa, and Australia all have regions with similar climates at similar latitudes, but those regions have very different plants and animals. The species found in one area of a continent are more closely allied with species found in other regions of that same continent than to species found on other continents. Darwin noted that barriers to migration played an important role in the differences between the species of different regions. The coastal sea life of the Atlantic and Pacific sides of Central America had almost no species in common even though the <a href="/wiki/Isthmus_of_Panama" title="Isthmus of Panama">Isthmus of Panama</a> was only a few miles wide. His explanation was a combination of migration and descent with modification. He went on to say: "On this principle of inheritance with modification, we can understand how it is that sections of genera, whole genera, and even families are confined to the same areas, as is so commonly and notoriously the case."<a href="#cite_note-165">[165]</a> Darwin explained how a volcanic island formed a few hundred miles from a continent might be colonised by a few species from that continent. These species would become modified over time, but would still be related to species found on the continent, and Darwin observed that this was a common pattern. Darwin discussed ways that species could be dispersed across oceans to colonise islands, many of which he had investigated experimentally.<a href="#cite_note-166">[166]</a> Chapter XII continues the discussion of biogeography. After a brief discussion of freshwater species, it returns to oceanic islands and their peculiarities; for example on some islands roles played by mammals on continents were played by other animals such as flightless birds or reptiles. The summary of both chapters says: <blockquote>... I think all the grand leading facts of geographical distribution are explicable on the theory of migration (generally of the more dominant forms of life), together with subsequent modification and the multiplication of new forms. We can thus understand the high importance of barriers, whether of land or water, which separate our several zoological and botanical provinces. We can thus understand the localisation of sub-genera, genera, and families; and how it is that under different latitudes, for instance in South America, the inhabitants of the plains and mountains, of the forests, marshes, and deserts, are in so mysterious a manner linked together by affinity, and are likewise linked to the extinct beings which formerly inhabited the same continent ... On these same principles, we can understand, as I have endeavoured to show, why oceanic islands should have few inhabitants, but of these a great number should be endemic or peculiar; ...<a href="#cite_note-167">[167]</a></blockquote> <div class="mw-heading mw-heading3"><h3 id="Classification,_morphology,_embryology,_rudimentary_organs">Classification, morphology, embryology, rudimentary organs</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=22" title="Edit section: Classification, morphology, embryology, rudimentary organs">edit</a>]</div> Chapter XIII starts by observing that classification depends on species being grouped together in a <a href="/wiki/Taxonomy_(biology)" title="Taxonomy (biology)">Taxonomy</a>, a multilevel system of groups and sub-groups based on varying degrees of resemblance. After discussing classification issues, Darwin concludes: <blockquote>All the foregoing rules and aids and difficulties in classification are explained, if I do not greatly deceive myself, on the view that the natural system is founded on descent with modification; that the characters which naturalists consider as showing true affinity between any two or more species, are those which have been inherited from a common parent, and, in so far, all true classification is genealogical; that community of descent is the hidden bond which naturalists have been unconsciously seeking, ...<a href="#cite_note-168">[168]</a></blockquote> Darwin discusses <a href="/wiki/Morphology_(biology)" title="Morphology (biology)">morphology</a>, including the importance of <a href="/wiki/Homology_(biology)" title="Homology (biology)">homologous structures</a>. He says, "What can be more curious than that the hand of a man, formed for grasping, that of a mole for digging, the leg of the horse, the paddle of the porpoise, and the wing of the bat, should all be constructed on the same pattern, and should include the same bones, in the same relative positions?" This made no sense under doctrines of independent creation of species, as even <a href="/wiki/Richard_Owen" title="Richard Owen">Richard Owen</a> had admitted, but the "explanation is manifest on the theory of the natural selection of successive slight modifications" showing <a href="/wiki/Common_descent" title="Common descent">common descent</a>.<a href="#cite_note-169">[169]</a> He notes that animals of the same class often have extremely similar <a href="/wiki/Embryology" title="Embryology">embryos</a>. Darwin discusses rudimentary organs, such as the wings of flightless birds and the rudiments of pelvis and leg bones found in some snakes. He remarks that some rudimentary organs, such as teeth in <a href="/wiki/Baleen_whales" class="mw-redirect" title="Baleen whales">baleen whales</a>, are found only in embryonic stages.<a href="#cite_note-170">[170]</a> These factors also supported his theory of descent with modification.<a href="#cite_note-wyhe-31">[31]</a> <div class="mw-heading mw-heading3"><h3 id="Concluding_remarks">Concluding remarks</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=23" title="Edit section: Concluding remarks">edit</a>]</div> The final chapter, "Recapitulation and Conclusion", reviews points from earlier chapters, and Darwin concludes by hoping that his theory might produce revolutionary changes in many fields of natural history.<a href="#cite_note-171">[171]</a> He suggests that psychology will be put on a new foundation and implies the relevance of his theory to the <a href="/wiki/Human_evolution" title="Human evolution">first appearance of humanity</a> with the sentence that "Light will be thrown on the origin of man and his history."<a href="#cite_note-wyhe-31">[31]</a><a href="#cite_note-172">[172]</a> Darwin ends with a passage that became well known and much quoted: <blockquote>It is interesting to contemplate an entangled bank, clothed with many plants of many kinds, with birds singing on the bushes, with various insects flitting about, and with worms crawling through the damp earth, and to reflect that these elaborately constructed forms, so different from each other, and dependent on each other in so complex a manner, have all been produced by laws acting around us ... Thus, from the war of nature, from famine and death, the most exalted object which we are capable of conceiving, namely, the production of the higher animals, directly follows. There is grandeur in this view of life, with its several powers, having been originally breathed into a few forms or into one; and that, whilst this planet has gone cycling on according to the fixed law of gravity, from so simple a beginning endless forms most beautiful and most wonderful have been, and are being, evolved.<a href="#cite_note-173">[173]</a></blockquote> Darwin added the phrase "by the Creator" from the 1860 second edition onwards, so that the ultimate sentence begins "There is grandeur in this view of life, with its several powers, having been originally breathed by the Creator into a few forms or into one".<a href="#cite_note-2nd_edn_p_190-174">[174]</a> <div class="mw-heading mw-heading2"><h2 id="Structure,_style,_and_themes">Structure, style, and themes</h2>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=24" title="Edit section: Structure, style, and themes">edit</a>]</div> <div class="mw-heading mw-heading3"><h3 id="Nature_and_structure_of_Darwin's_argument">Nature and structure of Darwin's argument</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=25" title="Edit section: Nature and structure of Darwin's argument">edit</a>]</div> Darwin's aims were twofold: to show that species had not been separately created, and to show that <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a> had been the chief agent of change.<a href="#cite_note-175">[175]</a> He knew that his readers were already familiar with the concept of transmutation of species from <a href="/wiki/Vestiges_of_the_Natural_History_of_Creation" title="Vestiges of the Natural History of Creation">Vestiges</a>, and his introduction ridicules that work as failing to provide a viable mechanism.<a href="#cite_note-sec-176">[176]</a> Therefore, the first four chapters lay out his case that selection in nature, caused by the struggle for existence, is analogous to the selection of variations under domestication, and that the accumulation of adaptive variations provides a scientifically testable mechanism for evolutionary <a href="/wiki/Speciation" title="Speciation">speciation</a>.<a href="#cite_note-177">[177]</a><a href="#cite_note-bowl180-178">[178]</a> Later chapters provide evidence that evolution has occurred, supporting the idea of branching, adaptive evolution without directly proving that selection is the mechanism. Darwin presents supporting facts drawn from many disciplines, showing that his theory could explain a myriad of observations from many fields of natural history that were inexplicable under the alternative concept that species had been individually created.<a href="#cite_note-bowl180-178">[178]</a><a href="#cite_note-179">[179]</a><a href="#cite_note-180">[180]</a> The structure of Darwin's argument showed the influence of <a href="/wiki/John_Herschel" title="John Herschel">John Herschel</a>, whose philosophy of science maintained that a mechanism could be called a vera causa (true cause) if three things could be demonstrated: its existence in nature, its ability to produce the effects of interest, and its ability to explain a wide range of observations.<a href="#cite_note-181">[181]</a> This reflected the influence of <a href="/wiki/William_Whewell" title="William Whewell">William Whewell</a>'s idea of a consilience of inductions, as explained in his work Philosophy of the Inductive Sciences, where if you could argue that a proposed mechanism successfully explained various phenomena you could then use those arguments as evidence for that mechanism.<a href="#cite_note-182">[182]</a> <div class="mw-heading mw-heading3"><h3 id="Literary_style">Literary style</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=26" title="Edit section: Literary style">edit</a>]</div> The Examiner review of 3 December 1859 commented, "Much of Mr. Darwin's volume is what ordinary readers would call 'tough reading;' that is, writing which to comprehend requires concentrated attention and some preparation for the task. All, however, is by no means of this description, and many parts of the book abound in information, easy to comprehend and both instructive and entertaining."<a href="#cite_note-sec-176">[176]</a><a href="#cite_note-183">[183]</a> While the book was readable enough to sell, its dryness ensured that it was seen as aimed at specialist scientists and could not be dismissed as mere journalism or imaginative fiction. Though Richard Owen did complain in the Quarterly Review that the style was too easy for a serious work of science.<a href="#cite_note-184">[184]</a> Unlike the still-popular Vestiges, it avoided the narrative style of the historical novel and cosmological speculation, though the closing sentence clearly hinted at cosmic progression. Darwin had long been immersed in the literary forms and practices of specialist science, and made effective use of his skills in structuring arguments.<a href="#cite_note-sec-176">[176]</a> <a href="/wiki/David_Quammen" title="David Quammen">David Quammen</a> has described the book as written in everyday language for a wide audience, but noted that Darwin's literary style was uneven: in some places he used convoluted sentences that are difficult to read, while in other places his writing was beautiful. Quammen advised that later editions were weakened by Darwin making concessions and adding details to address his critics, and recommended the first edition.<a href="#cite_note-185">[185]</a> James T. Costa said that because the book was an abstract produced in haste in response to Wallace's essay, it was more approachable than the big book on natural selection Darwin had been working on, which would have been encumbered by scholarly footnotes and much more technical detail. He added that some parts of Origin are dense, but other parts are almost lyrical, and the case studies and observations are presented in a narrative style unusual in serious scientific books, which broadened its audience.<a href="#cite_note-186">[186]</a> <div class="mw-heading mw-heading3"><h3 id="Human_evolution">Human evolution</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=27" title="Edit section: Human evolution">edit</a>]</div> From his early transmutation notebooks in the late 1830s onwards, Darwin considered <a href="/wiki/Human_evolution" title="Human evolution">human evolution</a> as part of the natural processes he was investigating,<a href="#cite_note-JC_51-187">[187]</a> and rejected divine intervention.<a href="#cite_note-188">[188]</a> In 1856, his "big book on species" titled <a href="/wiki/Natural_Selection_(manuscript)" title="Natural Selection (manuscript)">Natural Selection</a> was to include a "note on Man", but when Wallace enquired in December 1857, Darwin replied; "You ask whether I shall discuss 'man';—I think I shall avoid whole subject, as so surrounded with prejudices, though I fully admit that it is the highest & most interesting problem for the naturalist."<a href="#cite_note-189">[189]</a><a href="#cite_note-Letter2192-190">[190]</a> On 28 March 1859, with his manuscript for the book well under way, Darwin wrote to Lyell offering the suggested publisher <a href="/wiki/John_Murray_III" title="John Murray III">John Murray</a> assurances "That I do not discuss origin of man".<a href="#cite_note-Letter_2437-64">[64]</a><a href="#cite_note-FOOTNOTEDesmondMoore2009306-65">[65]</a> In the final chapter of On the Origin of Species, "<a href="#Concluding_remarks">Recapitulation and Conclusion</a>", Darwin briefly highlights the human implications of his theory: <blockquote>"In the distant future I see open fields for far more important researches. Psychology will be based on a new foundation, that of the necessary acquirement of each mental power and capacity by gradation. Light will be thrown on the origin of man and his history."<a href="#cite_note-191">[191]</a></blockquote> Discussing this in January 1860, Darwin assured Lyell that "by the sentence [Light will be thrown on the origin of man and his history] I show that I believe man is in same predicament with other animals.<a href="#cite_note-Letter_2647-192">[192]</a> Many modern writers have seen this sentence as Darwin's only reference to humans in the book;<a href="#cite_note-JC_51-187">[187]</a> <a href="/wiki/Janet_Browne" title="Janet Browne">Janet Browne</a> describes it as his only discussion there of human origins, while noting that the book makes other references to humanity.<a href="#cite_note-193">[193]</a> Some other statements in the book are quietly effective at pointing out the implication that humans are simply another species, evolving through the same processes and principles affecting other organisms. For example,<a href="#cite_note-JC_51-187">[187]</a> in Chapter III: "Struggle for Existence" Darwin includes "slow-breeding man" among other examples of <a href="/wiki/Malthusian_trap" class="mw-redirect" title="Malthusian trap">Malthusian population growth</a>.<a href="#cite_note-194">[194]</a> In his discussions on <a href="/wiki/Morphology_(biology)" title="Morphology (biology)">morphology</a>, Darwin compares and comments on bone structures that are <a href="/wiki/Homology_(biology)" title="Homology (biology)">homologous</a> between humans and other mammals.<a href="#cite_note-195">[195]</a> Darwin's early notebooks discussed how non-adaptive characteristics could be selected when animals or humans chose mates,<a href="#cite_note-196">[196]</a> with <a href="/wiki/Race_(human_classification)" class="mw-redirect" title="Race (human classification)">races of humans</a> differing over ideas of beauty.<a href="#cite_note-197">[197]</a> In his 1856 notes responding to <a href="/wiki/Robert_Knox_(surgeon)" title="Robert Knox (surgeon)">Robert Knox</a>'s The Races of Man: A Fragment, he called this effect <a href="/wiki/Sexual_selection" title="Sexual selection">sexual selection</a>.<a href="#cite_note-FOOTNOTERichards2017[httpsbooksgooglecombooksidPAF2DgAAQBAJpgPA315_315],_[httpsbooksgooglecombooksidPAF2DgAAQBAJpgPA324_323–324]-198">[198]</a> He added notes on sexual selection to his "big book on species", and in mid-1857 he added a section heading "Theory applied to Races of Man", but did not add text on this topic.<a href="#cite_note-199">[199]</a> In On the Origin of Species, Chapter VI: "Difficulties on Theory", Darwin mentions this in the context of "slight and unimportant variations":<a href="#cite_note-OtOOS_197-200">[200]</a> <blockquote>I might have adduced for this same purpose the differences between the races of man, which are so strongly marked; I may add that some little light can apparently be thrown on the origin of these differences, chiefly through sexual selection of a particular kind, but without here entering on copious details my reasoning would appear frivolous."<a href="#cite_note-OtOOS_197-200">[200]</a></blockquote> When Darwin published <a href="/wiki/The_Descent_of_Man,_and_Selection_in_Relation_to_Sex" title="The Descent of Man, and Selection in Relation to Sex">The Descent of Man, and Selection in Relation to Sex</a> twelve years later, he said that he had not gone into detail on human evolution in the Origin as he thought that would "only add to the prejudices against my views". He had not completely avoided the topic:<a href="#cite_note-DOM_1-201">[201]</a> <blockquote>It seemed to me sufficient to indicate, in the first edition of my "Origin of Species," that by this work "light would be thrown on the origin of man and his history;" and this implies that man must be included with other organic beings in any general conclusion respecting his manner of appearance on this earth.<a href="#cite_note-DOM_1-201">[201]</a><a href="#cite_note-202">[202]</a></blockquote> He also said that he had "merely alluded" in that book to sexual selection differentiating human races.<a href="#cite_note-203">[203]</a> <div class="mw-heading mw-heading2"><h2 id="Reception">Reception</h2>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=28" title="Edit section: Reception">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Reactions_to_On_the_Origin_of_Species" title="Reactions to On the Origin of Species">Reactions to On the Origin of Species</a></div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Editorial_cartoon_depicting_Charles_Darwin_as_an_ape_(1871).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6f/Editorial_cartoon_depicting_Charles_Darwin_as_an_ape_%281871%29.jpg/150px-Editorial_cartoon_depicting_Charles_Darwin_as_an_ape_%281871%29.jpg" decoding="async" width="150" height="202" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6f/Editorial_cartoon_depicting_Charles_Darwin_as_an_ape_%281871%29.jpg/225px-Editorial_cartoon_depicting_Charles_Darwin_as_an_ape_%281871%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/6f/Editorial_cartoon_depicting_Charles_Darwin_as_an_ape_%281871%29.jpg/300px-Editorial_cartoon_depicting_Charles_Darwin_as_an_ape_%281871%29.jpg 2x" data-file-width="2068" data-file-height="2782" /></a><figcaption>In the 1870s, British caricatures of Darwin with a non-human <a href="/wiki/Ape" title="Ape">ape</a> body contributed to the identification of <a href="/wiki/Evolutionism" title="Evolutionism">evolutionism</a> with <a href="/wiki/Darwinism" title="Darwinism">Darwinism</a>.<a href="#cite_note-204">[204]</a></figcaption></figure> The book aroused international interest<a href="#cite_note-wyhe48-205">[205]</a> and a widespread debate, with no sharp line between scientific issues and ideological, social and religious implications.<a href="#cite_note-bowler177-206">[206]</a> Much of the initial reaction was hostile, in a large part because very few reviewers actually understood his theory,<a href="#cite_note-207">[207]</a> but Darwin had to be taken seriously as a prominent and respected name in science. Bishop <a href="/wiki/Samuel_Wilberforce" title="Samuel Wilberforce">Samuel Wilberforce</a> wrote a review in Quarterly Review in 1860<a href="#cite_note-208">[208]</a> where he disagreed with Darwin's 'argument'. There was much less controversy than had greeted the 1844 publication <a href="/wiki/Vestiges_of_Creation" class="mw-redirect" title="Vestiges of Creation">Vestiges of Creation</a>, which had been rejected by scientists,<a href="#cite_note-wyhe48-205">[205]</a> but had influenced a wide public readership into believing that nature and human society were governed by natural laws.<a href="#cite_note-wyhe-31">[31]</a> The Origin of Species as a book of wide general interest became associated with ideas of social reform. Its proponents made full use of a surge in the publication of review journals, and it was given more popular attention than almost any other scientific work, though it failed to match the continuing sales of Vestiges.<a href="#cite_note-209">[209]</a> Darwin's book legitimised scientific discussion of evolutionary mechanisms, and the newly coined term '<a href="/wiki/Darwinism" title="Darwinism">Darwinism</a>' was used to cover the whole range of <a href="/wiki/Evolutionism" title="Evolutionism">evolutionism</a>, not just his own ideas. By the mid-1870s, evolutionism was triumphant.<a href="#cite_note-bowler177-206">[206]</a> While Darwin had been somewhat coy about human origins, not identifying any explicit conclusion on the matter in his book, he had dropped enough hints about human's animal ancestry for the inference to be made,<a href="#cite_note-210">[210]</a><a href="#cite_note-Rad_Hux_Ket-211">[211]</a> and the first review claimed it made a creed of the "men from monkeys" idea from Vestiges.<a href="#cite_note-212">[212]</a><a href="#cite_note-213">[213]</a> <a href="/wiki/Human_evolution" title="Human evolution">Human evolution</a> became central to the debate and was strongly argued by <a href="/wiki/Thomas_Henry_Huxley" title="Thomas Henry Huxley">Huxley</a> who featured it in his popular "working-men's lectures". Darwin did not publish his own views on this until 1871.<a href="#cite_note-214">[214]</a><a href="#cite_note-215">[215]</a> The <a href="/wiki/Naturalism_(philosophy)" title="Naturalism (philosophy)">naturalism</a> of <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a> conflicted with presumptions of <a href="/wiki/Teleology" title="Teleology">purpose in nature</a> and while this could be reconciled by <a href="/wiki/Theistic_evolution" title="Theistic evolution">theistic evolution</a>, other mechanisms implying more progress or purpose were more acceptable. <a href="/wiki/Herbert_Spencer" title="Herbert Spencer">Herbert Spencer</a> had already incorporated <a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a> into his popular philosophy of progressive <a href="/wiki/Free_market" title="Free market">free market</a> human society. He popularised the terms 'evolution' and '<a href="/wiki/Survival_of_the_fittest" title="Survival of the fittest">survival of the fittest</a>', and many thought Spencer was central to evolutionary thinking.<a href="#cite_note-216">[216]</a> <div class="mw-heading mw-heading3"><h3 id="Impact_on_the_scientific_community">Impact on the scientific community</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=29" title="Edit section: Impact on the scientific community">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">History of evolutionary thought</a></div> Scientific readers were already aware of arguments that species changed through processes that were subject to <a href="/wiki/Physical_law" class="mw-redirect" title="Physical law">laws of nature</a>, but the transmutational ideas of Lamarck and the vague "law of development" of Vestiges had not found scientific favour. Darwin presented <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a> as a scientifically testable mechanism while accepting that other mechanisms such as <a href="/wiki/Inheritance_of_acquired_characters" class="mw-redirect" title="Inheritance of acquired characters">inheritance of acquired characters</a> were possible. His strategy established that evolution through natural laws was worthy of scientific study, and by 1875, most scientists accepted that evolution occurred but few thought natural selection was significant. Darwin's scientific method was also disputed, with his proponents favouring the <a href="/wiki/Empiricism" title="Empiricism">empiricism</a> of <a href="/wiki/John_Stuart_Mill" title="John Stuart Mill">John Stuart Mill</a>'s <a href="/wiki/A_System_of_Logic" title="A System of Logic">A System of Logic</a>, while opponents held to the idealist school of <a href="/wiki/William_Whewell" title="William Whewell">William Whewell</a>'s Philosophy of the Inductive Sciences, in which investigation could begin with the intuitive idea that species were fixed objects created by design.<a href="#cite_note-217">[217]</a> Early support for Darwin's ideas came from the findings of field naturalists studying <a href="/wiki/Biogeography" title="Biogeography">biogeography</a> and ecology, including <a href="/wiki/Joseph_Dalton_Hooker" title="Joseph Dalton Hooker">Joseph Dalton Hooker</a> in 1860, and <a href="/wiki/Asa_Gray" title="Asa Gray">Asa Gray</a> in 1862. <a href="/wiki/Henry_Walter_Bates" title="Henry Walter Bates">Henry Walter Bates</a> presented research in 1861 that explained <a href="/wiki/Batesian_mimicry" title="Batesian mimicry">insect mimicry</a> using natural selection. <a href="/wiki/Alfred_Russel_Wallace" title="Alfred Russel Wallace">Alfred Russel Wallace</a> discussed evidence from his <a href="/wiki/Malay_Archipelago" title="Malay Archipelago">Malay Archipelago</a> research, including an 1864 paper with an evolutionary explanation for the <a href="/wiki/Wallace_line" class="mw-redirect" title="Wallace line">Wallace line</a>.<a href="#cite_note-218">[218]</a> <figure class="mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Huxley_-_Mans_Place_in_Nature.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/91/Huxley_-_Mans_Place_in_Nature.jpg/300px-Huxley_-_Mans_Place_in_Nature.jpg" decoding="async" width="300" height="180" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/91/Huxley_-_Mans_Place_in_Nature.jpg/450px-Huxley_-_Mans_Place_in_Nature.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/91/Huxley_-_Mans_Place_in_Nature.jpg/600px-Huxley_-_Mans_Place_in_Nature.jpg 2x" data-file-width="1103" data-file-height="660" /></a><figcaption><a href="/wiki/Thomas_Henry_Huxley" title="Thomas Henry Huxley">Huxley</a> used illustrations to show that humans and apes had the same basic skeletal structure.<a href="#cite_note-219">[219]</a></figcaption></figure> Evolution had less obvious applications to <a href="/wiki/Anatomy" title="Anatomy">anatomy</a> and <a href="/wiki/Morphology_(biology)" title="Morphology (biology)">morphology</a>, and at first had little impact on the research of the anatomist <a href="/wiki/Thomas_Henry_Huxley" title="Thomas Henry Huxley">Thomas Henry Huxley</a>.<a href="#cite_note-Bowler184-185-220">[220]</a> Despite this, Huxley strongly supported Darwin on evolution; though he called for experiments to show whether natural selection could form new species, and questioned if Darwin's <a href="/wiki/Gradualism" title="Gradualism">gradualism</a> was sufficient without <a href="/wiki/Saltation_(biology)" title="Saltation (biology)">sudden leaps</a> to cause <a href="/wiki/Speciation" title="Speciation">speciation</a>. Huxley wanted science to be secular, without religious interference, and his article in the April 1860 <a href="/wiki/Westminster_Review" class="mw-redirect" title="Westminster Review">Westminster Review</a> promoted <a href="/wiki/Naturalism_(philosophy)" title="Naturalism (philosophy)">scientific naturalism</a> over natural theology,<a href="#cite_note-221">[221]</a><a href="#cite_note-westminster-222">[222]</a> praising Darwin for "extending the domination of Science over regions of thought into which she has, as yet, hardly penetrated" and coining the term "<a href="/wiki/Darwinism" title="Darwinism">Darwinism</a>" as part of his efforts to secularise and professionalise science.<a href="#cite_note-223">[223]</a> Huxley gained influence, and initiated the <a href="/wiki/X_Club" title="X Club">X Club</a>, which used the journal <a href="/wiki/Nature_(journal)" title="Nature (journal)">Nature</a> to promote evolution and naturalism, shaping much of late-Victorian science. Later, the German morphologist <a href="/wiki/Ernst_Haeckel" title="Ernst Haeckel">Ernst Haeckel</a> would convince Huxley that comparative anatomy and <a href="/wiki/Paleontology" title="Paleontology">palaeontology</a> could be used to reconstruct <a href="/wiki/Phylogenetics" title="Phylogenetics">evolutionary genealogies</a>.<a href="#cite_note-Bowler184-185-220">[220]</a><a href="#cite_note-224">[224]</a> The leading naturalist in Britain was the anatomist <a href="/wiki/Richard_Owen" title="Richard Owen">Richard Owen</a>, an idealist who had shifted to the view in the 1850s that the history of life was the gradual unfolding of a divine plan.<a href="#cite_note-225">[225]</a> Owen's review of the Origin in the April 1860 Edinburgh Review bitterly attacked Huxley, Hooker and Darwin, but also signalled acceptance of a kind of evolution as a <a href="/wiki/Teleology" title="Teleology">teleological</a> plan in a continuous "ordained becoming", with new species appearing by natural birth. Others that rejected natural selection, but supported "creation by birth", included the <a href="/wiki/George_Campbell,_8th_Duke_of_Argyll" title="George Campbell, 8th Duke of Argyll">Duke of Argyll</a> who explained beauty in plumage by design.<a href="#cite_note-226">[226]</a><a href="#cite_note-227">[227]</a> Since 1858, Huxley had emphasised anatomical similarities between apes and humans, contesting Owen's view that humans were a separate sub-class. Their disagreement over human origins came to the fore at the <a href="/wiki/British_Association_for_the_Advancement_of_Science" class="mw-redirect" title="British Association for the Advancement of Science">British Association for the Advancement of Science</a> meeting featuring the legendary <a href="/wiki/1860_Oxford_evolution_debate" title="1860 Oxford evolution debate">1860 Oxford evolution debate</a>.<a href="#cite_note-oed-228">[228]</a><a href="#cite_note-229">[229]</a> In two years of acrimonious public dispute that <a href="/wiki/Charles_Kingsley" title="Charles Kingsley">Charles Kingsley</a> satirised as the "<a href="/wiki/Great_Hippocampus_Question" title="Great Hippocampus Question">Great Hippocampus Question</a>" and parodied in <a href="/wiki/The_Water-Babies,_A_Fairy_Tale_for_a_Land_Baby" class="mw-redirect" title="The Water-Babies, A Fairy Tale for a Land Baby">The Water-Babies</a> as the "great hippopotamus test", Huxley showed that Owen was incorrect in asserting that ape brains lacked a structure present in human brains.<a href="#cite_note-230">[230]</a> Others, including <a href="/wiki/Charles_Lyell" title="Charles Lyell">Charles Lyell</a> and <a href="/wiki/Alfred_Russel_Wallace" title="Alfred Russel Wallace">Alfred Russel Wallace</a>, thought that humans shared a common ancestor with apes, but higher mental faculties could not have evolved through a purely material process. Darwin published his own explanation in the <a href="/wiki/Descent_of_Man" class="mw-redirect" title="Descent of Man">Descent of Man</a> (1871).<a href="#cite_note-231">[231]</a> <div class="mw-heading mw-heading4"><h4 id="Impact_outside_Great_Britain">Impact outside Great Britain</h4>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=30" title="Edit section: Impact outside Great Britain">edit</a>]</div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Pedigree_of_man_(Haeckel_1874).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/da/Pedigree_of_man_%28Haeckel_1874%29.jpg/220px-Pedigree_of_man_%28Haeckel_1874%29.jpg" decoding="async" width="220" height="381" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/da/Pedigree_of_man_%28Haeckel_1874%29.jpg/330px-Pedigree_of_man_%28Haeckel_1874%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/da/Pedigree_of_man_%28Haeckel_1874%29.jpg/440px-Pedigree_of_man_%28Haeckel_1874%29.jpg 2x" data-file-width="1200" data-file-height="2077" /></a><figcaption>Haeckel showed a main trunk leading to mankind with minor branches to various animals, unlike Darwin's branching evolutionary tree.<a href="#cite_note-Bowler190_191-232">[232]</a></figcaption></figure> The German physiologist <a href="/wiki/Emil_du_Bois-Reymond" title="Emil du Bois-Reymond">Emil du Bois-Reymond</a> converted to Darwinism after reading an English copy of On the Origin of Species in the spring of 1860. Du Bois-Reymond was a committed supporter, securing Darwin an honorary degree from the University of Breslau, teaching his theory to students at the University of Berlin, and defending his name to paying audiences across Germany and The Netherlands. Du Bois-Reymond's exposition resembled Darwin's: he endorsed natural selection, rejected the inheritance of acquired characters, remained silent on the origin of variation, and identified "the altruism of bees, the regeneration of tissue, the effects of exercise, and the inheritance of disadvantageous traits" as puzzles presented by the theory.<a href="#cite_note-233">[233]</a> Evolutionary ideas, although not natural selection, were accepted by other German biologists accustomed to ideas of <a href="/wiki/Homology_(biology)" title="Homology (biology)">homology</a> in <a href="/wiki/Morphology_(biology)" title="Morphology (biology)">morphology</a> from <a href="/wiki/Johann_Wolfgang_von_Goethe" title="Johann Wolfgang von Goethe">Goethe</a>'s <a href="/wiki/Metamorphosis_of_Plants" title="Metamorphosis of Plants">Metamorphosis of Plants</a> and from their long tradition of comparative anatomy. <a href="/wiki/Heinrich_Georg_Bronn" title="Heinrich Georg Bronn">Bronn</a>'s alterations in his German translation added to the misgivings of conservatives but encouraged political radicals. <a href="/wiki/Ernst_Haeckel" title="Ernst Haeckel">Ernst Haeckel</a> was particularly ardent, aiming to synthesise Darwin's ideas with those of <a href="/wiki/Lamarckism" title="Lamarckism">Lamarck</a> and Goethe while still reflecting the spirit of <a href="/wiki/Naturphilosophie" title="Naturphilosophie">Naturphilosophie</a>.<a href="#cite_note-browne140-98">[98]</a><a href="#cite_note-bowler186-234">[234]</a> His ambitious programme to reconstruct the <a href="/wiki/Evolutionary_history_of_life" class="mw-redirect" title="Evolutionary history of life">evolutionary history of life</a> was joined by Huxley and supported by discoveries in <a href="/wiki/Paleontology" title="Paleontology">palaeontology</a>. Haeckel used <a href="/wiki/Embryology" title="Embryology">embryology</a> extensively in his <a href="/wiki/Recapitulation_theory" title="Recapitulation theory">recapitulation theory</a>, which embodied a progressive, almost linear model of evolution. Darwin was cautious about such histories, and had already noted that <a href="/wiki/Karl_Ernst_von_Baer" title="Karl Ernst von Baer">von Baer's</a> laws of embryology supported his idea of complex branching.<a href="#cite_note-Bowler190_191-232">[232]</a> <a href="/wiki/Asa_Gray" title="Asa Gray">Asa Gray</a> promoted and defended Origin against those American naturalists with an idealist approach, notably <a href="/wiki/Louis_Agassiz" title="Louis Agassiz">Louis Agassiz</a>, who viewed every species as a distinct fixed unit in the mind of the Creator, classifying as species what others considered merely varieties.<a href="#cite_note-235">[235]</a> <a href="/wiki/Edward_Drinker_Cope" title="Edward Drinker Cope">Edward Drinker Cope</a> and <a href="/wiki/Alpheus_Hyatt" title="Alpheus Hyatt">Alpheus Hyatt</a> reconciled this view with evolutionism in a form of <a href="/wiki/Lamarckism#Neo-Lamarckism" title="Lamarckism">neo-Lamarckism</a> involving recapitulation theory.<a href="#cite_note-bowler186-234">[234]</a> French-speaking naturalists in several countries showed appreciation of the much-modified French translation by <a href="/wiki/Cl%C3%A9mence_Royer" title="Clémence Royer">Clémence Royer</a>, but Darwin's ideas had little impact in France, where any scientists supporting evolutionary ideas opted for a form of Lamarckism.<a href="#cite_note-browne142-101">[101]</a> The intelligentsia in Russia had accepted the general phenomenon of evolution for several years before Darwin had published his theory, and scientists were quick to take it into account, although the <a href="/wiki/Malthusianism" title="Malthusianism">Malthusian</a> aspects were felt to be relatively unimportant. The political economy of struggle was criticised as a British stereotype by <a href="/wiki/Karl_Marx" title="Karl Marx">Karl Marx</a> and by <a href="/wiki/Leo_Tolstoy" title="Leo Tolstoy">Leo Tolstoy</a>, who had the character Levin in his novel <a href="/wiki/Anna_Karenina" title="Anna Karenina">Anna Karenina</a> voice sharp criticism of the morality of Darwin's views.<a href="#cite_note-browne256-97">[97]</a> <div class="mw-heading mw-heading4"><h4 id="Challenges_to_natural_selection">Challenges to natural selection</h4>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=31" title="Edit section: Challenges to natural selection">edit</a>]</div> There were serious scientific objections to the process of <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a> as the key mechanism of evolution, including <a href="/wiki/Carl_N%C3%A4geli" title="Carl Nägeli">Carl Nägeli</a>'s insistence that a trivial characteristic with no adaptive advantage could not be developed by selection. Darwin conceded that these could be linked to adaptive characteristics. His estimate that the <a href="/wiki/Age_of_the_Earth" class="mw-redirect" title="Age of the Earth">age of the Earth</a> allowed gradual evolution was disputed by <a href="/wiki/William_Thomson,_1st_Baron_Kelvin" class="mw-redirect" title="William Thomson, 1st Baron Kelvin">William Thomson</a> (later awarded the title Lord Kelvin), who calculated that the <a href="/wiki/Sun" title="Sun">Sun</a>, and therefore life on Earth, was only about 100 million years old.<a href="#cite_note-236">[236]</a> Darwin accepted <a href="/wiki/Blending_inheritance" title="Blending inheritance">blending inheritance</a>, but <a href="/wiki/Fleeming_Jenkin" title="Fleeming Jenkin">Fleeming Jenkin</a> calculated that as it mixed traits, natural selection could not accumulate useful traits. Darwin tried to meet these objections in the fifth edition. <a href="/wiki/St._George_Jackson_Mivart" title="St. George Jackson Mivart">Mivart</a> supported directed evolution, and compiled scientific and religious objections to natural selection. In response, Darwin made considerable changes to the sixth edition. The problems of the age of the Earth and heredity were only resolved in the 20th century.<a href="#cite_note-miv-89">[89]</a><a href="#cite_note-237">[237]</a> By the mid-1870s, most scientists accepted evolution, but relegated natural selection to a minor role as they believed evolution was purposeful and progressive. The range of evolutionary theories during "<a href="/wiki/The_eclipse_of_Darwinism" title="The eclipse of Darwinism">the eclipse of Darwinism</a>" included forms of "<a href="/wiki/Saltation_(biology)" title="Saltation (biology)">saltationism</a>" in which new species were thought to arise through "jumps" rather than gradual adaptation, forms of <a href="/wiki/Orthogenesis" title="Orthogenesis">orthogenesis</a> claiming that species had an inherent tendency to change in a particular direction, and forms of neo-Lamarckism in which inheritance of acquired characteristics led to progress. The minority view of <a href="/wiki/August_Weismann" title="August Weismann">August Weismann</a>, that natural selection was the only mechanism, was called <a href="/wiki/Neo-Darwinism" title="Neo-Darwinism">neo-Darwinism</a>. It was thought that the rediscovery of <a href="/wiki/Mendelian_inheritance" title="Mendelian inheritance">Mendelian inheritance</a> invalidated Darwin's views.<a href="#cite_note-238">[238]</a><a href="#cite_note-Quammen205-234-239">[239]</a> <div class="mw-heading mw-heading3"><h3 id="Impact_on_economic_and_political_debates">Impact on economic and political debates</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=32" title="Edit section: Impact on economic and political debates">edit</a>]</div> While some, like Spencer, used analogy from natural selection as an argument against government intervention in the economy to benefit the poor, others, including <a href="/wiki/Alfred_Russel_Wallace" title="Alfred Russel Wallace">Alfred Russel Wallace</a>, argued that action was needed to correct social and economic inequities to level the playing field before natural selection could improve humanity further. Some political commentaries, including <a href="/wiki/Walter_Bagehot" title="Walter Bagehot">Walter Bagehot</a>'s Physics and Politics (1872), attempted to extend the idea of natural selection to competition between nations and between human races. Such ideas were incorporated into what was already an ongoing effort by some working in <a href="/wiki/Anthropology" title="Anthropology">anthropology</a> to provide scientific evidence for the superiority of <a href="/wiki/Caucasian_race" title="Caucasian race">Caucasians</a> over non-white races and justify European <a href="/wiki/Imperialism" title="Imperialism">imperialism</a>. Historians write that most such political and economic commentators had only a superficial understanding of Darwin's scientific theory, and were as strongly influenced by other concepts about social progress and evolution, such as the Lamarckian ideas of Spencer and Haeckel, as they were by Darwin's work. Darwin objected to his ideas being used to justify military aggression and unethical business practices as he believed morality was part of fitness in humans, and he opposed <a href="/wiki/Polygenism" title="Polygenism">polygenism</a>, the idea that human races were fundamentally distinct and did not share a recent common ancestry.<a href="#cite_note-240">[240]</a> <div class="mw-heading mw-heading3"><h3 id="Religious_attitudes">Religious attitudes</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=33" title="Edit section: Religious attitudes">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Rev_Baden_Powell.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f2/Rev_Baden_Powell.jpg/220px-Rev_Baden_Powell.jpg" decoding="async" width="220" height="281" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f2/Rev_Baden_Powell.jpg/330px-Rev_Baden_Powell.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/f/f2/Rev_Baden_Powell.jpg 2x" data-file-width="350" data-file-height="447" /></a><figcaption>The liberal theologian <a href="/wiki/Baden_Powell_(mathematician)" title="Baden Powell (mathematician)">Baden Powell</a> defended evolutionary ideas by arguing that the introduction of new species should be considered a natural rather than a miraculous process.<a href="#cite_note-241">[241]</a></figcaption></figure> The book produced a wide range of religious responses at a time of changing ideas and increasing secularisation. The issues raised were complex and there was a large middle ground. <a href="/wiki/History_of_geology" title="History of geology">Developments in geology</a> meant that there was little opposition based on a literal reading of <a href="/wiki/Book_of_Genesis" title="Book of Genesis">Genesis</a>,<a href="#cite_note-bowl202-242">[242]</a> but defence of the <a href="/wiki/Teleological_argument" title="Teleological argument">argument from design</a> and <a href="/wiki/Natural_theology" title="Natural theology">natural theology</a> was central to debates over the book in the English-speaking world.<a href="#cite_note-Dewey26-243">[243]</a><a href="#cite_note-Larson89-92-244">[244]</a> Natural theology was not a unified doctrine, and while some such as Louis Agassiz were strongly opposed to the ideas in the book, others sought a reconciliation in which evolution was seen as purposeful.<a href="#cite_note-bowl202-242">[242]</a> In the Church of England, some <a href="/wiki/Liberal_Christianity" title="Liberal Christianity">liberal clergymen</a> interpreted natural selection as an instrument of God's design, with the cleric Charles Kingsley seeing it as "just as noble a conception of Deity".<a href="#cite_note-Darwinanddesign-245">[245]</a><a href="#cite_note-246">[246]</a> In the second edition of January 1860, Darwin quoted Kingsley as "a celebrated cleric", and added the phrase "by the Creator" to the closing sentence, which from then on read "life, with its several powers, having been originally breathed by the Creator into a few forms or into one".<a href="#cite_note-2nd_edn_p_190-174">[174]</a> While some commentators have taken this as a concession to religion that Darwin later regretted,<a href="#cite_note-b95-84">[84]</a> Darwin's view at the time was of God creating life through the laws of nature,<a href="#cite_note-247">[247]</a><a href="#cite_note-248">[248]</a> and even in the first edition there are several references to "creation".<a href="#cite_note-249">[249]</a> <a href="/wiki/Baden_Powell_(mathematician)" title="Baden Powell (mathematician)">Baden Powell</a> praised "Mr Darwin's masterly volume [supporting] the grand principle of the self-evolving powers of nature".<a href="#cite_note-250">[250]</a> In America, <a href="/wiki/Asa_Gray" title="Asa Gray">Asa Gray</a> argued that evolution is the secondary effect, or modus operandi, of the first cause, design,<a href="#cite_note-251">[251]</a> and published a pamphlet defending the book in terms of <a href="/wiki/Theistic_evolution" title="Theistic evolution">theistic evolution</a>, Natural Selection is not inconsistent with Natural Theology.<a href="#cite_note-Darwinanddesign-245">[245]</a><a href="#cite_note-miles-252">[252]</a><a href="#cite_note-253">[253]</a> Theistic evolution became a popular compromise, and <a href="/wiki/St._George_Jackson_Mivart" title="St. George Jackson Mivart">St. George Jackson Mivart</a> was among those accepting evolution but attacking Darwin's naturalistic mechanism. Eventually it was realised that supernatural intervention could not be a scientific explanation, and naturalistic mechanisms such as <a href="/wiki/Neo-Lamarckism" class="mw-redirect" title="Neo-Lamarckism">neo-Lamarckism</a> were favoured over <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a> as being more compatible with purpose.<a href="#cite_note-bowl202-242">[242]</a> Even though the book did not explicitly spell out Darwin's beliefs about <a href="/wiki/Human_evolution" title="Human evolution">human origins</a>, it had dropped a number of hints about human's animal ancestry<a href="#cite_note-Rad_Hux_Ket-211">[211]</a> and quickly became central to the debate, as mental and moral qualities were seen as spiritual aspects of the immaterial <a href="/wiki/Soul_(spirit)" class="mw-redirect" title="Soul (spirit)">soul</a>, and it was believed that animals did not have spiritual qualities. This conflict could be reconciled by supposing there was some supernatural intervention on the path leading to humans, or viewing evolution as a purposeful and progressive ascent to mankind's position at the head of nature.<a href="#cite_note-bowl202-242">[242]</a> While many conservative theologians accepted evolution, <a href="/wiki/Charles_Hodge" title="Charles Hodge">Charles Hodge</a> argued in his 1874 critique "What is Darwinism?" that "<a href="/wiki/Darwinism" title="Darwinism">Darwinism</a>", defined narrowly as including rejection of design, was atheism though he accepted that Asa Gray did not reject design.<a href="#cite_note-254">[254]</a><a href="#cite_note-255">[255]</a> Asa Gray responded that this charge misrepresented Darwin's text.<a href="#cite_note-GrayWhatisDarwinism?-256">[256]</a> By the early 20th century, four noted authors of <a href="/wiki/The_Fundamentals" title="The Fundamentals">The Fundamentals</a> were explicitly open to the possibility that God created through evolution,<a href="#cite_note-257">[257]</a> but <a href="/wiki/Fundamentalism" title="Fundamentalism">fundamentalism</a> inspired the American <a href="/wiki/Creation%E2%80%93evolution_controversy" class="mw-redirect" title="Creation–evolution controversy">creation–evolution controversy</a> that began in the 1920s. Some conservative <a href="/wiki/Roman_Catholic" class="mw-redirect" title="Roman Catholic">Roman Catholic</a> writers and influential <a href="/wiki/Jesuit" class="mw-redirect" title="Jesuit">Jesuits</a> opposed evolution in the late 19th and early 20th century, but other Catholic writers, starting with Mivart, pointed out that early <a href="/wiki/Church_Fathers" title="Church Fathers">Church Fathers</a> had not interpreted Genesis literally in this area.<a href="#cite_note-258">[258]</a> The English cardinal <a href="/wiki/John_Henry_Newman" title="John Henry Newman">John Henry Newman</a> stated in a letter that “… Darwin's theory need not then to be atheistical, be it true or not; it may simply be suggesting a larger idea of Divine Prescience and Skill.”<a href="#cite_note-259">[259]</a> The <a href="/wiki/Holy_See" title="Holy See">Vatican</a> stated its <a href="/wiki/Evolution_and_the_Roman_Catholic_Church" class="mw-redirect" title="Evolution and the Roman Catholic Church">official position</a> in a <a href="/wiki/Humani_generis" title="Humani generis">1950 papal encyclical</a>, which held that evolution was not inconsistent with Catholic teaching.<a href="#cite_note-260">[260]</a><a href="#cite_note-261">[261]</a> <div class="mw-heading mw-heading2"><h2 id="Modern_influence">Modern influence</h2>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=34" title="Edit section: Modern influence">edit</a>]</div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:CollapsedtreeLabels-simplified.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/12/CollapsedtreeLabels-simplified.svg/300px-CollapsedtreeLabels-simplified.svg.png" decoding="async" width="300" height="198" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/12/CollapsedtreeLabels-simplified.svg/450px-CollapsedtreeLabels-simplified.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/12/CollapsedtreeLabels-simplified.svg/600px-CollapsedtreeLabels-simplified.svg.png 2x" data-file-width="800" data-file-height="528" /></a><figcaption>A modern <a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">phylogenetic tree</a> based on <a href="/wiki/Genome" title="Genome">genome</a> analysis shows the <a href="/wiki/Domain_(biology)" title="Domain (biology)">three-domain system</a>.</figcaption></figure> Various alternative evolutionary mechanisms favoured during "<a href="/wiki/The_eclipse_of_Darwinism" title="The eclipse of Darwinism">the eclipse of Darwinism</a>" became untenable as more was learned about inheritance and <a href="/wiki/Mutation" title="Mutation">mutation</a>. The full significance of <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a> was at last accepted in the 1930s and 1940s as part of the <a href="/wiki/Extended_evolutionary_synthesis" title="Extended evolutionary synthesis">modern evolutionary synthesis</a>. During that synthesis biologists and statisticians, including <a href="/wiki/R._A._Fisher" class="mw-redirect" title="R. A. Fisher">R. A. Fisher</a>, <a href="/wiki/Sewall_Wright" title="Sewall Wright">Sewall Wright</a> and <a href="/wiki/J._B._S._Haldane" title="J. B. S. Haldane">J. B. S. Haldane</a>, merged Darwinian selection with a statistical understanding of <a href="/wiki/Mendelian_genetics" class="mw-redirect" title="Mendelian genetics">Mendelian genetics</a>.<a href="#cite_note-Quammen205-234-239">[239]</a> <a href="/wiki/Evolution" title="Evolution">Modern evolutionary theory</a> continues to develop. Darwin's theory of evolution by natural selection, with its tree-like model of branching <a href="/wiki/Common_descent" title="Common descent">common descent</a>, has become the unifying theory of the <a href="/wiki/Life_science" class="mw-redirect" title="Life science">life sciences</a>. The theory explains the diversity of living organisms and their adaptation to the environment. It makes sense of the <a href="/wiki/Geological_record" class="mw-redirect" title="Geological record">geological record</a>, biogeography, parallels in <a href="/wiki/Embryo" title="Embryo">embryonic</a> development, <a href="/wiki/Homology_(biology)" title="Homology (biology)">biological homologies</a>, <a href="/wiki/Vestigiality" title="Vestigiality">vestigiality</a>, <a href="/wiki/Cladistics" title="Cladistics">cladistics</a>, <a href="/wiki/Phylogenetics" title="Phylogenetics">phylogenetics</a> and other fields, with unrivalled explanatory power; it has also become essential to applied sciences such as <a href="/wiki/Evolutionary_medicine" title="Evolutionary medicine">medicine</a> and agriculture.<a href="#cite_note-262">[262]</a><a href="#cite_note-263">[263]</a> Despite the scientific consensus, a religion-based political <a href="/wiki/Creation%E2%80%93evolution_controversy" class="mw-redirect" title="Creation–evolution controversy">controversy</a> has developed over how evolution is taught in schools, especially in the United States.<a href="#cite_note-264">[264]</a> Interest in Darwin's writings continues, and scholars have generated an extensive literature, the <a href="/wiki/Darwin_Industry" title="Darwin Industry">Darwin Industry</a>, about his life and work. The text of Origin itself has been subject to much analysis including a <a href="/wiki/Variorum" title="Variorum">variorum</a>, detailing the changes made in every edition, first published in 1959,<a href="#cite_note-265">[265]</a> and a <a href="/wiki/Concordance_(publishing)" title="Concordance (publishing)">concordance</a>, an exhaustive external index published in 1981.<a href="#cite_note-266">[266]</a> Worldwide commemorations of the 150th anniversary of the publication of On the Origin of Species and the bicentenary of Darwin's birth were scheduled for 2009.<a href="#cite_note-267">[267]</a> They celebrated the ideas which "over the last 150 years have revolutionised our understanding of nature and our place within it".<a href="#cite_note-268">[268]</a> In a survey conducted by a group of academic booksellers, publishers and librarians in advance of Academic Book Week in the United Kingdom, On the Origin of Species was voted the most influential academic book ever written.<a href="#cite_note-269">[269]</a> It was hailed as "the supreme demonstration of why academic books matter" and "a book which has changed the way we think about everything".<a href="#cite_note-270">[270]</a> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=35" title="Edit section: See also">edit</a>]</div> <style data-mw-deduplicate="TemplateStyles:r1266661725">.mw-parser-output .portalbox{padding:0;margin:0.5em 0;display:table;box-sizing:border-box;max-width:175px;list-style:none}.mw-parser-output .portalborder{border:1px solid var(--border-color-base,#a2a9b1);padding:0.1em;background:var(--background-color-neutral-subtle,#f8f9fa)}.mw-parser-output .portalbox-entry{display:table-row;font-size:85%;line-height:110%;height:1.9em;font-style:italic;font-weight:bold}.mw-parser-output .portalbox-image{display:table-cell;padding:0.2em;vertical-align:middle;text-align:center}.mw-parser-output .portalbox-link{display:table-cell;padding:0.2em 0.2em 0.2em 0.3em;vertical-align:middle}@media(min-width:720px){.mw-parser-output .portalleft{margin:0.5em 1em 0.5em 0}.mw-parser-output .portalright{clear:right;float:right;margin:0.5em 0 0.5em 1em}}</style><ul role="navigation" aria-label="Portals" class="noprint portalbox portalborder portalright"> <li class="portalbox-entry"><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/32px-Tree_of_life.svg.png" decoding="async" width="32" height="27" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/09/Tree_of_life.svg/48px-Tree_of_life.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/09/Tree_of_life.svg/64px-Tree_of_life.svg.png 2x" data-file-width="915" data-file-height="760" /></a><a href="/wiki/Portal:Evolutionary_biology" title="Portal:Evolutionary biology">Evolutionary biology portal</a></li></ul> <ul><li><a href="/wiki/Charles_Darwin_bibliography" title="Charles Darwin bibliography">Charles Darwin bibliography</a></li> <li><a href="/wiki/The_Complete_Works_of_Charles_Darwin_Online" title="The Complete Works of Charles Darwin Online">The Complete Works of Charles Darwin Online</a></li> <li><a href="/wiki/The_Descent_of_Man,_and_Selection_in_Relation_to_Sex" title="The Descent of Man, and Selection in Relation to Sex">The Descent of Man, and Selection in Relation to Sex</a>, published in 1871; his second major book on <a href="/wiki/Evolutionary_theory" class="mw-redirect" title="Evolutionary theory">evolutionary theory</a>.</li> <li><a href="/wiki/History_of_biology" title="History of biology">History of biology</a></li> <li><a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">History of evolutionary thought</a></li> <li><a href="/wiki/History_of_speciation" title="History of speciation">History of speciation</a></li> <li><a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern evolutionary synthesis</a></li> <li><a href="/wiki/Transmutation_of_species" title="Transmutation of species">Transmutation of species</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=36" title="Edit section: References">edit</a>]</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" style="column-width: 25em;"> <ol class="references"> <li id="cite_note-1"><a href="#cite_ref-1">^</a> <a href="#CITEREFDarwin1859">Darwin 1859</a>, p. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?viewtype=side&itemID=F373&pageseq=8">iii</a> </li> <li id="cite_note-Freeman_1977-2">^ <a href="#cite_ref-Freeman_1977_2-0">a</a> <a href="#cite_ref-Freeman_1977_2-1">b</a> <a href="#cite_ref-Freeman_1977_2-2">c</a> <a href="#cite_ref-Freeman_1977_2-3">d</a> <a href="#cite_ref-Freeman_1977_2-4">e</a> <a href="#CITEREFFreeman1977">Freeman 1977</a> </li> <li id="cite_note-titles-3">^ <a href="#cite_ref-titles_3-0">a</a> <a href="#cite_ref-titles_3-1">b</a> <a href="#cite_ref-titles_3-2">c</a> The book's full original title was On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life. In the 1872 sixth edition, "On" was omitted, so the full title is The origin of species by means of natural selection, or the preservation of favoured races in the struggle for life. This edition is usually known as The Origin of Species. The 6th is Darwin's final edition; there were minor modifications in the text of certain subsequent issues. See Freeman, R. B. "<a rel="nofollow" class="external text" href="http://darwin-online.org.uk/EditorialIntroductions/Freeman_OntheOriginofSpecies.html">The works of Charles Darwin: an annotated bibliographical handlist.</a>" In Van Wyhe, John, ed. Darwin Online: On the Origin of Species, 2002. </li> <li id="cite_note-FOOTNOTEDesmondMoore1991477-4"><a href="#cite_ref-FOOTNOTEDesmondMoore1991477_4-0">^</a> <a href="#CITEREFDesmondMoore1991">Desmond & Moore 1991</a>, p. 477. </li> <li id="cite_note-5"><a href="#cite_ref-5">^</a> <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 class="citation web cs1"><a rel="nofollow" class="external text" href="http://cudl.lib.cam.ac.uk/collections/darwin_mss">"Darwin Manuscripts (Digitised notes on Origin)"</a>. Cambridge Digital Library. Retrieved 24 November 2014.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Darwin+Manuscripts+%28Digitised+notes+on+Origin%29&rft.pub=Cambridge+Digital+Library&rft_id=http%3A%2F%2Fcudl.lib.cam.ac.uk%2Fcollections%2Fdarwin_mss&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> </li> <li id="cite_note-6"><a href="#cite_ref-6">^</a> <a href="#CITEREFMayr1982">Mayr 1982</a>, pp. 479–480 </li> <li id="cite_note-7"><a href="#cite_ref-7">^</a> <a href="#CITEREFDarwin1872">Darwin 1872</a>, p. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=F391&pageseq=18">xiii</a> </li> <li id="cite_note-8"><a href="#cite_ref-8">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAristotle" class="citation cs2">Aristotle, <a rel="nofollow" class="external text" href="http://classics.mit.edu/Aristotle/physics.2.ii.html">Physics</a>, translated by Hardie, R. 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Retrieved 16 January 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Letter+2443+%E2%80%94+Murray%2C+John+%28b%29+to+Darwin%2C+C.+R.%2C+1+April+1859&rft.pub=Darwin+Correspondence+Project&rft_id=http%3A%2F%2Fwww.darwinproject.ac.uk%2FDCP-LETT-2443&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> </li> <li id="cite_note-Letter_2445-69"><a href="#cite_ref-Letter_2445_69-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://www.darwinproject.ac.uk/DCP-LETT-2445">"Letter 2445 — Darwin, C. R. to Murray, John (b), 2 April (1859)"</a>. Darwin Correspondence Project. Retrieved 16 January 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Letter+2445+%E2%80%94+Darwin%2C+C.+R.+to+Murray%2C+John+%28b%29%2C+2+April+%281859%29&rft.pub=Darwin+Correspondence+Project&rft_id=http%3A%2F%2Fwww.darwinproject.ac.uk%2FDCP-LETT-2445&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> </li> <li id="cite_note-Darwin_and_his_publisher-70"><a href="#cite_ref-Darwin_and_his_publisher_70-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20101007052027/http://www.darwinproject.ac.uk/charles-darwin-and-john-murray">"Charles Darwin and his publisher"</a>. Darwin Correspondence Project. 2010. Archived from <a rel="nofollow" class="external text" href="http://www.darwinproject.ac.uk/charles-darwin-and-john-murray">the original</a> on 7 October 2010. Retrieved 7 September 2010.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Charles+Darwin+and+his+publisher&rft.pub=Darwin+Correspondence+Project&rft.date=2010&rft_id=http%3A%2F%2Fwww.darwinproject.ac.uk%2Fcharles-darwin-and-john-murray&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> </li> <li id="cite_note-Letter_2447-71"><a href="#cite_ref-Letter_2447_71-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://www.darwinproject.ac.uk/DCP-LETT-2447">"Letter 2447 — Darwin, C. R. to Murray, John (b), 5 April (1859)"</a>. Darwin Correspondence Project. Retrieved 16 January 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Letter+2447+%E2%80%94+Darwin%2C+C.+R.+to+Murray%2C+John+%28b%29%2C+5+April+%281859%29&rft.pub=Darwin+Correspondence+Project&rft_id=http%3A%2F%2Fwww.darwinproject.ac.uk%2FDCP-LETT-2447&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> </li> <li id="cite_note-72"><a href="#cite_ref-72">^</a> Darwin, C. R. [early draft title of Origin] <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?pageseq=1&itemID=CUL-DAR205.1.70&viewtype=side">On the mutability of species</a> [& other notes] CUL-DAR205.1.70 Transcribed by Kees Rookmaaker, edited by John van Wyhe </li> <li id="cite_note-Letter_2457A-73"><a href="#cite_ref-Letter_2457A_73-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://www.darwinproject.ac.uk/DCP-LETT-2457A">"Letter 2457A — Elwin, Whitwell, to Murray, John (b), 3 May 1859"</a>. Darwin Correspondence Project. Retrieved 16 January 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Letter+2457A+%E2%80%94+Elwin%2C+Whitwell%2C+to+Murray%2C+John+%28b%29%2C+3+May+1859&rft.pub=Darwin+Correspondence+Project&rft_id=http%3A%2F%2Fwww.darwinproject.ac.uk%2FDCP-LETT-2457A&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> </li> <li id="cite_note-Letter_2459-74"><a href="#cite_ref-Letter_2459_74-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://www.darwinproject.ac.uk/DCP-LETT-2459">"Letter 2459 — Darwin, C. R. to Murray, John (b), 6 May (1859)"</a>. Darwin Correspondence Project. Retrieved 16 January 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Letter+2459+%E2%80%94+Darwin%2C+C.+R.+to+Murray%2C+John+%28b%29%2C+6+May+%281859%29&rft.pub=Darwin+Correspondence+Project&rft_id=http%3A%2F%2Fwww.darwinproject.ac.uk%2FDCP-LETT-2459&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> </li> <li id="cite_note-Letter_2448-75"><a href="#cite_ref-Letter_2448_75-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://www.darwinproject.ac.uk/DCP-LETT-2488">"Letter 2448 — Darwin, C. R. to Murray, John (b), 10 September (1859)"</a>. Darwin Correspondence Project. Retrieved 16 January 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Letter+2448+%E2%80%94+Darwin%2C+C.+R.+to+Murray%2C+John+%28b%29%2C+10+September+%281859%29&rft.pub=Darwin+Correspondence+Project&rft_id=http%3A%2F%2Fwww.darwinproject.ac.uk%2FDCP-LETT-2488&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> </li> <li id="cite_note-NCSE_define_evo-76"><a href="#cite_ref-NCSE_define_evo_76-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://ncse.com/library-resource/defining-evolution-0">"Defining Evolution"</a>. <a href="/wiki/National_Center_for_Science_Education" title="National Center for Science Education">National Center for Science Education</a>. 24 August 2000. Retrieved 27 June 2016.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=National+Center+for+Science+Education&rft.atitle=Defining+Evolution&rft.date=2000-08-24&rft_id=https%3A%2F%2Fncse.com%2Flibrary-resource%2Fdefining-evolution-0&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> </li> <li id="cite_note-BernasconiLott2000-77"><a href="#cite_ref-BernasconiLott2000_77-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRobert_BernasconiTommy_Lee_Lott2000" class="citation book cs1">Robert Bernasconi; Tommy Lee Lott (2000). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=di2YVLKgIC8C&pg=PA54">The Idea of Race</a>. Hackett Publishing. p. 54. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/0-87220-458-8" title="Special:BookSources/0-87220-458-8"><bdi>0-87220-458-8</bdi></a>. <q>The full title [of the book] employs the term 'race' only in the broad biological use of the word, which refers to varieties throughout organic life; however, speculation about the implications of his views specifically for the question of the human races began almost as soon as the book was published.</q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Idea+of+Race&rft.pages=54&rft.pub=Hackett+Publishing&rft.date=2000&rft.isbn=0-87220-458-8&rft.au=Robert+Bernasconi&rft.au=Tommy+Lee+Lott&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3Ddi2YVLKgIC8C%26pg%3DPA54&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> </li> <li id="cite_note-Sober45-78"><a href="#cite_ref-Sober45_78-0">^</a> <a href="#CITEREFSober2011">Sober 2011</a>, p. <a rel="nofollow" class="external text" href="https://books.google.com/books?id=nHEGJY9soYMC&pg=PT38">45</a>, Quote: "There nonetheless are a few cases in which Darwin does discuss selection processes in which groups are the units, and these will be the focus of the present chapter. But even here it does not matter whether the groups are from different 'races' or from the same race. It is nests of honeybees that compete with each other, and human tribes that compete with other human tribes. For Darwin, the question of group selection had nothing special to do with 'race.' Still, writing in the heyday of empire, Darwin saw European nations outcompeting the nations, kingdoms, and tribes that occupy the rest of the globe. In this one very salient example, Darwin did see races struggling with each other. In any event, the word race in Darwin's subtitle needs to be understood very broadly; it encompasses competition among individuals, competition among groups in the same 'race,' and competition from groups from different 'races.' This is a much broader meaning than the word 'race' tends to have today." </li> <li id="cite_note-79"><a href="#cite_ref-79">^</a> <a href="#CITEREFDarwin1859">Darwin 1859</a>, p. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=F373&pageseq=30">15</a> </li> <li id="cite_note-80"><a href="#cite_ref-80">^</a> the three instances of the phrase "races of man" are found on <a href="#CITEREFDarwin1859">Darwin 1859</a>, pp. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?pageseq=217&itemID=F373&viewtype=text">199</a>, <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?pageseq=400&itemID=F373&viewtype=text">382</a>, <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?pageseq=440&itemID=F373&viewtype=text">422</a> </li> <li id="cite_note-dupree-81"><a href="#cite_ref-dupree_81-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFDupree1988" class="citation book cs1">Dupree, A. Hunter (1988). Asa Gray, American Botanist, Friend of Darwin. Baltimore, MD: Johns Hopkins University Press. p. 267. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-8018-3741-8" title="Special:BookSources/978-0-8018-3741-8"><bdi>978-0-8018-3741-8</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Asa+Gray%2C+American+Botanist%2C+Friend+of+Darwin&rft.place=Baltimore%2C+MD&rft.pages=267&rft.pub=Johns+Hopkins+University+Press&rft.date=1988&rft.isbn=978-0-8018-3741-8&rft.aulast=Dupree&rft.aufirst=A.+Hunter&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> </li> <li id="cite_note-82"><a href="#cite_ref-82">^</a> <a href="#CITEREFBrowne2002">Browne 2002</a>, p. 89 </li> <li id="cite_note-83"><a href="#cite_ref-83">^</a> <a href="#CITEREFDarwin1958">Darwin 1958</a>, p. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?itemID=F1497&viewtype=text&pageseq=126">122</a> </li> <li id="cite_note-b95-84">^ <a href="#cite_ref-b95_84-0">a</a> <a href="#cite_ref-b95_84-1">b</a> <a href="#cite_ref-b95_84-2">c</a> <a href="#CITEREFBrowne2002">Browne 2002</a>, pp. 95–96 </li> <li id="cite_note-85"><a href="#cite_ref-85">^</a> <a href="#CITEREFDarwin1861">Darwin 1861</a>, p. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?itemID=F381&viewtype=text&pageseq=20">xiii</a> </li> <li id="cite_note-old_Darwin_manuscript_2022-86"><a href="#cite_ref-old_Darwin_manuscript_2022_86-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://news.nus.edu.sg/secret-of-157-year-old-darwin-manuscript/">"Science ahead of its time: Secret of 157-year old Darwin manuscript"</a>. <a href="/wiki/National_University_of_Singapore" title="National University of Singapore">National University of Singapore</a> News. 24 November 2022. Retrieved 25 November 2022.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=National+University+of+Singapore+News&rft.atitle=Science+ahead+of+its+time%3A+Secret+of+157-year+old+Darwin+manuscript&rft.date=2022-11-24&rft_id=https%3A%2F%2Fnews.nus.edu.sg%2Fsecret-of-157-year-old-darwin-manuscript%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFDarwin1865" class="citation web cs1">Darwin, C. R. (October 1865). <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?pageseq=1&itemID=Sothebys-N11124&viewtype=side">"Signed autograph paragraph from Origin of species 3d ed. for Hermann Kindt"</a>. Darwin Online. Retrieved 25 November 2022. <q>Introduction by John van Wyhe</q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Darwin+Online&rft.atitle=Signed+autograph+paragraph+from+Origin+of+species+3d+ed.+for+Hermann+Kindt&rft.date=1865-10&rft.aulast=Darwin&rft.aufirst=C.+R.&rft_id=http%3A%2F%2Fdarwin-online.org.uk%2Fcontent%2Fframeset%3Fpageseq%3D1%26itemID%3DSothebys-N11124%26viewtype%3Dside&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> </li> <li id="cite_note-87"><a href="#cite_ref-87">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation news cs1"><a rel="nofollow" class="external text" href="https://www.bbc.com/news/science-environment-63741207">"Charles Darwin: Autographed document could fetch record price"</a>. BBC News. 25 November 2022. Retrieved 25 November 2022.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=BBC+News&rft.atitle=Charles+Darwin%3A+Autographed+document+could+fetch+record+price&rft.date=2022-11-25&rft_id=https%3A%2F%2Fwww.bbc.com%2Fnews%2Fscience-environment-63741207&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> </li> <li id="cite_note-sotf-88"><a href="#cite_ref-sotf_88-0">^</a> "This survival of the fittest, which I have here sought to express in mechanical terms, is that which Mr. Darwin has called 'natural selection', or the preservation of favoured races in the struggle for life." <a href="#CITEREFSpencer1864">Spencer 1864</a>, pp. <a rel="nofollow" class="external text" href="https://books.google.com/books?id=SRkRAAAAYAAJ&pg=PA444">444–445</a> </li> <li id="cite_note-miv-89">^ <a href="#cite_ref-miv_89-0">a</a> <a href="#cite_ref-miv_89-1">b</a> <a href="#CITEREFMivart1871">Mivart 1871</a> </li> <li id="cite_note-90"><a href="#cite_ref-90">^</a> <a href="#CITEREFBrowne2002">Browne 2002</a>, p. 59 </li> <li id="cite_note-91"><a href="#cite_ref-91">^</a> <a href="#CITEREFFreeman1977">Freeman 1977</a>, pp. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?pageseq=80&itemID=A1&viewtype=text">79–80</a>. "Evolution" in the transformist sense had been used by <a href="/wiki/Charles_Lyell" title="Charles Lyell">Charles Lyell</a> in 1832, <a href="/wiki/Principles_of_Geology" title="Principles of Geology">Principles of Geology</a> vol 2, <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=A505.2&pageseq=24">p. 11</a>; and was used by Darwin in <a href="/wiki/The_Descent_of_Man" class="mw-redirect" title="The Descent of Man">The Descent of Man</a> in 1871, <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=F937.1&pageseq=15">p. 2</a> onwards. </li> <li id="cite_note-6th-92">^ <a href="#cite_ref-6th_92-0">a</a> <a href="#cite_ref-6th_92-1">b</a> <a href="#CITEREFDesmondMoore1991">Desmond & Moore 1991</a>, pp. 577, 582, 590, 592–593 </li> <li id="cite_note-Letter2592-93"><a href="#cite_ref-Letter2592_93-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation cs2"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20090213042956/http://www.darwinproject.ac.uk/darwinletters/calendar/entry-2592.html#mark-2592.f3">Darwin Correspondence Project – Letter 2592—Darwin, C. R. to Gray, Asa, 21 December (1859)</a>, archived from <a rel="nofollow" class="external text" href="http://www.darwinproject.ac.uk/darwinletters/calendar/entry-2592.html#mark-2592.f3">the original</a> on 13 February 2009, retrieved 6 December 2008</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Darwin+Correspondence+Project+%E2%80%93+Letter+2592%E2%80%94Darwin%2C+C.+R.+to+Gray%2C+Asa%2C+21+December+%281859%29&rft_id=http%3A%2F%2Fwww.darwinproject.ac.uk%2Fdarwinletters%2Fcalendar%2Fentry-2592.html%23mark-2592.f3&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> </li> <li id="cite_note-Letter2665-94"><a href="#cite_ref-Letter2665_94-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation cs2"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20090213042959/http://www.darwinproject.ac.uk/darwinletters/calendar/entry-2665.html#back-mark-2665.f9">Darwin Correspondence Project – Letter 2665—Darwin, C. 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Biol. 117 (4): 665–90. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1985JThBi.117..665B">1985JThBi.117..665B</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2FS0022-5193%2885%2980246-0">10.1016/S0022-5193(85)80246-0</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/4094459">4094459</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=J.+Theor.+Biol.&rft.atitle=Sex+and+the+emergence+of+species&rft.volume=117&rft.issue=4&rft.pages=%3Cspan+class%3D%22nowrap%22%3E665-%3C%2Fspan%3E90&rft.date=1985-12&rft_id=info%3Apmid%2F4094459&rft_id=info%3Adoi%2F10.1016%2FS0022-5193%2885%2980246-0&rft_id=info%3Abibcode%2F1985JThBi.117..665B&rft.au=Bernstein+H.&rft.au=Byerly+H.C.&rft.au=Hopf+F.A.&rft.au=Michod+R.E.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> </li> <li id="cite_note-146"><a href="#cite_ref-146">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMichod,_Richard_E.1995" class="citation book cs1">Michod, Richard E. 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href="http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=F373&pageseq=324">306–308</a> </li> <li id="cite_note-161"><a href="#cite_ref-161">^</a> <a href="#CITEREFSchopf2000">Schopf 2000</a> </li> <li id="cite_note-darX-162">^ <a href="#cite_ref-darX_162-0">a</a> <a href="#cite_ref-darX_162-1">b</a> <a href="#CITEREFDarwin1859">Darwin 1859</a>, pp. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=F373&pageseq=330">312–345</a> </li> <li id="cite_note-163"><a href="#cite_ref-163">^</a> <a href="#CITEREFRhodes1987">Rhodes 1987</a> </li> <li id="cite_note-164"><a href="#cite_ref-164">^</a> <a href="#CITEREFDarwinCosta2009">Darwin & Costa 2009</a>, p. 108 </li> <li id="cite_note-165"><a href="#cite_ref-165">^</a> <a href="#CITEREFDarwin1859">Darwin 1859</a>, pp. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=F373&pageseq=368">350–351</a> </li> <li id="cite_note-166"><a href="#cite_ref-166">^</a> <a href="#CITEREFDarwin1859">Darwin 1859</a>, pp. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=F373&pageseq=364">346–382</a> </li> <li id="cite_note-167"><a href="#cite_ref-167">^</a> <a href="#CITEREFDarwin1859">Darwin 1859</a>, pp. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=F373&pageseq=426">408–409</a> </li> <li id="cite_note-168"><a href="#cite_ref-168">^</a> <a href="#CITEREFDarwin1859">Darwin 1859</a>, p. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=F373&pageseq=438">420</a> </li> <li id="cite_note-169"><a href="#cite_ref-169">^</a> <a href="#CITEREFDarwin1859">Darwin 1859</a>, pp. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=F373&pageseq=452">434–435</a> </li> <li id="cite_note-170"><a href="#cite_ref-170">^</a> <a href="#CITEREFDarwin1859">Darwin 1859</a>, pp. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?pageseq=468&itemID=F373&viewtype=text">450–451</a> </li> <li id="cite_note-171"><a href="#cite_ref-171">^</a> <a href="#CITEREFDarwin1859">Darwin 1859</a>, pp. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?pageseq=502&itemID=F373&viewtype=text">484–488</a>, Quote: "When the views entertained in this volume on the origin of species, or when analogous views are generally admitted, we can dimly foresee that there will be a considerable revolution in natural history. ..." </li> <li id="cite_note-172"><a href="#cite_ref-172">^</a> <a href="#CITEREFDarwin1859">Darwin 1859</a>, p. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=F373&pageseq=506">488</a> <a href="#CITEREFDarwin1871">Darwin 1871</a>, p. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?pageseq=14&itemID=F937.1&viewtype=text">1</a>, Quote: "… this implies that man must be included with other organic beings in any general conclusion respecting his manner of appearance on this earth." </li> <li id="cite_note-173"><a href="#cite_ref-173">^</a> <a href="#CITEREFDarwin1859">Darwin 1859</a>, pp. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=F373&pageseq=507">489–490</a> </li> <li id="cite_note-2nd_edn_p_190-174">^ <a href="#cite_ref-2nd_edn_p_190_174-0">a</a> <a href="#cite_ref-2nd_edn_p_190_174-1">b</a> <a href="#CITEREFDarwin1860">Darwin 1860</a>, p. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=F376&pageseq=508">490</a> </li> <li id="cite_note-175"><a href="#cite_ref-175">^</a> <a href="#CITEREFDarwin1871">Darwin 1871</a>, p. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=F937.1&pageseq=165">152</a> </li> <li id="cite_note-sec-176">^ <a href="#cite_ref-sec_176-0">a</a> <a href="#cite_ref-sec_176-1">b</a> <a href="#cite_ref-sec_176-2">c</a> <a href="#CITEREFSecord2000">Secord 2000</a>, pp. 508–511 </li> <li id="cite_note-177"><a href="#cite_ref-177">^</a> <a href="#CITEREFQuammen2006">Quammen 2006</a>, pp. 183–188 </li> <li id="cite_note-bowl180-178">^ <a href="#cite_ref-bowl180_178-0">a</a> <a href="#cite_ref-bowl180_178-1">b</a> <a href="#CITEREFBowler2003">Bowler 2003</a>, pp. 180–181 </li> <li id="cite_note-179"><a href="#cite_ref-179">^</a> <a href="#CITEREFQuammen2006">Quammen 2006</a>, pp. 190, 200–201 </li> <li id="cite_note-180"><a href="#cite_ref-180">^</a> <a href="#CITEREFLarson2004">Larson 2004</a>, pp. 88–89 </li> <li id="cite_note-181"><a href="#cite_ref-181">^</a> <a href="#CITEREFDarwinCosta2009">Darwin & Costa 2009</a>, p. xvii </li> <li id="cite_note-182"><a href="#cite_ref-182">^</a> <a href="#CITEREFRuse2009">Ruse 2009</a>, pp. 22–23 </li> <li id="cite_note-183"><a href="#cite_ref-183">^</a> <a href="#CITEREFCrawford1859">Crawford 1859</a> </li> <li id="cite_note-184"><a href="#cite_ref-184">^</a> <a href="#CITEREFRuse2009">Ruse 2009</a>, p. 18 </li> <li id="cite_note-185"><a href="#cite_ref-185">^</a> <a href="#CITEREFQuammen2006">Quammen 2006</a>, pp. 176–181 </li> <li id="cite_note-186"><a href="#cite_ref-186">^</a> <a href="#CITEREFDarwinCosta2009">Darwin & Costa 2009</a>, p. ix </li> <li id="cite_note-JC_51-187">^ <a href="#cite_ref-JC_51_187-0">a</a> <a href="#cite_ref-JC_51_187-1">b</a> <a href="#cite_ref-JC_51_187-2">c</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCarroll2003" class="citation book cs1"><a href="/wiki/Joseph_Carroll_(scholar)" title="Joseph Carroll (scholar)">Carroll, Joseph</a> (2003). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=eTfRotZTXI0C&pg=PA51">On the Origin of Species / Charles Darwin</a>. Broadview Press. pp. 51–52. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/1-55111-337-6" title="Special:BookSources/1-55111-337-6"><bdi>1-55111-337-6</bdi></a>. <q>Following Darwin's lead, most commentators cite this one passage as the only reference to man in the Origin, but they thus overlook, as did Darwin himself, two sentences that are, in their own quiet way, even more effective.</q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=On+the+Origin+of+Species+%2F+Charles+Darwin&rft.pages=%3Cspan+class%3D%22nowrap%22%3E51-%3C%2Fspan%3E52&rft.pub=Broadview+Press&rft.date=2003&rft.isbn=1-55111-337-6&rft.aulast=Carroll&rft.aufirst=Joseph&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DeTfRotZTXI0C%26pg%3DPA51&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> </li> <li id="cite_note-188"><a href="#cite_ref-188">^</a> <a href="#CITEREFBrowne2007">Browne 2007</a>, p. 42, quoting <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?pageseq=172&itemID=CUL-DAR122.-&viewtype=text">Darwin, C. R. Notebook C</a> (February to July 1838) pp. 196–197 "Man in his arrogance thinks himself a great work worthy the interposition of a deity, more humble & I believe truer to consider him created from animals." </li> <li id="cite_note-189"><a href="#cite_ref-189">^</a> <a href="#CITEREFDesmondMoore1991">Desmond & Moore 1991</a>, pp. 412–441, 457–458, 462–463 <a href="#CITEREFDesmondMoore2009">Desmond & Moore 2009</a>, pp. 283–284, 290–292, 295 </li> <li id="cite_note-Letter2192-190"><a href="#cite_ref-Letter2192_190-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://www.darwinproject.ac.uk/DCP-LETT-2192">"Letter 2192 – Darwin, C. R. to Wallace, A. R., 22 December 1857"</a>. Darwin Correspondence Project.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Letter+2192+%E2%80%93+Darwin%2C+C.+R.+to+Wallace%2C+A.+R.%2C+22+December+1857&rft.pub=Darwin+Correspondence+Project&rft_id=http%3A%2F%2Fwww.darwinproject.ac.uk%2FDCP-LETT-2192&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> </li> <li id="cite_note-191"><a href="#cite_ref-191">^</a> <a href="#CITEREFDarwin1871">Darwin 1871</a>, p. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=F373&pageseq=506">488</a> </li> <li id="cite_note-Letter_2647-192"><a href="#cite_ref-Letter_2647_192-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://www.darwinproject.ac.uk/DCP-LETT-2647">"Letter 2647 – Darwin, C. R. to Charles Lyell, 10 January (1860)"</a>. Darwin Correspondence Project. Retrieved 18 September 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Letter+2647+%E2%80%93+Darwin%2C+C.+R.+to+Charles+Lyell%2C+10+January+%281860%29&rft.pub=Darwin+Correspondence+Project&rft_id=http%3A%2F%2Fwww.darwinproject.ac.uk%2FDCP-LETT-2647&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> </li> <li id="cite_note-193"><a href="#cite_ref-193">^</a> For example, <a href="#CITEREFBrowne2002">Browne 2002</a>, p. 60, "In this book, he was completely silent on the subject of human origins, although he did refer in several places to mankind as an example of biological details. The only words he allowed himself—and these out of a sense of duty that he must somewhere refer to human beings–were gnomic in their brevity. 'Light will be thrown on the origin of man and his history'." </li> <li id="cite_note-194"><a href="#cite_ref-194">^</a> <a href="#CITEREFDarwin1859">Darwin 1859</a>, p. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?pageseq=79&itemID=F373&viewtype=text">64</a>, Quote: "There is no exception to the rule that every organic being naturally increases at so high a rate, that if not destroyed, the earth would soon be covered by the progeny of a single pair. Even slow-breeding man has doubled in twenty-five years, and at this rate, in a few thousand years, there would literally not be standing room for his progeny." </li> <li id="cite_note-195"><a href="#cite_ref-195">^</a> <a href="#CITEREFvan_Wyhe2008">van Wyhe 2008</a> <a href="#CITEREFDarwin1859">Darwin 1859</a>, p. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?pageseq=452&itemID=F373&viewtype=text">434</a>, Quote: "What can be more curious than that the hand of a man, formed for grasping, that of a mole for digging, the leg of the horse, the paddle of the porpoise, and the wing of the bat, should all be constructed on the same pattern, and should include the same bones, in the same relative positions?" <a href="#CITEREFDarwin1859">Darwin 1859</a>, p. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?pageseq=497&itemID=F373&viewtype=text">479</a>, Quote: "The framework of bones being the same in the hand of a man, wing of a bat, fin of the porpoise, and leg of the horse … at once explain themselves on the theory of descent with slow and slight successive modifications." </li> <li id="cite_note-196"><a href="#cite_ref-196">^</a> Darwin, C. R. Notebook C, CUL-DAR122.- Transcribed by Kees Rookmaaker. (Darwin Online), notes from de Beer, Gavin ed. 1960. Darwin's notebooks on transmutation of species. Part II. Second notebook [C] (February to July 1838). Bulletin of the British Museum (Natural History). Historical Series 2, No. 3 (May): pp. <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?pageseq=5&itemID=F1574b&viewtype=text">79</a> </li> <li id="cite_note-197"><a href="#cite_ref-197">^</a> <a href="#CITEREFDesmondMoore2009">Desmond & Moore 2009</a>, pp. 139–141, quotes "our acquiring the instinct one notion of beauty & negroes another" from Darwin, C. R. Notebook M : [Metaphysics on morals and speculations on expression (1838)]. CUL-DAR125.- Transcribed by Kees Rookmaaker, edited by Paul Barrett. 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Retrieved 25 October 2024.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=inters.org&rft.atitle=John+Henry+Newman+to+J.+Walker+of+Scarborough+on+Darwin%27s+Theory+of+Evolution+%7C+Inters.org&rft_id=https%3A%2F%2Finters.org%2FNewman-Scarborough-Darwin-Evolution&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> </li> <li id="cite_note-260"><a href="#cite_ref-260">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPius_XII1950" class="citation cs2">Pius XII (1950), <a rel="nofollow" class="external text" href="https://www.vatican.va/holy_father/pius_xii/encyclicals/documents/hf_p-xii_enc_12081950_humani-generis_en.html">Humani generis</a>, Vatican, retrieved 8 June 2009</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Humani+generis&rft.pub=Vatican&rft.date=1950&rft.au=Pius+XII&rft_id=https%3A%2F%2Fwww.vatican.va%2Fholy_father%2Fpius_xii%2Fencyclicals%2Fdocuments%2Fhf_p-xii_enc_12081950_humani-generis_en.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> </li> <li id="cite_note-261"><a href="#cite_ref-261">^</a> <a href="#CITEREFKreeft2001">Kreeft 2001</a>, p. 49 </li> <li id="cite_note-262"><a href="#cite_ref-262">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation cs2"><a rel="nofollow" class="external text" href="http://darwin-online.org.uk/biography.html">Biography</a>, <a href="/wiki/The_Complete_Works_of_Charles_Darwin_Online" title="The Complete Works of Charles Darwin Online">The Complete Works of Charles Darwin Online</a>, 21 January 2009, retrieved 23 April 2009</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Biography&rft.pub=The+Complete+Works+of+Charles+Darwin+Online&rft.date=2009-01-21&rft_id=http%3A%2F%2Fdarwin-online.org.uk%2Fbiography.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> </li> <li id="cite_note-263"><a href="#cite_ref-263">^</a> <a href="#CITEREFLarson2004">Larson 2004</a>, pp. 287–288, 295 </li> <li id="cite_note-264"><a href="#cite_ref-264">^</a> <a href="#CITEREFDarwinCosta2009">Darwin & Costa 2009</a>, p. x </li> <li id="cite_note-265"><a href="#cite_ref-265">^</a> <a href="#CITEREFPeckham1959">Peckham 1959</a>, recently reprinted. </li> <li id="cite_note-266"><a href="#cite_ref-266">^</a> <a href="#CITEREFQuammen2006">Quammen 2006</a>, pp. 179, 271–283 </li> <li id="cite_note-267"><a href="#cite_ref-267">^</a> The <a href="/wiki/International_Standard_Text_Code" title="International Standard Text Code">ISTC</a> of On the Origin of Species is A02-2009-00000001-4. As a tribute to its influence, this work has been the first one to be registered by The International ISTC Agency. </li> <li id="cite_note-268"><a href="#cite_ref-268">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation cs2"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20090228095055/http://www.darwin200.org/what-is.html">Darwin 200: Celebrating Charles Darwin's bicentenary – What is Darwin200?</a>, The <a href="/wiki/Natural_History_Museum,_London" title="Natural History Museum, London">Natural History Museum</a>, archived from <a rel="nofollow" class="external text" href="http://www.darwin200.org/what-is.html">the original</a> on 28 February 2009, retrieved 23 April 2009</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Darwin+200%3A+Celebrating+Charles+Darwin%27s+bicentenary+%E2%80%93+What+is+Darwin200%3F&rft.pub=The+Natural+History+Museum&rft_id=http%3A%2F%2Fwww.darwin200.org%2Fwhat-is.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> </li> <li id="cite_note-269"><a href="#cite_ref-269">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://www.livescience.com/52756-darwins-book-most-influential.html">"Darwin's 'Origin of Species' Voted Most Influential Academic Book"</a>. Tia Ghose. 11 November 2015. Retrieved 2 February 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Darwin%27s+%27Origin+of+Species%27+Voted+Most+Influential+Academic+Book&rft.pub=Tia+Ghose&rft.date=2015-11-11&rft_id=http%3A%2F%2Fwww.livescience.com%2F52756-darwins-book-most-influential.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> </li> <li id="cite_note-270"><a href="#cite_ref-270">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.theguardian.com/books/2015/nov/10/on-the-origin-of-species-voted-most-influential-academic-book-charles-darwin">"On the Origin of Species voted most influential academic book in history"</a>. Alison Flood. 10 November 2015. Retrieved 2 February 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=On+the+Origin+of+Species+voted+most+influential+academic+book+in+history&rft.pub=Alison+Flood&rft.date=2015-11-10&rft_id=https%3A%2F%2Fwww.theguardian.com%2Fbooks%2F2015%2Fnov%2F10%2Fon-the-origin-of-species-voted-most-influential-academic-book-charles-darwin&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> </li> </ol></div> <div class="mw-heading mw-heading3"><h3 id="Works_cited">Works cited</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=37" title="Edit section: Works cited">edit</a>]</div> <style data-mw-deduplicate="TemplateStyles:r1239549316">.mw-parser-output .refbegin{margin-bottom:0.5em}.mw-parser-output .refbegin-hanging-indents>ul{margin-left:0}.mw-parser-output .refbegin-hanging-indents>ul>li{margin-left:0;padding-left:3.2em;text-indent:-3.2em}.mw-parser-output .refbegin-hanging-indents ul,.mw-parser-output .refbegin-hanging-indents ul li{list-style:none}@media(max-width:720px){.mw-parser-output .refbegin-hanging-indents>ul>li{padding-left:1.6em;text-indent:-1.6em}}.mw-parser-output .refbegin-columns{margin-top:0.3em}.mw-parser-output .refbegin-columns ul{margin-top:0}.mw-parser-output .refbegin-columns li{page-break-inside:avoid;break-inside:avoid-column}@media screen{.mw-parser-output .refbegin{font-size:90%}}</style><div class="refbegin refbegin-columns references-column-width" style="column-width: 30em"> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBarlow1963" class="citation cs2"><a href="/wiki/Nora_Barlow" title="Nora Barlow">Barlow, Nora</a>, ed. (1963), <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=F1577&pageseq=1">"Darwin's Ornithological Notes"</a>, Bulletin of the British Museum (Natural History), Historical Series, vol. 2, no. 7, pp. 201–278, retrieved 10 June 2009</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Bulletin+of+the+British+Museum+%28Natural+History%29%2C+Historical+Series&rft.atitle=Darwin%27s+Ornithological+Notes&rft.volume=2&rft.issue=7&rft.pages=%3Cspan+class%3D%22nowrap%22%3E201-%3C%2Fspan%3E278&rft.date=1963&rft_id=http%3A%2F%2Fdarwin-online.org.uk%2Fcontent%2Fframeset%3Fviewtype%3Dtext%26itemID%3DF1577%26pageseq%3D1&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBowler1989" class="citation cs2"><a href="/wiki/Peter_J._Bowler" title="Peter J. Bowler">Bowler, Peter J.</a> (1989), "The Mendelian Revolution: The Emergence of Hereditarian Concepts in Modern Science and Society", The Yale Journal of Biology and Medicine, 63 (4), Baltimore: Johns Hopkins University Press: 335, <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/0-485-11375-9" title="Special:BookSources/0-485-11375-9"><bdi>0-485-11375-9</bdi></a>, <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2589185">2589185</a></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Yale+Journal+of+Biology+and+Medicine&rft.atitle=The+Mendelian+Revolution%3A+The+Emergence+of+Hereditarian+Concepts+in+Modern+Science+and+Society&rft.volume=63&rft.issue=4&rft.pages=335&rft.date=1989&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2589185%23id-name%3DPMC&rft.isbn=0-485-11375-9&rft.aulast=Bowler&rft.aufirst=Peter+J.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBowler1996" class="citation cs2">Bowler, Peter J (1996), Charles Darwin : the man and his influence, Cambridge University Press, <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/0-521-56668-1" title="Special:BookSources/0-521-56668-1"><bdi>0-521-56668-1</bdi></a></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Charles+Darwin+%3A+the+man+and+his+influence&rft.pub=Cambridge+University+Press&rft.date=1996&rft.isbn=0-521-56668-1&rft.aulast=Bowler&rft.aufirst=Peter+J&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBowler2003" class="citation cs2">Bowler, Peter J. (2003), <a rel="nofollow" class="external text" href="https://archive.org/details/evolutionhistory0000bowl_n7y8">Evolution: The History of an Idea</a> (3rd ed.), University of California Press, <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/0-520-23693-9" title="Special:BookSources/0-520-23693-9"><bdi>0-520-23693-9</bdi></a></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Evolution%3A+The+History+of+an+Idea&rft.edition=3rd&rft.pub=University+of+California+Press&rft.date=2003&rft.isbn=0-520-23693-9&rft.aulast=Bowler&rft.aufirst=Peter+J.&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fevolutionhistory0000bowl_n7y8&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBowler2013" class="citation cs2">Bowler, Peter J. (2013), Darwin Deleted: Imagining a World without Darwin, The University of Chicago Press, <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-226-00984-1" title="Special:BookSources/978-0-226-00984-1"><bdi>978-0-226-00984-1</bdi></a></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Darwin+Deleted%3A+Imagining+a+World+without+Darwin&rft.pub=The+University+of+Chicago+Press&rft.date=2013&rft.isbn=978-0-226-00984-1&rft.aulast=Bowler&rft.aufirst=Peter+J.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBrowne1995" class="citation cs2"><a href="/wiki/Janet_Browne" title="Janet Browne">Browne, E. Janet</a> (1995), Charles Darwin: Vol. 1 Voyaging, London: Jonathan Cape, <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/1-84413-314-1" title="Special:BookSources/1-84413-314-1"><bdi>1-84413-314-1</bdi></a></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Charles+Darwin%3A+Vol.+1+Voyaging&rft.place=London&rft.pub=Jonathan+Cape&rft.date=1995&rft.isbn=1-84413-314-1&rft.aulast=Browne&rft.aufirst=E.+Janet&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBrowne2002" class="citation cs2">Browne, E. 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(1977), <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/EditorialIntroductions/Freeman_OntheOriginofSpecies.html">"On the Origin of Species"</a>, The Works of Charles Darwin: An Annotated Bibliographical Handlist (2nd ed.), Folkestone, England: Dawson, <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/0-7129-0740-8" title="Special:BookSources/0-7129-0740-8"><bdi>0-7129-0740-8</bdi></a></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=On+the+Origin+of+Species&rft.btitle=The+Works+of+Charles+Darwin%3A+An+Annotated+Bibliographical+Handlist&rft.place=Folkestone%2C+England&rft.edition=2nd&rft.pub=Dawson&rft.date=1977&rft.isbn=0-7129-0740-8&rft.aulast=Freeman&rft.aufirst=Richard+B.&rft_id=http%3A%2F%2Fdarwin-online.org.uk%2FEditorialIntroductions%2FFreeman_OntheOriginofSpecies.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHerbert1980" class="citation cs2">Herbert, Sandra, ed. (1980), <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?viewtype=side&itemID=F1583e&pageseq=1">"The Red Notebook of Charles Darwin"</a>, Bulletin of the British Museum (Natural History), Historical Series, 7: 1–164, <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.5962%2Fp.272299">10.5962/p.272299</a></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Bulletin+of+the+British+Museum+%28Natural+History%29%2C+Historical+Series&rft.atitle=The+Red+Notebook+of+Charles+Darwin&rft.volume=7&rft.pages=%3Cspan+class%3D%22nowrap%22%3E1-%3C%2Fspan%3E164&rft.date=1980&rft_id=info%3Adoi%2F10.5962%2Fp.272299&rft_id=http%3A%2F%2Fdarwin-online.org.uk%2Fcontent%2Fframeset%3Fviewtype%3Dside%26itemID%3DF1583e%26pageseq%3D1&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> Also available here <a rel="nofollow" class="external autonumber" href="https://archive.org/details/bulletinofbritis07brit">[1]</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHodge1874" class="citation cs2"><a href="/wiki/Charles_Hodge" title="Charles Hodge">Hodge, Charles</a> (1874), <a rel="nofollow" class="external text" href="http://www.gutenberg.org/files/19192/19192-8.txt">What is Darwinism?</a>, Scribner Armstrong, retrieved 14 January 2007</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=What+is+Darwinism%3F&rft.pub=Scribner+Armstrong&rft.date=1874&rft.aulast=Hodge&rft.aufirst=Charles&rft_id=http%3A%2F%2Fwww.gutenberg.org%2Ffiles%2F19192%2F19192-8.txt&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHuxleyKettlewell1965" class="citation book cs1"><a href="/wiki/Julian_Huxley" title="Julian Huxley">Huxley, Julian</a>; <a href="/wiki/Bernard_Kettlewell" title="Bernard Kettlewell">Kettlewell, H.B.D.</a> (1965). Charles Darwin and His World. New York: the Viking Press.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Charles+Darwin+and+His+World&rft.place=New+York&rft.pub=the+Viking+Press&rft.date=1965&rft.aulast=Huxley&rft.aufirst=Julian&rft.au=Kettlewell%2C+H.B.D.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHuxley1860" class="citation cs2">Huxley, Thomas Henry (1860), <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?itemID=A32&viewtype=text&pageseq=1">"Darwin on the Origin of Species"</a>, <a href="/wiki/Westminster_Review" class="mw-redirect" title="Westminster Review">Westminster Review</a>, 17 (April 1860): 541–570</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Westminster+Review&rft.atitle=Darwin+on+the+Origin+of+Species&rft.volume=17&rft.issue=April+1860&rft.pages=%3Cspan+class%3D%22nowrap%22%3E541-%3C%2Fspan%3E570&rft.date=1860&rft.aulast=Huxley&rft.aufirst=Thomas+Henry&rft_id=http%3A%2F%2Fdarwin-online.org.uk%2Fcontent%2Fframeset%3FitemID%3DA32%26viewtype%3Dtext%26pageseq%3D1&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988">. Published anonymously.</li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHuxley1863" class="citation cs2"><a href="/wiki/Thomas_Huxley" class="mw-redirect" title="Thomas Huxley">Huxley, Thomas</a> (1863), <a rel="nofollow" class="external text" href="http://aleph0.clarku.edu/huxley/CE2/Phen.html">Six Lectures to Working Men "On Our Knowledge of the Causes of the Phenomena of Organic Nature" (Republished in Volume II of his Collected Essays, Darwiniana)</a>, retrieved 15 December 2006</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Six+Lectures+to+Working+Men+%22On+Our+Knowledge+of+the+Causes+of+the+Phenomena+of+Organic+Nature%22+%28Republished+in+Volume+II+of+his+Collected+Essays%2C+Darwiniana%29&rft.date=1863&rft.aulast=Huxley&rft.aufirst=Thomas&rft_id=http%3A%2F%2Faleph0.clarku.edu%2Fhuxley%2FCE2%2FPhen.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKeynes2000" class="citation cs2">Keynes, Richard, ed. (2000), Charles Darwin's Zoology Notes & Specimen Lists from HMS Beagle, Cambridge University Press, <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/0-521-67350-X" title="Special:BookSources/0-521-67350-X"><bdi>0-521-67350-X</bdi></a></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Charles+Darwin%27s+Zoology+Notes+%26+Specimen+Lists+from+HMS+Beagle&rft.pub=Cambridge+University+Press&rft.date=2000&rft.isbn=0-521-67350-X&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKragh2016" class="citation cs2"><a href="/wiki/Helge_Kragh" title="Helge Kragh">Kragh, Helge</a> (2016), <a rel="nofollow" class="external text" href="https://link.springer.com/article/10.1140/epjh/e2016-70045-7">"The source of solar energy, ca. 1840–1910: From meteoric hypothesis to radioactive speculations"</a>, The European Physical Journal H, 41 (4–5), Springer Nature: 365–394, <a href="/wiki/ArXiv_(identifier)" class="mw-redirect" title="ArXiv (identifier)">arXiv</a>:<a rel="nofollow" class="external text" href="https://arxiv.org/abs/1609.02834">1609.02834</a>, <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2016EPJH...41..365K">2016EPJH...41..365K</a>, <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1140%2Fepjh%2Fe2016-70045-7">10.1140/epjh/e2016-70045-7</a></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+European+Physical+Journal+H&rft.atitle=The+source+of+solar+energy%2C+ca.+1840%E2%80%931910%3A+From+meteoric+hypothesis+to+radioactive+speculations&rft.volume=41&rft.issue=%3Cspan+class%3D%22nowrap%22%3E4%E2%80%93%3C%2Fspan%3E5&rft.pages=%3Cspan+class%3D%22nowrap%22%3E365-%3C%2Fspan%3E394&rft.date=2016&rft_id=info%3Aarxiv%2F1609.02834&rft_id=info%3Adoi%2F10.1140%2Fepjh%2Fe2016-70045-7&rft_id=info%3Abibcode%2F2016EPJH...41..365K&rft.aulast=Kragh&rft.aufirst=Helge&rft_id=https%3A%2F%2Flink.springer.com%2Farticle%2F10.1140%2Fepjh%2Fe2016-70045-7&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKreeft2001" class="citation cs2"><a href="/wiki/Peter_Kreeft" title="Peter Kreeft">Kreeft, Peter</a> (2001), <a rel="nofollow" class="external text" href="https://archive.org/details/catholicchristia00kree">Catholic Christianity</a>, San Francisco: Ignatius Press, <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/0-89870-798-6" title="Special:BookSources/0-89870-798-6"><bdi>0-89870-798-6</bdi></a></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Catholic+Christianity&rft.place=San+Francisco&rft.pub=Ignatius+Press&rft.date=2001&rft.isbn=0-89870-798-6&rft.aulast=Kreeft&rft.aufirst=Peter&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fcatholicchristia00kree&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLarson2004" class="citation cs2"><a href="/wiki/Edward_Larson" class="mw-redirect" title="Edward Larson">Larson, Edward J.</a> (2004), Evolution: The Remarkable History of a Scientific Theory, New York: Modern Library, <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/0-8129-6849-2" title="Special:BookSources/0-8129-6849-2"><bdi>0-8129-6849-2</bdi></a></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Evolution%3A+The+Remarkable+History+of+a+Scientific+Theory&rft.place=New+York&rft.pub=Modern+Library&rft.date=2004&rft.isbn=0-8129-6849-2&rft.aulast=Larson&rft.aufirst=Edward+J.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLeifchild1859" class="citation cs2">Leifchild (19 November 1859), <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?viewtype=image&itemID=CUL-DAR226.1.8&pageseq=1">"Review of 'Origin'"</a>, <a href="/wiki/Athenaeum_(British_magazine)" class="mw-redirect" title="Athenaeum (British magazine)">Athenaeum</a>, no. 1673, retrieved 22 November 2008</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Athenaeum&rft.atitle=Review+of+%27Origin%27&rft.issue=1673&rft.date=1859-11-19&rft.au=Leifchild&rft_id=http%3A%2F%2Fdarwin-online.org.uk%2Fcontent%2Fframeset%3Fviewtype%3Dimage%26itemID%3DCUL-DAR226.1.8%26pageseq%3D1&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLucas1979" class="citation cs2"><a href="/wiki/John_Lucas_(philosopher)" title="John Lucas (philosopher)">Lucas, John R.</a> (1979), <a rel="nofollow" class="external text" href="https://web.archive.org/web/20200627072321/http://users.ox.ac.uk/~jrlucas/legend.html">"Wilberforce and Huxley: A Legendary Encounter"</a>, The Historical Journal, vol. 22, no. 2, pp. 313–330, <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1017%2FS0018246X00016848">10.1017/S0018246X00016848</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/11617072">11617072</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:19198585">19198585</a>, archived from <a rel="nofollow" class="external text" href="http://users.ox.ac.uk/~jrlucas/legend.html">the original</a> on 27 June 2020, retrieved 22 November 2008</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Historical+Journal&rft.atitle=Wilberforce+and+Huxley%3A+A+Legendary+Encounter&rft.volume=22&rft.issue=2&rft.pages=%3Cspan+class%3D%22nowrap%22%3E313-%3C%2Fspan%3E330&rft.date=1979&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A19198585%23id-name%3DS2CID&rft_id=info%3Apmid%2F11617072&rft_id=info%3Adoi%2F10.1017%2FS0018246X00016848&rft.aulast=Lucas&rft.aufirst=John+R.&rft_id=http%3A%2F%2Fusers.ox.ac.uk%2F~jrlucas%2Flegend.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMayr1982" class="citation cs2"><a href="/wiki/Ernst_Mayr" title="Ernst Mayr">Mayr, Ernst</a> (1982), <a href="/wiki/The_Growth_of_Biological_Thought" title="The Growth of Biological Thought">The Growth of Biological Thought</a>, Harvard University Press, <a 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href="http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=A544&pageseq=1">"Mind the gap: Did Darwin Avoid Publishing his Theory for Many Years?"</a>, Notes and Records of the Royal Society, 61 (2): 177–205, <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1098%2Frsnr.2006.0171">10.1098/rsnr.2006.0171</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:202574857">202574857</a>, retrieved 5 January 2009</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Notes+and+Records+of+the+Royal+Society&rft.atitle=Mind+the+gap%3A+Did+Darwin+Avoid+Publishing+his+Theory+for+Many+Years%3F&rft.volume=61&rft.issue=2&rft.pages=%3Cspan+class%3D%22nowrap%22%3E177-%3C%2Fspan%3E205&rft.date=2007&rft_id=info%3Adoi%2F10.1098%2Frsnr.2006.0171&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A202574857%23id-name%3DS2CID&rft.aulast=van+Wyhe&rft.aufirst=John&rft_id=http%3A%2F%2Fdarwin-online.org.uk%2Fcontent%2Fframeset%3Fviewtype%3Dtext%26itemID%3DA544%26pageseq%3D1&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFvan_Wyhe2008" class="citation cs2">van Wyhe, John (2008), Darwin: The Story of the Man and His Theories of Evolution, London: Andre Deutsch, <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-233-00251-4" title="Special:BookSources/978-0-233-00251-4"><bdi>978-0-233-00251-4</bdi></a></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Darwin%3A+The+Story+of+the+Man+and+His+Theories+of+Evolution&rft.place=London&rft.pub=Andre+Deutsch&rft.date=2008&rft.isbn=978-0-233-00251-4&rft.aulast=van+Wyhe&rft.aufirst=John&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFvan_Wyhe2009" class="citation cs2">van Wyhe, John (2009), <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/darwin.html">Charles Darwin: Gentleman Naturalist: A Biographical Sketch</a>, <a href="/wiki/The_Complete_Works_of_Charles_Darwin_Online" title="The Complete Works of Charles Darwin Online">The Complete Works of Charles Darwin Online</a>, retrieved 6 June 2009</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Charles+Darwin%3A+Gentleman+Naturalist%3A+A+Biographical+Sketch&rft.pub=The+Complete+Works+of+Charles+Darwin+Online&rft.date=2009&rft.aulast=van+Wyhe&rft.aufirst=John&rft_id=http%3A%2F%2Fdarwin-online.org.uk%2Fdarwin.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"></li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=38" title="Edit section: Further reading">edit</a>]</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><div class="side-box metadata side-box-right"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1126788409"> <div class="side-box-abovebelow"> <a href="/wiki/Wikipedia:The_Wikipedia_Library" title="Wikipedia:The Wikipedia Library">Library resources</a> about On the origin of species <hr /></div> <div class="side-box-flex"> <div class="side-box-text plainlist"><ul><li><a class="external text" href="https://ftl.toolforge.org/cgi-bin/ftl?st=&su=Darwin%2C+Charles%2C+1809%E2%80%931882.+%27%27On+the+origin+of+species%27%27&library=OLBP">Online books</a></li> <li><a class="external text" href="https://ftl.toolforge.org/cgi-bin/ftl?st=&su=Darwin%2C+Charles%2C+1809%E2%80%931882.+%27%27On+the+origin+of+species%27%27">Resources in your library</a></li> <li><a class="external text" href="https://ftl.toolforge.org/cgi-bin/ftl?st=&su=Darwin%2C+Charles%2C+1809%E2%80%931882.+%27%27On+the+origin+of+species%27%27&library=0CHOOSE0">Resources in other libraries</a></li> </ul></div></div> </div> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBrowne2007" class="citation cs2"><a href="/wiki/Janet_Browne" title="Janet Browne">Browne, Janet</a> (2007), <a rel="nofollow" class="external text" href="https://archive.org/details/darwinsoriginofs00brow">Darwin's Origin of Species: A Biography</a>, Grove Press, <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-87113-953-5" title="Special:BookSources/978-0-87113-953-5"><bdi>978-0-87113-953-5</bdi></a></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Darwin%27s+Origin+of+Species%3A+A+Biography&rft.pub=Grove+Press&rft.date=2007&rft.isbn=978-0-87113-953-5&rft.aulast=Browne&rft.aufirst=Janet&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fdarwinsoriginofs00brow&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation cs2"><a href="/wiki/Thomas_Malthus" class="mw-redirect" title="Thomas Malthus">Malthus, Thomas Robert</a> (1826), <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?pageseq=1&itemID=A545.1&viewtype=text">An Essay on the Principle of Population: A View of its Past and Present Effects on Human Happiness; with an Inquiry into Our Prospects Respecting the Future Removal or Mitigation of the Evils which It Occasions</a>, vol. 1 (6th ed.), London: John Murray, retrieved 13 November 2017</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=An+Essay+on+the+Principle+of+Population%3A+A+View+of+its+Past+and+Present+Effects+on+Human+Happiness%3B+with+an+Inquiry+into+Our+Prospects+Respecting+the+Future+Removal+or+Mitigation+of+the+Evils+which+It+Occasions&rft.place=London&rft.edition=6th&rft.pub=John+Murray&rft.date=1826&rft.aulast=Malthus&rft.aufirst=Thomas+Robert&rft_id=http%3A%2F%2Fdarwin-online.org.uk%2Fcontent%2Fframeset%3Fpageseq%3D1%26itemID%3DA545.1%26viewtype%3Dtext&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"> (<a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?pageseq=1&itemID=A545.2&viewtype=text">Vol. 2</a>)</li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation cs2">Reznick, David N. (2009), The Origin Then and Now: An Interpretive Guide to the Origin of Species, Princeton University Press, <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-691-12978-5" title="Special:BookSources/978-0-691-12978-5"><bdi>978-0-691-12978-5</bdi></a></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Origin+Then+and+Now%3A+An+Interpretive+Guide+to+the+Origin+of+Species&rft.pub=Princeton+University+Press&rft.date=2009&rft.isbn=978-0-691-12978-5&rft.aulast=Reznick&rft.aufirst=David+N.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation cs2"><a href="/wiki/J._William_Schopf" title="J. William Schopf">Schopf, J. William</a>; Scheibel, Arnold B. (1997), The Origin and Evolution of Intelligence, Boston: Jones and Bartlett, <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/0-7637-0365-6" title="Special:BookSources/0-7637-0365-6"><bdi>0-7637-0365-6</bdi></a></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Origin+and+Evolution+of+Intelligence&rft.place=Boston&rft.pub=Jones+and+Bartlett&rft.date=1997&rft.isbn=0-7637-0365-6&rft.aulast=Schopf&rft.aufirst=J.+William&rft.au=Scheibel%2C+Arnold+B.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation cs2">van Hoorn, Marijn (2009), <a rel="nofollow" class="external text" href="https://web.archive.org/web/20111202091528/http://api.ning.com/files/aT0Fpi7QEAJXZty6RRr0*ayjT5aN-CLIR1HAnom7DRBt*Z64Ko2B*I88LPx*h2otwLvz5Vp*IKVqHQunTD*sItKN3P7MUMve/TeylerWinklerDarwin.doc">Teyler, Winkler, Darwin (Lecture given at the Congress of the European Botanical and Horticultural Libraries Group, Prague, 23 April 2009)</a>, Teyler Net (Weblog of the Teylers Museum, Haarlem), archived from <a rel="nofollow" class="external text" href="http://api.ning.com/files/aT0Fpi7QEAJXZty6RRr0*ayjT5aN-CLIR1HAnom7DRBt*Z64Ko2B*I88LPx*h2otwLvz5Vp*IKVqHQunTD*sItKN3P7MUMve/TeylerWinklerDarwin.doc">the original</a> on 2 December 2011, retrieved 27 April 2010</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Teyler%2C+Winkler%2C+Darwin+%28Lecture+given+at+the+Congress+of+the+European+Botanical+and+Horticultural+Libraries+Group%2C+Prague%2C+23+April+2009%29&rft.pub=Teyler+Net+%28Weblog+of+the+Teylers+Museum%2C+Haarlem%29&rft.date=2009&rft.aulast=van+Hoorn&rft.aufirst=Marijn&rft_id=http%3A%2F%2Fapi.ning.com%2Ffiles%2FaT0Fpi7QEAJXZty6RRr0%2AayjT5aN-CLIR1HAnom7DRBt%2AZ64Ko2B%2AI88LPx%2Ah2otwLvz5Vp%2AIKVqHQunTD%2AsItKN3P7MUMve%2FTeylerWinklerDarwin.doc&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPechenik2023" class="citation cs2">Pechenik, Jan A. (2023), <a rel="nofollow" class="external text" href="https://books.google.com/books?id=xAamEAAAQBAJ">The Readable Darwin: The Origin of Species Edited for Modern Readers</a> (2 ed.), Oxford University Press, <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-19-757526-0" title="Special:BookSources/978-0-19-757526-0"><bdi>978-0-19-757526-0</bdi></a></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Readable+Darwin%3A+The+Origin+of+Species+Edited+for+Modern+Readers&rft.edition=2&rft.pub=Oxford+University+Press&rft.date=2023&rft.isbn=978-0-19-757526-0&rft.aulast=Pechenik&rft.aufirst=Jan+A.&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DxAamEAAAQBAJ&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"></li></ul> <div class="mw-heading mw-heading3"><h3 id="Contemporary_reviews">Contemporary reviews</h3>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=39" title="Edit section: Contemporary reviews">edit</a>]</div> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation cs2"><a href="/wiki/William_Benjamin_Carpenter" title="William Benjamin Carpenter">Carpenter, William Benjamin</a> (1859), <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?itemID=A17&viewtype=text&pageseq=1">"Darwin on the Origin of Species"</a>, <a href="/wiki/National_Review_(1855)" title="National Review (1855)">National Review</a>, vol. 10, no. December 1859, pp. 188–214</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=National+Review&rft.atitle=Darwin+on+the+Origin+of+Species&rft.volume=10&rft.issue=December+1859&rft.pages=%3Cspan+class%3D%22nowrap%22%3E188-%3C%2Fspan%3E214&rft.date=1859&rft.aulast=Carpenter&rft.aufirst=William+Benjamin&rft_id=http%3A%2F%2Fdarwin-online.org.uk%2Fcontent%2Fframeset%3FitemID%3DA17%26viewtype%3Dtext%26pageseq%3D1&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988">. Published anonymously.</li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation cs2"><a href="/wiki/Asa_Gray" title="Asa Gray">Gray, Asa</a> (1860), <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?itemID=A213&viewtype=text&pageseq=1">"(Review of) The Origin of Species"</a>, <a href="/wiki/Athenaeum_(British_magazine)" class="mw-redirect" title="Athenaeum (British magazine)">Athenaeum</a> (1710: 4 August 1860): 161</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Athenaeum&rft.atitle=%28Review+of%29+The+Origin+of+Species&rft.issue=1710%3A+4+August+1860&rft.pages=161&rft.date=1860&rft.aulast=Gray&rft.aufirst=Asa&rft_id=http%3A%2F%2Fdarwin-online.org.uk%2Fcontent%2Fframeset%3FitemID%3DA213%26viewtype%3Dtext%26pageseq%3D1&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988">. Extract from Proceedings of the American Academy of Arts and Sciences 4 (1860): 411–415.</li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation cs2"><a href="/wiki/Thomas_Henry_Huxley" title="Thomas Henry Huxley">Huxley, Thomas Henry</a> (1859), <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?itemID=A43&viewtype=image&pageseq=1">"Time and Life: Mr Darwin's Origin of Species"</a>, Macmillan's Magazine, 1: 142–148</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Macmillan%27s+Magazine&rft.atitle=Time+and+Life%3A+Mr+Darwin%27s+Origin+of+Species&rft.volume=1&rft.pages=%3Cspan+class%3D%22nowrap%22%3E142-%3C%2Fspan%3E148&rft.date=1859&rft.aulast=Huxley&rft.aufirst=Thomas+Henry&rft_id=http%3A%2F%2Fdarwin-online.org.uk%2Fcontent%2Fframeset%3FitemID%3DA43%26viewtype%3Dimage%26pageseq%3D1&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988">.</li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation cs2">Huxley, Thomas Henry (1859), <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?itemID=A166&viewtype=text&pageseq=1">"Darwin on the Origin of Species"</a>, <a href="/wiki/The_Times" title="The Times">The Times</a> (26 December 1859): 8–9</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Times&rft.atitle=Darwin+on+the+Origin+of+Species&rft.issue=26+December+1859&rft.pages=%3Cspan+class%3D%22nowrap%22%3E8-%3C%2Fspan%3E9&rft.date=1859&rft.aulast=Huxley&rft.aufirst=Thomas+Henry&rft_id=http%3A%2F%2Fdarwin-online.org.uk%2Fcontent%2Fframeset%3FitemID%3DA166%26viewtype%3Dtext%26pageseq%3D1&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988">. Published anonymously.</li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation cs2"><a href="/wiki/Fleeming_Jenkin" title="Fleeming Jenkin">Jenkin, Fleeming</a> (1867), <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?itemID=A24&viewtype=text&pageseq=1">"(Review of) The Origin of Species"</a>, North British Review, 46 (June 1867): 277–318</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=North+British+Review&rft.atitle=%28Review+of%29+The+Origin+of+Species&rft.volume=46&rft.issue=June+1867&rft.pages=%3Cspan+class%3D%22nowrap%22%3E277-%3C%2Fspan%3E318&rft.date=1867&rft.aulast=Jenkin&rft.aufirst=Fleeming&rft_id=http%3A%2F%2Fdarwin-online.org.uk%2Fcontent%2Fframeset%3FitemID%3DA24%26viewtype%3Dtext%26pageseq%3D1&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988">. Published anonymously.</li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation cs2"><a href="/wiki/Andrew_Murray_(naturalist)" title="Andrew Murray (naturalist)">Murray, Andrew</a> (1860), <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?itemID=A14&viewtype=text&pageseq=1">"On Mr Darwin's Theory of the Origin of Species"</a>, Proceedings of the Royal Society of Edinburgh, 4: 274–291, <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1017%2FS0370164600034246">10.1017/S0370164600034246</a></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proceedings+of+the+Royal+Society+of+Edinburgh&rft.atitle=On+Mr+Darwin%27s+Theory+of+the+Origin+of+Species&rft.volume=4&rft.pages=%3Cspan+class%3D%22nowrap%22%3E274-%3C%2Fspan%3E291&rft.date=1860&rft_id=info%3Adoi%2F10.1017%2FS0370164600034246&rft.aulast=Murray&rft.aufirst=Andrew&rft_id=http%3A%2F%2Fdarwin-online.org.uk%2Fcontent%2Fframeset%3FitemID%3DA14%26viewtype%3Dtext%26pageseq%3D1&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988">.</li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation cs2"><a href="/wiki/Richard_Owen" title="Richard Owen">Owen, Richard</a> (1860), <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?itemID=A30&viewtype=text&pageseq=1">"Review of Darwin's Origin of Species"</a>, <a href="/wiki/Edinburgh_Review" title="Edinburgh Review">Edinburgh Review</a>, 3 (April 1860): 487–532</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Edinburgh+Review&rft.atitle=Review+of+Darwin%27s+Origin+of+Species&rft.volume=3&rft.issue=April+1860&rft.pages=%3Cspan+class%3D%22nowrap%22%3E487-%3C%2Fspan%3E532&rft.date=1860&rft.aulast=Owen&rft.aufirst=Richard&rft_id=http%3A%2F%2Fdarwin-online.org.uk%2Fcontent%2Fframeset%3FitemID%3DA30%26viewtype%3Dtext%26pageseq%3D1&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988">. Published anonymously.</li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation cs2"><a href="/wiki/Samuel_Wilberforce" title="Samuel Wilberforce">Wilberforce, Samuel</a> (1860), <a rel="nofollow" class="external text" href="http://darwin-online.org.uk/content/frameset?itemID=A19&viewtype=text&pageseq=1">"(Review of) On the Origin of Species, by means of Natural Selection; or the Preservation of Favoured Races in the Struggle for Life"</a>, <a href="/wiki/Quarterly_Review" title="Quarterly Review">Quarterly Review</a>, 108 (215: July 1860): 225–264</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Quarterly+Review&rft.atitle=%28Review+of%29+On+the+Origin+of+Species%2C+by+means+of+Natural+Selection%3B+or+the+Preservation+of+Favoured+Races+in+the+Struggle+for+Life&rft.volume=108&rft.issue=215%3A+July+1860&rft.pages=%3Cspan+class%3D%22nowrap%22%3E225-%3C%2Fspan%3E264&rft.date=1860&rft.aulast=Wilberforce&rft.aufirst=Samuel&rft_id=http%3A%2F%2Fdarwin-online.org.uk%2Fcontent%2Fframeset%3FitemID%3DA19%26viewtype%3Dtext%26pageseq%3D1&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988">. Published anonymously.</li></ul> <dl><dd><dl><dd>For further reviews, see <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation cs2"><a rel="nofollow" class="external text" href="http://darwin-online.org.uk/reviews.html">Darwin Online: Reviews & Responses to Darwin</a>, Darwin Online, 10 March 2009, retrieved 18 June 2009</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Darwin+Online%3A+Reviews+%26+Responses+to+Darwin&rft.pub=Darwin+Online&rft.date=2009-03-10&rft_id=http%3A%2F%2Fdarwin-online.org.uk%2Freviews.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3AOn+the+Origin+of+Species" class="Z3988"></dd></dl></dd></dl> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2>[<a href="/w/index.php?title=On_the_Origin_of_Species&action=edit&section=40" title="Edit section: External links">edit</a>]</div> <link rel="mw-deduplicated-inline-style" 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Polish, Russian and Spanish</li> <li><a rel="nofollow" class="external text" href="http://darwin-online.org.uk/Variorum/index.html">Online Variorum</a>, showing every change between the six British editions</li></ul></li> <li><a rel="nofollow" class="external text" href="https://standardebooks.org/ebooks/charles-darwin/the-origin-of-species">On the Origin of Species</a> at <a href="/wiki/Standard_Ebooks" title="Standard Ebooks">Standard Ebooks</a></li> <li><a rel="nofollow" class="external text" href="https://www.gutenberg.org/ebooks/1228">On the Origin of Species</a> eBook provided by <a href="/wiki/Project_Gutenberg" title="Project Gutenberg">Project Gutenberg</a></li> <li><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/2/21/Speaker_Icon.svg/15px-Speaker_Icon.svg.png" decoding="async" width="15" height="15" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/21/Speaker_Icon.svg/23px-Speaker_Icon.svg.png 1.5x, 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Appleton and Company, New York, with front insert by H. E. Barker, Lincolniana</li> <li><a rel="nofollow" class="external text" href="http://cudl.lib.cam.ac.uk/collections/darwin_mss">Darwin's notes on the creation of On the Origin of Species</a> digitised in <a href="/wiki/Cambridge_Digital_Library" title="Cambridge Digital Library">Cambridge Digital Library</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 .navbox-group,.mw-parser-output .navbox-title,.mw-parser-output .navbox-abovebelow{padding:0.25em 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.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="Charles_Darwin69" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Charles_Darwin" title="Template:Charles Darwin"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Charles_Darwin" title="Template talk:Charles Darwin"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Charles_Darwin" title="Special:EditPage/Template:Charles Darwin"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Charles_Darwin69" style="font-size:114%;margin:0 4em"><a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Life</th><td class="navbox-list-with-group navbox-list navbox-odd wraplinks" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Darwin%E2%80%93Wedgwood_family" title="Darwin–Wedgwood family">Darwin–Wedgwood family</a> <ul><li><a href="/wiki/Erasmus_Darwin" title="Erasmus Darwin">Erasmus Darwin</a></li> <li><a href="/wiki/Josiah_Wedgwood" title="Josiah Wedgwood">Josiah Wedgwood</a></li></ul></li> <li><a href="/wiki/Charles_Darwin%27s_education" title="Charles Darwin's education">Education</a></li> <li><a href="/wiki/Second_voyage_of_HMS_Beagle" title="Second voyage of HMS Beagle">Voyage on HMS Beagle</a></li> <li><a href="/wiki/Inception_of_Darwin%27s_theory" title="Inception of Darwin's theory">Inception of theory</a></li> <li><a href="/wiki/Development_of_Darwin%27s_theory" title="Development of Darwin's theory">Development of theory</a></li> <li><a href="/wiki/Publication_of_Darwin%27s_theory" title="Publication of Darwin's theory">Publication of theory</a></li> <li><a href="/wiki/Reactions_to_On_the_Origin_of_Species" title="Reactions to On the Origin of Species">Reactions to On the Origin of Species</a></li> <li><a href="/wiki/Darwin_from_Orchids_to_Variation" title="Darwin from Orchids to Variation">Orchids to Variation</a></li> <li><a href="/wiki/Darwin_from_Descent_of_Man_to_Emotions" title="Darwin from Descent of Man to Emotions">Descent of Man to Emotions</a></li> <li><a href="/wiki/Darwin_from_Insectivorous_Plants_to_Worms" title="Darwin from Insectivorous Plants to Worms">Insectivorous Plants to Worms</a></li> <li><a href="/wiki/Religious_views_of_Charles_Darwin" title="Religious views of Charles Darwin">Religious views</a></li> <li><a href="/wiki/Health_of_Charles_Darwin" title="Health of Charles Darwin">Health</a></li> <li><a href="/wiki/Darwin_and_women" title="Darwin and women">Women</a></li> <li><a href="/wiki/Portraits_of_Charles_Darwin" title="Portraits of Charles Darwin">Portraits of Darwin</a> (<a href="/wiki/Caricatures_of_Charles_Darwin_and_his_evolutionary_theory_in_19th-century_England" title="Caricatures of Charles Darwin and his evolutionary theory in 19th-century England">caricatures</a>)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Charles_Darwin_bibliography" title="Charles Darwin bibliography">Writings</a></th><td class="navbox-list-with-group navbox-list navbox-even wraplinks" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Extracts_from_Letters_to_Henslow" title="Extracts from Letters to Henslow">Extracts from Letters to Henslow</a> (1835)</li> <li><a href="/wiki/The_Voyage_of_the_Beagle" title="The Voyage of the Beagle">The Voyage of the Beagle</a> (1839)</li> <li><a href="/wiki/The_Structure_and_Distribution_of_Coral_Reefs" title="The Structure and Distribution of Coral Reefs">The Structure and Distribution of Coral Reefs</a> (1842)</li> <li><a href="/wiki/Zoology_of_the_Voyage_of_H.M.S._Beagle" title="Zoology of the Voyage of H.M.S. Beagle">Zoology of the Voyage of H.M.S. Beagle</a> (1838–1843)</li> <li><a href="/wiki/Geological_Observations_on_the_Volcanic_Islands" title="Geological Observations on the Volcanic Islands">Geological Observations on the Volcanic Islands</a> (1844)</li> <li><a href="/wiki/Geological_Observations_on_South_America" title="Geological Observations on South America">Geological Observations on South America</a> (1846)</li> <li>"<a href="/wiki/On_the_Tendency_of_Species_to_form_Varieties;_and_on_the_Perpetuation_of_Varieties_and_Species_by_Natural_Means_of_Selection" title="On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection">On the Tendency of Species to form Varieties</a>" (1858)</li> <li><a class="mw-selflink selflink">On the Origin of Species</a> (1859)</li> <li><a href="/wiki/Fertilisation_of_Orchids" title="Fertilisation of Orchids">Fertilisation of Orchids</a> (1862)</li> <li><a href="/wiki/The_Variation_of_Animals_and_Plants_Under_Domestication" title="The Variation of Animals and Plants Under Domestication">The Variation of Animals and Plants Under Domestication</a> (1868) <ul><li><a href="/wiki/Natural_Selection_(manuscript)" title="Natural Selection (manuscript)">Natural Selection</a></li></ul></li> <li><a href="/wiki/The_Descent_of_Man,_and_Selection_in_Relation_to_Sex" title="The Descent of Man, and Selection in Relation to Sex">The Descent of Man, and Selection in Relation to Sex</a> (1871)</li> <li><a href="/wiki/The_Expression_of_the_Emotions_in_Man_and_Animals" title="The Expression of the Emotions in Man and Animals">The Expression of the Emotions in Man and Animals</a> (1872)</li> <li><a href="/wiki/Insectivorous_Plants_(book)" class="mw-redirect" title="Insectivorous Plants (book)">Insectivorous Plants</a> (1875)</li> <li><a href="/wiki/On_the_Movements_and_Habits_of_Climbing_Plants" title="On the Movements and Habits of Climbing Plants">On the Movements and Habits of Climbing Plants</a> (1875)</li> <li><a href="/wiki/The_Effects_of_Cross_and_Self_Fertilisation_in_the_Vegetable_Kingdom" title="The Effects of Cross and Self Fertilisation in the Vegetable Kingdom">The Effects of Cross and Self Fertilisation in the Vegetable Kingdom</a> (1876)</li> <li><a href="/wiki/The_Different_Forms_of_Flowers_on_Plants_of_the_Same_Species" title="The Different Forms of Flowers on Plants of the Same Species">The Different Forms of Flowers on Plants of the Same Species</a> (1877)</li> <li><a href="/wiki/The_Power_of_Movement_in_Plants" title="The Power of Movement in Plants">The Power of Movement in Plants</a> (1880)</li> <li><a href="/wiki/The_Formation_of_Vegetable_Mould_Through_the_Action_of_Worms" title="The Formation of Vegetable Mould Through the Action of Worms">The Formation of Vegetable Mould Through the Action of Worms</a> (1881)</li> <li><a href="/wiki/The_Autobiography_of_Charles_Darwin" title="The Autobiography of Charles Darwin">The Autobiography of Charles Darwin</a> (1887)</li> <li><a href="/wiki/Correspondence_of_Charles_Darwin" title="Correspondence of Charles Darwin">Correspondence</a> <ul><li><a href="/wiki/The_Life_and_Letters_of_Charles_Darwin" title="The Life and Letters of Charles Darwin">The Life and Letters of Charles Darwin</a>, 1887</li> <li><a href="/wiki/More_Letters_of_Charles_Darwin" title="More Letters of Charles Darwin">More Letters of Charles Darwin</a>, 1903</li></ul></li> <li><a href="/wiki/List_of_taxa_described_by_Charles_Darwin" title="List of taxa described by Charles Darwin">List of described taxa</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related</th><td class="navbox-list-with-group navbox-list navbox-odd wraplinks" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Eugenics" title="Eugenics">Eugenics</a></li> <li><a href="/wiki/Dysgenics" title="Dysgenics">Dysgenics</a></li> <li><a href="/wiki/Vestiges_of_the_Natural_History_of_Creation#Influence_and_effects" title="Vestiges of the Natural History of Creation">Vestiges of the Natural History of Creation</a></li> <li><a href="/wiki/The_Genetical_Theory_of_Natural_Selection" title="The Genetical Theory of Natural Selection">The Genetical Theory of Natural Selection</a> (1930)</li> <li><a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">History of evolutionary thought</a></li> <li><a href="/wiki/Pangenesis" title="Pangenesis">Pangenesis</a></li> <li><a href="/wiki/Darwin_Industry" title="Darwin Industry">Darwin Industry</a></li> <li><a href="/wiki/Commemoration_of_Charles_Darwin" title="Commemoration of Charles Darwin">Commemoration</a> <ul><li><a href="/wiki/List_of_things_named_after_Charles_Darwin" title="List of things named after Charles Darwin">things named for Darwin</a></li></ul></li> <li><a href="/wiki/Darwinism" title="Darwinism">Darwinism</a> <ul><li><a href="/wiki/Neural_Darwinism" title="Neural Darwinism">Neural</a></li> <li><a href="/wiki/Quantum_Darwinism" title="Quantum Darwinism">Quantum</a></li> <li><a href="/wiki/Social_Darwinism" title="Social Darwinism">Social</a></li> <li><a href="/wiki/Universal_Darwinism" title="Universal Darwinism">Universal</a></li> <li><a href="/wiki/Alternatives_to_Darwinian_evolution" title="Alternatives to Darwinian evolution">Alternatives to</a></li> <li><a href="/wiki/The_eclipse_of_Darwinism" title="The eclipse of Darwinism">Eclipse of</a></li></ul></li> <li><a href="/wiki/HMS_Beagle" title="HMS Beagle">HMS Beagle</a></li> <li><a href="/wiki/Darwin_Medal" title="Darwin Medal">Darwin Medal</a></li> <li><a href="/wiki/Darwin_Awards" title="Darwin Awards">Darwin Awards</a></li> <li><a href="/wiki/Huxley_family" title="Huxley family">Huxley family</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Evolutionary_biology753" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Evolution" title="Template:Evolution"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Evolution" title="Template talk:Evolution"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Evolution" title="Special:EditPage/Template:Evolution"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Evolutionary_biology753" style="font-size:114%;margin:0 4em"><a href="/wiki/Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><a href="/wiki/Introduction_to_evolution" title="Introduction to evolution">Introduction</a></li> <li><a href="/wiki/Outline_of_evolution" title="Outline of evolution">Outline</a></li> <li><a href="/wiki/Timeline_of_the_evolutionary_history_of_life" title="Timeline of the evolutionary history of life">Timeline of evolution</a></li> <li><a href="/wiki/History_of_life" title="History of life">History of life</a></li> <li><a href="/wiki/Index_of_evolutionary_biology_articles" title="Index of evolutionary biology articles">Index</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Evolution" title="Evolution">Evolution</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Abiogenesis" title="Abiogenesis">Abiogenesis</a></li> <li><a href="/wiki/Adaptation" title="Adaptation">Adaptation</a></li> <li><a href="/wiki/Adaptive_radiation" title="Adaptive radiation">Adaptive radiation</a></li> <li><a href="/wiki/Altruism_(biology)" title="Altruism (biology)">Altruism</a> <ul><li><a href="/wiki/Cheating_(biology)" title="Cheating (biology)">Cheating</a></li> <li><a href="/wiki/Reciprocal_altruism" title="Reciprocal altruism">Reciprocal</a></li></ul></li> <li><a href="/wiki/Baldwin_effect" title="Baldwin effect">Baldwin effect</a></li> <li><a href="/wiki/Cladistics" title="Cladistics">Cladistics</a></li> <li><a href="/wiki/Coevolution" title="Coevolution">Coevolution</a> <ul><li><a href="/wiki/Mutualism_(biology)" title="Mutualism (biology)">Mutualism</a></li></ul></li> <li><a href="/wiki/Common_descent" title="Common descent">Common descent</a></li> <li><a href="/wiki/Convergent_evolution" title="Convergent evolution">Convergence</a></li> <li><a href="/wiki/Divergent_evolution" title="Divergent evolution">Divergence</a></li> <li><a href="/wiki/Earliest_known_life_forms" title="Earliest known life forms">Earliest known life forms</a></li> <li><a href="/wiki/Evidence_of_common_descent" title="Evidence of common descent">Evidence of evolution</a></li> <li><a href="/wiki/Evolutionary_arms_race" title="Evolutionary arms race">Evolutionary arms race</a></li> <li><a href="/wiki/Evolutionary_pressure" title="Evolutionary pressure">Evolutionary pressure</a></li> <li><a href="/wiki/Exaptation" title="Exaptation">Exaptation</a></li> <li><a href="/wiki/Extinction" title="Extinction">Extinction</a> <ul><li><a href="/wiki/Extinction_event" title="Extinction event">Event</a></li></ul></li> <li><a href="/wiki/Homology_(biology)" title="Homology (biology)">Homology</a></li> <li><a href="/wiki/Last_universal_common_ancestor" title="Last universal common ancestor">Last universal common ancestor</a></li> <li><a href="/wiki/Macroevolution" title="Macroevolution">Macroevolution</a></li> <li><a href="/wiki/Microevolution" title="Microevolution">Microevolution</a></li> <li><a href="/wiki/Evolutionary_mismatch" title="Evolutionary mismatch">Mismatch</a></li> <li><a href="/wiki/Nonadaptive_radiation" title="Nonadaptive radiation">Non-adaptive radiation</a></li> <li><a href="/wiki/Abiogenesis" title="Abiogenesis">Origin of life</a></li> <li><a href="/wiki/Panspermia" title="Panspermia">Panspermia</a></li> <li><a href="/wiki/Parallel_evolution" title="Parallel evolution">Parallel evolution</a></li> <li><a href="/wiki/Signalling_theory" title="Signalling theory">Signalling theory</a> <ul><li><a href="/wiki/Handicap_principle" title="Handicap principle">Handicap principle</a></li></ul></li> <li><a href="/wiki/Speciation" title="Speciation">Speciation</a> <ul><li><a href="/wiki/Species" title="Species">Species</a></li> <li><a href="/wiki/Species_complex" title="Species complex">Species complex</a></li></ul></li> <li><a href="/wiki/Evolutionary_taxonomy" title="Evolutionary taxonomy">Taxonomy</a></li> <li><a href="/wiki/Unit_of_selection" title="Unit of selection">Unit of selection</a> <ul><li><a href="/wiki/Gene-centered_view_of_evolution" title="Gene-centered view of evolution">Gene-centered view of evolution</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Population_genetics" title="Population genetics">Population genetics</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Selective_breeding" title="Selective breeding">Artificial selection</a></li> <li><a href="/wiki/Biodiversity" title="Biodiversity">Biodiversity</a></li> <li><a href="/wiki/Evolutionarily_stable_strategy" title="Evolutionarily stable strategy">Evolutionarily stable strategy</a></li> <li><a href="/wiki/Fisher%27s_principle" title="Fisher's principle">Fisher's principle</a></li> <li><a href="/wiki/Fitness_(biology)" title="Fitness (biology)">Fitness</a> <ul><li><a href="/wiki/Inclusive_fitness" title="Inclusive fitness">Inclusive</a></li></ul></li> <li><a href="/wiki/Gene_flow" title="Gene flow">Gene flow</a></li> <li><a href="/wiki/Genetic_drift" title="Genetic drift">Genetic drift</a></li> <li><a href="/wiki/Kin_selection" title="Kin selection">Kin selection</a> <ul><li><a href="/wiki/Parental_investment" title="Parental investment">Parental investment</a></li> <li><a href="/wiki/Parent%E2%80%93offspring_conflict" title="Parent–offspring conflict">Parent–offspring conflict</a></li></ul></li> <li><a href="/wiki/Mutation" title="Mutation">Mutation</a></li> <li><a href="/wiki/Population" title="Population">Population</a></li> <li><a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a></li> <li><a href="/wiki/Sexual_dimorphism" title="Sexual dimorphism">Sexual dimorphism</a></li> <li><a href="/wiki/Sexual_selection" title="Sexual selection">Sexual selection</a> <ul><li><a href="/wiki/Sexual_selection_in_flowering_plants" title="Sexual selection in flowering plants">Flowering plants</a></li> <li><a href="/wiki/Sexual_selection_in_fungi" title="Sexual selection in fungi">Fungi</a></li> <li><a href="/wiki/Mate_choice" title="Mate choice">Mate choice</a></li></ul></li> <li><a href="/wiki/Social_selection" title="Social selection">Social selection</a></li> <li><a href="/wiki/Trivers%E2%80%93Willard_hypothesis" title="Trivers–Willard hypothesis">Trivers–Willard hypothesis</a></li> <li><a href="/wiki/Genetic_variation" title="Genetic variation">Variation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Development</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Canalisation_(genetics)" title="Canalisation (genetics)">Canalisation</a></li> <li><a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evolutionary developmental biology</a></li> <li><a href="/wiki/Genetic_assimilation" title="Genetic assimilation">Genetic assimilation</a></li> <li><a href="/wiki/Inversion_(evolutionary_biology)" title="Inversion (evolutionary biology)">Inversion</a></li> <li><a href="/wiki/Modularity_(biology)" title="Modularity (biology)">Modularity</a></li> <li><a href="/wiki/Phenotypic_plasticity" title="Phenotypic plasticity">Phenotypic plasticity</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Taxon" title="Taxon">taxa</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_bacteria" title="Evolution of bacteria">Bacteria</a></li> <li><a href="/wiki/Evolution_of_birds" title="Evolution of birds">Birds</a> <ul><li><a href="/wiki/Origin_of_birds" title="Origin of birds">origin</a></li></ul></li> <li><a href="/wiki/Evolution_of_brachiopods" title="Evolution of brachiopods">Brachiopods</a></li> <li><a href="/wiki/Evolution_of_molluscs" title="Evolution of molluscs">Molluscs</a> <ul><li><a href="/wiki/Evolution_of_cephalopods" title="Evolution of cephalopods">Cephalopods</a></li></ul></li> <li><a href="/wiki/Dinosaur#Evolutionary_history" title="Dinosaur">Dinosaurs</a></li> <li><a href="/wiki/Evolution_of_fish" title="Evolution of fish">Fish</a></li> <li><a href="/wiki/Evolution_of_fungi" title="Evolution of fungi">Fungi</a></li> <li><a href="/wiki/Evolution_of_insects" title="Evolution of insects">Insects</a> <ul><li><a href="/wiki/Evolution_of_butterflies" title="Evolution of butterflies">butterflies</a></li></ul></li> <li><a href="/wiki/History_of_life" title="History of life">Life</a></li> <li><a href="/wiki/Evolution_of_mammals" title="Evolution of mammals">Mammals</a> <ul><li><a href="/wiki/Cat_gap" title="Cat gap">cats</a></li> <li><a href="/wiki/Canidae#Evolution" title="Canidae">canids</a> <ul><li><a href="/wiki/Evolution_of_the_wolf" title="Evolution of the wolf">wolves</a></li> <li><a href="/wiki/Domestication_of_the_dog" title="Domestication of the dog">dogs</a></li></ul></li> <li><a href="/wiki/Hyena#Evolution" title="Hyena">hyenas</a></li> <li><a href="/wiki/Evolution_of_cetaceans" title="Evolution of cetaceans">dolphins and whales</a></li> <li><a href="/wiki/Evolution_of_the_horse" title="Evolution of the horse">horses</a></li> <li><a href="/wiki/Evolution_of_Macropodidae" title="Evolution of Macropodidae">Kangaroos</a></li> <li><a href="/wiki/Evolution_of_primates" title="Evolution of primates">primates</a> <ul><li><a href="/wiki/Human_evolution" title="Human evolution">humans</a></li> <li><a href="/wiki/Evolution_of_lemurs" title="Evolution of lemurs">lemurs</a></li></ul></li> <li><a href="/wiki/Evolution_of_sirenians" title="Evolution of sirenians">sea cows</a></li></ul></li> <li><a href="/wiki/Evolutionary_history_of_plants" title="Evolutionary history of plants">Plants</a> <ul><li><a href="/wiki/Pollinator-mediated_selection" title="Pollinator-mediated selection">pollinator-mediated</a></li></ul></li> <li><a href="/wiki/Evolution_of_reptiles" title="Evolution of reptiles">Reptiles</a></li> <li><a href="/wiki/Evolution_of_spiders" title="Evolution of spiders">Spiders</a></li> <li><a href="/wiki/Evolution_of_tetrapods" title="Evolution of tetrapods">Tetrapods</a></li> <li><a href="/wiki/Viral_evolution" title="Viral evolution">Viruses</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Organ_(anatomy)" class="mw-redirect" title="Organ (anatomy)">organs</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_cells" title="Evolution of cells">Cell</a></li> <li><a href="/wiki/Models_of_DNA_evolution" title="Models of DNA evolution">DNA</a></li> <li><a href="/wiki/Evolution_of_flagella" title="Evolution of flagella">Flagella</a></li> <li><a href="/wiki/Eukaryote#Origin_of_eukaryotes" title="Eukaryote">Eukaryotes</a> <ul><li><a href="/wiki/Symbiogenesis" title="Symbiogenesis">symbiogenesis</a></li> <li><a href="/wiki/Chromosome#Eukaryotes" title="Chromosome">chromosome</a></li> <li><a href="/wiki/Endomembrane_system#Evolution" title="Endomembrane system">endomembrane system</a></li> <li><a href="/wiki/Mitochondrion#Origin_and_evolution" title="Mitochondrion">mitochondria</a></li> <li><a href="/wiki/Cell_nucleus#Evolution" title="Cell nucleus">nucleus</a></li> <li><a href="/wiki/Plastid#Origin" title="Plastid">plastids</a></li></ul></li> <li>In animals <ul><li><a href="/wiki/Evolution_of_the_eye" title="Evolution of the eye">eye</a></li> <li><a href="/wiki/Hair#Evolution" title="Hair">hair</a></li> <li><a href="/wiki/Evolution_of_mammalian_auditory_ossicles" title="Evolution of mammalian auditory ossicles">auditory ossicle</a></li> <li><a href="/wiki/Evolution_of_nervous_systems" title="Evolution of nervous systems">nervous system</a></li> <li><a href="/wiki/Evolution_of_the_brain" title="Evolution of the brain">brain</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Biological_process" title="Biological process">processes</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_ageing" title="Evolution of ageing">Aging</a> <ul><li><a href="/wiki/Death#Evolution_of_aging_and_mortality" title="Death">Death</a></li> <li><a href="/wiki/Programmed_cell_death#Evolutionary_origin_of_mitochondrial_apoptosis" title="Programmed cell death">Programmed cell death</a></li></ul></li> <li><a href="/wiki/Origin_of_avian_flight" title="Origin of avian flight">Avian flight</a></li> <li><a href="/wiki/Evolution_of_biological_complexity" title="Evolution of biological complexity">Biological complexity</a></li> <li><a href="/wiki/Cooperation_(evolution)" title="Cooperation (evolution)">Cooperation</a></li> <li><a href="/wiki/Evolution_of_color_vision" title="Evolution of color vision">Color vision</a> <ul><li><a href="/wiki/Evolution_of_color_vision_in_primates" title="Evolution of color vision in primates">in primates</a></li></ul></li> <li><a href="/wiki/Evolution_of_emotion" title="Evolution of emotion">Emotion</a></li> <li><a href="/wiki/Empathy#Evolution_across_species" title="Empathy">Empathy</a></li> <li><a href="/wiki/Evolutionary_ethics" title="Evolutionary ethics">Ethics</a></li> <li><a href="/wiki/Evolution_of_eusociality" title="Evolution of eusociality">Eusociality</a></li> <li><a href="/wiki/Immune_system#Evolution_and_other_mechanisms" title="Immune system">Immune system</a></li> <li><a href="/wiki/Metabolism#Evolution" title="Metabolism">Metabolism</a></li> <li><a href="/wiki/Monogamy_in_animals" title="Monogamy in animals">Monogamy</a></li> <li><a href="/wiki/Evolution_of_morality" title="Evolution of morality">Morality</a></li> <li><a href="/wiki/Mosaic_evolution" title="Mosaic evolution">Mosaic evolution</a></li> <li><a href="/wiki/Multicellular_organism#Evolutionary_history" title="Multicellular organism">Multicellularity</a></li> <li><a href="/wiki/Evolution_of_sexual_reproduction" title="Evolution of sexual reproduction">Sexual reproduction</a> <ul><li><a href="/wiki/Anisogamy#Evolution" title="Anisogamy">Gamete differentiation/sexes</a></li> <li><a href="/wiki/Biological_life_cycle" title="Biological life cycle">Life cycles/nuclear phases</a></li> <li><a href="/wiki/Mating_type" title="Mating type">Mating types</a></li> <li><a href="/wiki/Origin_and_function_of_meiosis" title="Origin and function of meiosis">Meiosis</a></li> <li><a href="/wiki/Sex-determination_system#Evolution" title="Sex-determination system">Sex-determination</a></li></ul></li> <li><a href="/wiki/Evolution_of_snake_venom" title="Evolution of snake venom">Snake venom</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Tempo_and_Mode_in_Evolution" title="Tempo and Mode in Evolution">Tempo and modes</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Phyletic_gradualism" title="Phyletic gradualism">Gradualism</a>/<a href="/wiki/Punctuated_equilibrium" title="Punctuated equilibrium">Punctuated equilibrium</a>/<a href="/wiki/Saltation_(biology)" title="Saltation (biology)">Saltationism</a></li> <li><a href="/wiki/Point_mutation" title="Point mutation">Micromutation</a>/<a href="/wiki/Mutationism#Later_mutationist_theories" title="Mutationism">Macromutation</a></li> <li><a href="/wiki/Uniformitarianism" title="Uniformitarianism">Uniformitarianism</a>/<a href="/wiki/Catastrophism" title="Catastrophism">Catastrophism</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Speciation" title="Speciation">Speciation</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Allopatric_speciation" title="Allopatric speciation">Allopatric</a></li> <li><a href="/wiki/Anagenesis" title="Anagenesis">Anagenesis</a></li> <li><a href="/wiki/Catagenesis_(biology)" title="Catagenesis (biology)">Catagenesis</a></li> <li><a href="/wiki/Cladogenesis" title="Cladogenesis">Cladogenesis</a></li> <li><a href="/wiki/Cospeciation" title="Cospeciation">Cospeciation</a></li> <li><a href="/wiki/Ecological_speciation" title="Ecological speciation">Ecological</a></li> <li><a href="/wiki/Hybrid_speciation" title="Hybrid speciation">Hybrid</a></li> <li><a href="/wiki/Nonecological_speciation" title="Nonecological speciation">Non-ecological</a></li> <li><a href="/wiki/Parapatric_speciation" title="Parapatric speciation">Parapatric</a></li> <li><a href="/wiki/Peripatric_speciation" title="Peripatric speciation">Peripatric</a></li> <li><a href="/wiki/Reinforcement_(speciation)" title="Reinforcement (speciation)">Reinforcement</a></li> <li><a href="/wiki/Sympatric_speciation" title="Sympatric speciation">Sympatric</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">History</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolutionary_ideas_of_the_Renaissance_and_Enlightenment" title="Evolutionary ideas of the Renaissance and Enlightenment">Renaissance and Enlightenment</a></li> <li><a href="/wiki/Transmutation_of_species" title="Transmutation of species">Transmutation of species</a></li> <li><a href="/wiki/David_Hume" title="David Hume">David Hume</a> <ul><li><a href="/wiki/Dialogues_Concerning_Natural_Religion" title="Dialogues Concerning Natural Religion">Dialogues Concerning Natural Religion</a></li></ul></li> <li><a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a> <ul><li><a class="mw-selflink selflink">On the Origin of Species</a></li></ul></li> <li><a href="/wiki/History_of_paleontology" title="History of paleontology">History of paleontology</a></li> <li><a href="/wiki/Transitional_fossil" title="Transitional fossil">Transitional fossil</a></li> <li><a href="/wiki/Blending_inheritance" title="Blending inheritance">Blending inheritance</a></li> <li><a href="/wiki/Mendelian_inheritance" title="Mendelian inheritance">Mendelian inheritance</a></li> <li><a href="/wiki/The_eclipse_of_Darwinism" title="The eclipse of Darwinism">The eclipse of Darwinism</a></li> <li><a href="/wiki/Neo-Darwinism" title="Neo-Darwinism">Neo-Darwinism</a></li> <li><a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis</a></li> <li><a href="/wiki/History_of_molecular_evolution" title="History of molecular evolution">History of molecular evolution</a></li> <li><a href="/wiki/Extended_evolutionary_synthesis" title="Extended evolutionary synthesis">Extended evolutionary synthesis</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Philosophy</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Darwinism" title="Darwinism">Darwinism</a></li> <li><a href="/wiki/Alternatives_to_Darwinian_evolution" title="Alternatives to Darwinian evolution">Alternatives</a> <ul><li><a href="/wiki/Catastrophism" title="Catastrophism">Catastrophism</a></li> <li><a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a></li> <li><a href="/wiki/Orthogenesis" title="Orthogenesis">Orthogenesis</a></li> <li><a href="/wiki/Mutationism" title="Mutationism">Mutationism</a></li> <li><a href="/wiki/Saltation_(biology)" title="Saltation (biology)">Saltationism</a></li> <li><a href="/wiki/Structuralism_(biology)" title="Structuralism (biology)">Structuralism</a> <ul><li><a href="/wiki/Spandrel_(biology)" title="Spandrel (biology)">Spandrel</a></li></ul></li> <li><a href="/wiki/Theistic_evolution" title="Theistic evolution">Theistic</a></li> <li><a href="/wiki/Vitalism" title="Vitalism">Vitalism</a></li></ul></li> <li><a href="/wiki/Teleology_in_biology" title="Teleology in biology">Teleology in biology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Biogeography" title="Biogeography">Biogeography</a></li> <li><a href="/wiki/Ecological_genetics" title="Ecological genetics">Ecological genetics</a></li> <li><a href="/wiki/Evolutionary_medicine" title="Evolutionary medicine">Evolutionary medicine</a></li> <li><a href="/wiki/Group_selection" title="Group selection">Group selection</a> <ul><li><a href="/wiki/Cultural_evolution" title="Cultural evolution">Cultural evolution</a></li> <li><a href="/wiki/Cultural_group_selection" title="Cultural group selection">Cultural group selection</a></li> <li><a href="/wiki/Dual_inheritance_theory" title="Dual inheritance theory">Dual inheritance theory</a></li></ul></li> <li><a href="/wiki/Hologenome_theory_of_evolution" title="Hologenome theory of evolution">Hologenome theory of evolution</a></li> <li><a href="/wiki/Missing_heritability_problem" title="Missing heritability problem">Missing heritability problem</a></li> <li><a href="/wiki/Molecular_evolution" title="Molecular evolution">Molecular evolution</a></li> <li><a href="/wiki/Astrobiology" title="Astrobiology">Astrobiology</a></li> <li><a href="/wiki/Phylogenetics" title="Phylogenetics">Phylogenetics</a> <ul><li><a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">Tree</a></li></ul></li> <li><a href="/wiki/Polymorphism_(biology)" title="Polymorphism (biology)">Polymorphism</a></li> <li><a href="/wiki/Protocell" title="Protocell">Protocell</a></li> <li><a href="/wiki/Systematics" title="Systematics">Systematics</a></li> <li><a href="/wiki/Transgenerational_epigenetic_inheritance" title="Transgenerational epigenetic inheritance">Transgenerational epigenetic inheritance</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><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" /> <a href="/wiki/Category:Evolutionary_biology" title="Category:Evolutionary biology">Category</a></li> <li><a href="/wiki/File:Symbol_portal_class.svg" class="mw-file-description" title="Portal"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/16px-Symbol_portal_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/23px-Symbol_portal_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/31px-Symbol_portal_class.svg.png 2x" data-file-width="180" data-file-height="185" /></a> <a href="/wiki/Portal:Evolutionary_biology" title="Portal:Evolutionary biology">Portal</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Natural_history50" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="3"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Natural_history" title="Template:Natural history"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Natural_history" title="Template talk:Natural history"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Natural_history" title="Special:EditPage/Template:Natural history"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Natural_history50" style="font-size:114%;margin:0 4em"><a href="/wiki/Natural_history" title="Natural history">Natural history</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Pioneering naturalists</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:1%"><a href="/wiki/Classical_antiquity" title="Classical antiquity">Classical antiquity</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/Aristotle" title="Aristotle">Aristotle</a> (<a href="/wiki/History_of_Animals" title="History of Animals">History of Animals</a>)</li> <li><a href="/wiki/Theophrastus" title="Theophrastus">Theophrastus</a> (<a href="/wiki/Historia_Plantarum_(Theophrastus_book)" class="mw-redirect" title="Historia Plantarum (Theophrastus book)">Historia Plantarum</a>)</li> <li><a href="/wiki/Claudius_Aelianus" title="Claudius Aelianus">Aelian</a> (De Natura Animalium)</li> <li><a href="/wiki/Pliny_the_Elder" title="Pliny the Elder">Pliny the Elder</a> (<a href="/wiki/Natural_History_(Pliny)" title="Natural History (Pliny)">Natural History</a>)</li> <li><a href="/wiki/Pedanius_Dioscorides" class="mw-redirect" title="Pedanius Dioscorides">Dioscorides</a> (<a href="/wiki/De_Materia_Medica" class="mw-redirect" title="De Materia Medica">De Materia Medica</a>)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Renaissance" title="Renaissance">Renaissance</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/Ulisse_Aldrovandi" title="Ulisse Aldrovandi">Ulisse Aldrovandi</a></li> <li><a href="/wiki/Gaspard_Bauhin" title="Gaspard Bauhin">Gaspard Bauhin</a> (Pinax theatri botanici)</li> <li><a href="/wiki/Otto_Brunfels" title="Otto Brunfels">Otto Brunfels</a></li> <li><a href="/wiki/Hieronymus_Bock" title="Hieronymus Bock">Hieronymus Bock</a></li> <li><a href="/wiki/Andrea_Cesalpino" title="Andrea Cesalpino">Andrea Cesalpino</a></li> <li><a href="/wiki/Valerius_Cordus" title="Valerius Cordus">Valerius Cordus</a></li> <li><a href="/wiki/Leonhart_Fuchs" title="Leonhart Fuchs">Leonhart Fuchs</a></li> <li><a href="/wiki/Conrad_Gessner" title="Conrad Gessner">Conrad Gessner</a> (<a href="/wiki/Historia_animalium_(Gessner_book)" title="Historia animalium (Gessner book)">Historia animalium</a>)</li> <li><a href="/wiki/Frederik_Ruysch" title="Frederik Ruysch">Frederik Ruysch</a></li> <li><a href="/wiki/William_Turner_(naturalist)" title="William Turner (naturalist)">William Turner</a> (Avium Praecipuarum, New Herball)</li> <li><a href="/wiki/John_Gerard" title="John Gerard">John Gerard</a> (Herball, or Generall Historie of Plantes)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Age_of_Enlightenment" title="Age of Enlightenment">Enlightenment</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/Robert_Hooke" title="Robert Hooke">Robert Hooke</a> (<a href="/wiki/Micrographia" title="Micrographia">Micrographia</a>)</li> <li><a href="/wiki/Marcello_Malpighi" title="Marcello Malpighi">Marcello Malpighi</a></li> <li><a href="/wiki/Antonie_van_Leeuwenhoek" title="Antonie van Leeuwenhoek">Antonie van Leeuwenhoek</a></li> <li><a href="/wiki/William_Derham" title="William Derham">William Derham</a></li> <li><a href="/wiki/Hans_Sloane" title="Hans Sloane">Hans Sloane</a></li> <li><a href="/wiki/Jan_Swammerdam" title="Jan Swammerdam">Jan Swammerdam</a></li> <li><a href="/wiki/Regnier_de_Graaf" title="Regnier de Graaf">Regnier de Graaf</a></li> <li><a href="/wiki/Carl_Linnaeus" title="Carl Linnaeus">Carl Linnaeus</a> (<a href="/wiki/Systema_Naturae" title="Systema Naturae">Systema Naturae</a>)</li> <li><a href="/wiki/Georg_Wilhelm_Steller" title="Georg Wilhelm Steller">Georg Steller</a></li> <li><a href="/wiki/Joseph_Banks" title="Joseph Banks">Joseph Banks</a></li> <li><a href="/wiki/Johan_Christian_Fabricius" title="Johan Christian Fabricius">Johan Christian Fabricius</a></li> <li><a href="/wiki/James_Hutton" title="James Hutton">James Hutton</a></li> <li><a href="/wiki/John_Ray" title="John Ray">John Ray</a> (<a href="/wiki/Historia_Plantarum_(Ray_book)" title="Historia Plantarum (Ray book)">Historia Plantarum</a>)</li> <li><a href="/wiki/Georges-Louis_Leclerc,_Comte_de_Buffon" title="Georges-Louis Leclerc, Comte de Buffon">Comte de Buffon</a> (<a href="/wiki/Histoire_Naturelle" title="Histoire Naturelle">Histoire Naturelle</a>)</li> <li><a href="/wiki/Bernard_Germain_de_Lac%C3%A9p%C3%A8de" title="Bernard Germain de Lacépède">Bernard Germain de Lacépède</a></li> <li><a href="/wiki/Gilbert_White" title="Gilbert White">Gilbert White</a> (<a href="/wiki/The_Natural_History_and_Antiquities_of_Selborne" title="The Natural History and Antiquities of Selborne">The Natural History of Selborne</a>)</li> <li><a href="/wiki/Thomas_Bewick" title="Thomas Bewick">Thomas Bewick</a> (<a href="/wiki/A_History_of_British_Birds" title="A History of British Birds">A History of British Birds</a>)</li> <li><a href="/wiki/Jean-Baptiste_Lamarck" title="Jean-Baptiste Lamarck">Jean-Baptiste Lamarck</a> (<a href="/wiki/Philosophie_zoologique" title="Philosophie zoologique">Philosophie zoologique</a>)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">19th century</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/George_Montagu_(naturalist)" title="George Montagu (naturalist)">George Montagu</a> (<a href="/wiki/Ornithological_Dictionary" title="Ornithological Dictionary">Ornithological Dictionary</a>)</li> <li><a href="/wiki/Georges_Cuvier" title="Georges Cuvier">Georges Cuvier</a> (<a href="/wiki/Le_R%C3%A8gne_Animal" title="Le Règne Animal">Le Règne Animal</a>)</li> <li><a href="/wiki/William_Smith_(geologist)" title="William Smith (geologist)">William Smith</a></li> <li><a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a> (<a class="mw-selflink selflink">On the Origin of Species</a>)</li> <li><a href="/wiki/Alfred_Russel_Wallace" title="Alfred Russel Wallace">Alfred Russel Wallace</a> (<a href="/wiki/The_Malay_Archipelago" title="The Malay Archipelago">The Malay Archipelago</a>)</li> <li><a href="/wiki/Henry_Walter_Bates" title="Henry Walter Bates">Henry Walter Bates</a> (<a href="/wiki/The_Naturalist_on_the_River_Amazons" title="The Naturalist on the River Amazons">The Naturalist on the River Amazons</a>)</li> <li><a href="/wiki/Alexander_von_Humboldt" title="Alexander von Humboldt">Alexander von Humboldt</a></li> <li><a href="/wiki/John_James_Audubon" title="John James Audubon">John James Audubon</a> (<a href="/wiki/The_Birds_of_America" title="The Birds of America">The Birds of America</a>)</li> <li><a href="/wiki/William_Buckland" title="William Buckland">William Buckland</a></li> <li><a href="/wiki/Charles_Lyell" title="Charles Lyell">Charles Lyell</a></li> <li><a href="/wiki/Mary_Anning" title="Mary Anning">Mary Anning</a></li> <li><a href="/wiki/Jean-Henri_Fabre" title="Jean-Henri Fabre">Jean-Henri Fabre</a></li> <li><a href="/wiki/Louis_Agassiz" title="Louis Agassiz">Louis Agassiz</a></li> <li><a href="/wiki/Philip_Henry_Gosse" title="Philip Henry Gosse">Philip Henry Gosse</a></li> <li><a href="/wiki/Asa_Gray" title="Asa Gray">Asa Gray</a></li> <li><a href="/wiki/William_Jackson_Hooker" title="William Jackson Hooker">William Jackson Hooker</a></li> <li><a href="/wiki/Joseph_Dalton_Hooker" title="Joseph Dalton Hooker">Joseph Dalton Hooker</a></li> <li><a href="/wiki/Sir_William_Jardine,_7th_Baronet" title="Sir William Jardine, 7th Baronet">William Jardine</a> (The Naturalist's Library)</li> <li><a href="/wiki/Ernst_Haeckel" title="Ernst Haeckel">Ernst Haeckel</a> (<a href="/wiki/Kunstformen_der_Natur" title="Kunstformen der Natur">Kunstformen der Natur</a>)</li> <li><a href="/wiki/Richard_Lydekker" title="Richard Lydekker">Richard Lydekker</a> (The Royal Natural History)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">20th century</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/Martinus_Beijerinck" title="Martinus Beijerinck">Martinus Beijerinck</a></li> <li><a href="/wiki/Abbott_Handerson_Thayer" title="Abbott Handerson Thayer">Abbott Thayer</a> (<a href="/wiki/Concealing-Coloration_in_the_Animal_Kingdom" title="Concealing-Coloration in the Animal Kingdom">Concealing-Coloration in the Animal Kingdom</a>)</li> <li><a href="/wiki/Hugh_B._Cott" title="Hugh B. Cott">Hugh B. Cott</a> (<a href="/wiki/Adaptive_Coloration_in_Animals" title="Adaptive Coloration in Animals">Adaptive Coloration in Animals</a>)</li> <li><a href="/wiki/Nikolaas_Tinbergen" title="Nikolaas Tinbergen">Niko Tinbergen</a> (The Study of Instinct)</li> <li><a href="/wiki/Konrad_Lorenz" title="Konrad Lorenz">Konrad Lorenz</a> (<a href="/wiki/On_Aggression" title="On Aggression">On Aggression</a>)</li> <li><a href="/wiki/Karl_von_Frisch" title="Karl von Frisch">Karl von Frisch</a> (The Dancing Bees)</li> <li><a href="/wiki/Ronald_Lockley" title="Ronald Lockley">Ronald Lockley</a> (Shearwaters)</li></ul> </div></td></tr></tbody></table><div></div></td><td class="noviewer navbox-image" rowspan="2" style="width:1px;padding:0 0 0 2px"><div><a href="/wiki/File:Naturalist_on_the_River_Amazons_figure_17.png" class="mw-file-description" title="Sceliphron wasp building nest"><img alt="Sceliphron wasp building nest" src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f8/Naturalist_on_the_River_Amazons_figure_17.png/85px-Naturalist_on_the_River_Amazons_figure_17.png" decoding="async" width="85" height="123" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f8/Naturalist_on_the_River_Amazons_figure_17.png/128px-Naturalist_on_the_River_Amazons_figure_17.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f8/Naturalist_on_the_River_Amazons_figure_17.png/170px-Naturalist_on_the_River_Amazons_figure_17.png 2x" data-file-width="887" data-file-height="1280" /></a></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Topics</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Natural_history_museum" title="Natural history museum">Natural history museums</a> (<a href="/wiki/List_of_natural_history_museums" title="List of natural history museums">List</a>)</li> <li><a href="/wiki/Parson-naturalist" title="Parson-naturalist">Parson-naturalists</a> (<a href="/wiki/List_of_parson-naturalists" title="List of parson-naturalists">List</a>)</li> <li><a href="/wiki/Category:Natural_history_societies" title="Category:Natural history societies">Natural History Societies</a></li> <li><a href="/wiki/List_of_natural_history_dealers" title="List of natural history dealers">List of natural history dealers</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="History_of_biology_(timeline)253" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:History_of_biology" title="Template:History of biology"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:History_of_biology" title="Template talk:History of biology"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:History_of_biology" title="Special:EditPage/Template:History of biology"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="History_of_biology_(timeline)253" style="font-size:114%;margin:0 4em"><a href="/wiki/History_of_biology" title="History of biology">History of biology</a> (<a href="/wiki/Timeline_of_biology_and_organic_chemistry" title="Timeline of biology and organic chemistry">timeline</a>)</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Fields, disciplines</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/History_of_agricultural_science" title="History of agricultural science">Agricultural science</a></li> <li><a href="/wiki/History_of_anatomy" title="History of anatomy">Anatomy</a></li> <li><a href="/wiki/History_of_biochemistry" title="History of biochemistry">Biochemistry</a></li> <li><a href="/wiki/History_of_biotechnology" title="History of biotechnology">Biotechnology</a></li> <li><a href="/wiki/History_of_botany" title="History of botany">Botany</a></li> <li><a href="/wiki/History_of_ecology" title="History of ecology">Ecology</a></li> <li><a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">Evolutionary thought</a></li> <li><a href="/wiki/History_of_genetics" title="History of genetics">Genetics</a></li> <li><a href="/wiki/History_of_geology" title="History of geology">Geology</a></li> <li><a href="/wiki/Timeline_of_immunology" title="Timeline of immunology">Immunology</a></li> <li><a href="/wiki/History_of_medicine" title="History of medicine">Medicine</a></li> <li><a href="/wiki/History_of_model_organisms" title="History of model organisms">Model organisms</a></li> <li><a href="/wiki/History_of_molecular_biology" title="History of molecular biology">Molecular biology</a></li> <li><a href="/wiki/History_of_molecular_evolution" title="History of molecular evolution">Molecular evolution</a></li> <li><a href="/wiki/History_of_paleontology" title="History of paleontology">Paleontology</a></li> <li><a href="/wiki/History_of_phycology" title="History of phycology">Phycology</a></li> <li><a href="/wiki/History_of_plant_systematics" title="History of plant systematics">Plant systematics</a></li> <li><a href="/wiki/History_of_RNA_biology" title="History of RNA biology">RNA biology</a></li> <li><a href="/wiki/History_of_zoology_through_1859" title="History of zoology through 1859">Zoology (through 1859)</a></li> <li><a href="/wiki/History_of_zoology_since_1859" class="mw-redirect" title="History of zoology since 1859">Zoology (since 1859)</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Theories, concepts</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/Germ_theory_of_disease" title="Germ theory of disease">Germ theory of disease</a></li> <li><a href="/wiki/Central_dogma_of_molecular_biology" title="Central dogma of molecular biology">Central dogma of molecular biology</a></li> <li><a href="/wiki/Darwinism" title="Darwinism">Darwinism</a></li> <li><a href="/wiki/Great_chain_of_being" title="Great chain of being">Great chain of being</a></li> <li><a href="/wiki/Biological_organisation" title="Biological organisation">Hierarchy of life</a></li> <li><a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a></li> <li><a href="/wiki/One_gene%E2%80%93one_enzyme_hypothesis" title="One gene–one enzyme hypothesis">One gene–one enzyme hypothesis</a></li> <li><a href="/wiki/Protocell" title="Protocell">Protocell</a></li> <li><a href="/wiki/RNA_world" title="RNA world">RNA world</a></li> <li><a href="/wiki/Sequence_hypothesis" title="Sequence hypothesis">Sequence hypothesis</a></li> <li><a href="/wiki/Spontaneous_generation" title="Spontaneous generation">Spontaneous generation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/History_of_science" title="History of science">History of science</a></li> <li><a href="/wiki/Philosophy_of_biology" title="Philosophy of biology">Philosophy of biology</a> <ul><li><a href="/wiki/Teleology_in_biology" title="Teleology in biology">Teleology</a></li></ul></li> <li><a href="/wiki/Ethnobotany" title="Ethnobotany">Ethnobotany</a></li> <li><a href="/wiki/Eugenics" title="Eugenics">Eugenics</a></li> <li><a href="/wiki/Dysgenics" title="Dysgenics">Dysgenics</a></li> <li><a href="/wiki/History_of_the_creation%E2%80%93evolution_controversy" title="History of the creation–evolution controversy">History of the creation-evolution controversy</a></li> <li><a href="/wiki/Human_Genome_Project" title="Human Genome Project">Human Genome Project</a></li> <li><a href="/wiki/Humboldtian_science" title="Humboldtian science">Humboldtian science</a></li> <li><a href="/wiki/Natural_history" title="Natural history">Natural history</a></li> <li><a href="/wiki/Natural_philosophy" title="Natural philosophy">Natural philosophy</a></li> <li><a href="/wiki/Natural_theology" title="Natural theology">Natural theology</a></li> <li><a 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</div><script>(RLQ=window.RLQ||[]).push(function(){mw.config.set({"wgHostname":"mw-web.codfw.canary-55fd9d4764-gtn8x","wgBackendResponseTime":164,"wgPageParseReport":{"limitreport":{"cputime":"2.382","walltime":"2.676","ppvisitednodes":{"value":12976,"limit":1000000},"postexpandincludesize":{"value":400868,"limit":2097152},"templateargumentsize":{"value":8585,"limit":2097152},"expansiondepth":{"value":13,"limit":100},"expensivefunctioncount":{"value":14,"limit":500},"unstrip-depth":{"value":1,"limit":20},"unstrip-size":{"value":545182,"limit":5000000},"entityaccesscount":{"value":1,"limit":400},"timingprofile":["100.00% 2210.725 1 -total"," 37.07% 819.476 1 Template:Reflist"," 23.50% 519.572 89 Template:Citation"," 11.52% 254.772 208 Template:Harvnb"," 11.34% 250.644 29 Template:Cite_web"," 6.63% 146.567 1 Template:Evolutionary_biology"," 6.49% 143.560 1 Template:Sidebar_with_collapsible_lists"," 5.88% 129.966 7 Template:Sfn"," 4.58% 101.233 1 Template:Infobox_book"," 3.69% 81.643 1 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[\"CITEREFDarwin2006\"] = 1,\n [\"CITEREFDarwinCosta2009\"] = 1,\n [\"CITEREFDesmond1989\"] = 1,\n [\"CITEREFDesmondMoore1991\"] = 1,\n [\"CITEREFDesmondMoore2009\"] = 1,\n [\"CITEREFDewey1994\"] = 1,\n [\"CITEREFDupree1988\"] = 1,\n [\"CITEREFEldredge2006\"] = 1,\n [\"CITEREFFinkelstein2013\"] = 1,\n [\"CITEREFForsterMarston1999\"] = 1,\n [\"CITEREFFreeman1977\"] = 1,\n [\"CITEREFGray1860\"] = 1,\n [\"CITEREFGray1874\"] = 1,\n [\"CITEREFHerbert1980\"] = 1,\n [\"CITEREFHodge1874\"] = 1,\n [\"CITEREFHuxley1860\"] = 1,\n [\"CITEREFHuxley1863\"] = 1,\n [\"CITEREFHuxleyKettlewell1965\"] = 1,\n [\"CITEREFJin2018\"] = 1,\n [\"CITEREFKeynes2000\"] = 1,\n [\"CITEREFKragh2016\"] = 1,\n [\"CITEREFKreeft2001\"] = 1,\n [\"CITEREFLarson2004\"] = 1,\n [\"CITEREFLeifchild1859\"] = 1,\n [\"CITEREFLucas1979\"] = 1,\n [\"CITEREFMayr1982\"] = 1,\n [\"CITEREFMichod,_Richard_E.1995\"] = 1,\n [\"CITEREFMiles2001\"] = 1,\n [\"CITEREFMivart1871\"] = 1,\n [\"CITEREFMoore2006\"] = 1,\n [\"CITEREFPechenik2023\"] = 1,\n [\"CITEREFPeckham1959\"] = 1,\n [\"CITEREFPhipps1983\"] = 1,\n [\"CITEREFPius_XII1950\"] = 1,\n [\"CITEREFQuammen2006\"] = 1,\n [\"CITEREFRadick2013\"] = 1,\n [\"CITEREFRhodes1987\"] = 1,\n [\"CITEREFRichard_Dawkins2003\"] = 1,\n [\"CITEREFRichards2017\"] = 1,\n [\"CITEREFRobert_BernasconiTommy_Lee_Lott2000\"] = 1,\n [\"CITEREFRuse2009\"] = 1,\n [\"CITEREFSchopf2000\"] = 1,\n [\"CITEREFSecord2000\"] = 1,\n [\"CITEREFSober2011\"] = 1,\n [\"CITEREFSpencer1864\"] = 1,\n [\"CITEREFWallace1855\"] = 1,\n [\"CITEREFWesley_R._Elsberry1996\"] = 1,\n [\"CITEREFWilberforce\"] = 1,\n [\"CITEREFvan_Wyhe2007\"] = 1,\n [\"CITEREFvan_Wyhe2008\"] = 1,\n [\"CITEREFvan_Wyhe2009\"] = 1,\n}\ntemplate_list = table#1 {\n [\"!\"] = 1,\n [\"Authority control\"] = 1,\n [\"Citation\"] = 89,\n [\"Cite book\"] = 7,\n [\"Cite journal\"] = 2,\n [\"Cite news\"] = 1,\n [\"Cite web\"] = 29,\n [\"Commons category\"] = 1,\n [\"Darwin\"] = 1,\n [\"Evolution\"] = 1,\n [\"Evolutionary biology\"] = 1,\n [\"Featured article\"] = 1,\n [\"Harvnb\"] = 208,\n [\"History of biology\"] = 1,\n [\"In Our Time\"] = 1,\n [\"Infobox book\"] = 1,\n [\"Library resources box\"] = 1,\n [\"Librivox book\"] = 1,\n [\"Main\"] = 1,\n [\"Natural history\"] = 1,\n [\"Portal\"] = 1,\n [\"Pp-move\"] = 1,\n [\"Redirect2\"] = 1,\n [\"Refbegin\"] = 1,\n [\"Refend\"] = 1,\n [\"Reflist\"] = 1,\n [\"See also\"] = 5,\n [\"Sfn\"] = 7,\n [\"Short description\"] = 1,\n [\"StandardEbooks\"] = 1,\n [\"Use British English\"] = 1,\n [\"Use dmy dates\"] = 1,\n [\"Webarchive\"] = 2,\n [\"Wikisource\"] = 1,\n}\narticle_whitelist = table#1 {\n}\nciteref_patterns = table#1 {\n}\n","limitreport-profile":[["?","280","19.7"],["recursiveClone 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Species","url":"https:\/\/en.wikipedia.org\/wiki\/On_the_Origin_of_Species","sameAs":"http:\/\/www.wikidata.org\/entity\/Q20124","mainEntity":"http:\/\/www.wikidata.org\/entity\/Q20124","author":{"@type":"Organization","name":"Contributors to Wikimedia projects"},"publisher":{"@type":"Organization","name":"Wikimedia Foundation, Inc.","logo":{"@type":"ImageObject","url":"https:\/\/www.wikimedia.org\/static\/images\/wmf-hor-googpub.png"}},"datePublished":"2001-03-25T23:03:16Z","dateModified":"2025-01-27T18:58:25Z","image":"https:\/\/upload.wikimedia.org\/wikipedia\/commons\/c\/cd\/Origin_of_Species_title_page.jpg","headline":"treatise on natural selection, written by Charles Darwin"}</script> </body> </html>