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<ul id="toc-History_and_significance-sublist" class="vector-toc-list"> <li id="toc-Dating_the_Cambrian" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Dating_the_Cambrian"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1</span> <span>Dating the Cambrian</span> </div> </a> <ul id="toc-Dating_the_Cambrian-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Body_fossils" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Body_fossils"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2</span> <span>Body fossils</span> </div> </a> <ul id="toc-Body_fossils-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Trace_fossils" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Trace_fossils"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.3</span> <span>Trace fossils</span> </div> </a> <ul id="toc-Trace_fossils-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Geochemical_observations" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Geochemical_observations"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.4</span> <span>Geochemical observations</span> </div> </a> <ul id="toc-Geochemical_observations-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Phylogenetic_techniques" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Phylogenetic_techniques"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.5</span> <span>Phylogenetic techniques</span> </div> </a> <ul id="toc-Phylogenetic_techniques-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Explanation_of_key_scientific_terms" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Explanation_of_key_scientific_terms"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Explanation of key scientific terms</span> </div> </a> <button aria-controls="toc-Explanation_of_key_scientific_terms-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Explanation of key scientific terms subsection</span> </button> <ul id="toc-Explanation_of_key_scientific_terms-sublist" class="vector-toc-list"> <li id="toc-Phylum" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Phylum"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>Phylum</span> </div> </a> <ul id="toc-Phylum-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Stem_group" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Stem_group"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>Stem group</span> </div> </a> <ul id="toc-Stem_group-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Triploblastic" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Triploblastic"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3</span> <span>Triploblastic</span> </div> </a> <ul id="toc-Triploblastic-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Bilaterian" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Bilaterian"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.4</span> <span>Bilaterian</span> </div> </a> <ul id="toc-Bilaterian-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Coelomate" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Coelomate"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.5</span> <span>Coelomate</span> </div> </a> <ul id="toc-Coelomate-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Precambrian_life" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Precambrian_life"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Precambrian life</span> </div> </a> <button aria-controls="toc-Precambrian_life-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Precambrian life subsection</span> </button> <ul id="toc-Precambrian_life-sublist" class="vector-toc-list"> <li id="toc-Evidence_of_animals_around_1_billion_years_ago" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Evidence_of_animals_around_1_billion_years_ago"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>Evidence of animals around 1 billion years ago</span> </div> </a> <ul id="toc-Evidence_of_animals_around_1_billion_years_ago-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Ediacaran_organisms" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Ediacaran_organisms"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Ediacaran organisms</span> </div> </a> <ul id="toc-Ediacaran_organisms-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Cambrian_life" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Cambrian_life"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Cambrian life</span> </div> </a> <button aria-controls="toc-Cambrian_life-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Cambrian life subsection</span> </button> <ul id="toc-Cambrian_life-sublist" class="vector-toc-list"> <li id="toc-Trace_fossils_2" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Trace_fossils_2"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.1</span> <span>Trace fossils</span> </div> </a> <ul id="toc-Trace_fossils_2-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Small_shelly_fauna" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Small_shelly_fauna"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.2</span> <span>Small shelly fauna</span> </div> </a> <ul id="toc-Small_shelly_fauna-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Cnidarians" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cnidarians"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.3</span> <span>Cnidarians</span> </div> </a> <ul id="toc-Cnidarians-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Trilobites" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Trilobites"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.4</span> <span>Trilobites</span> </div> </a> <ul id="toc-Trilobites-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Crustaceans" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Crustaceans"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.5</span> <span>Crustaceans</span> </div> </a> <ul id="toc-Crustaceans-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Echinoderms" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Echinoderms"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.6</span> <span>Echinoderms</span> </div> </a> <ul id="toc-Echinoderms-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Burrowing" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Burrowing"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.7</span> <span>Burrowing</span> </div> </a> <ul id="toc-Burrowing-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Skeletonisation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Skeletonisation"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.8</span> <span>Skeletonisation</span> </div> </a> <ul id="toc-Skeletonisation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Burgess_Shale_type_faunas" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Burgess_Shale_type_faunas"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.9</span> <span>Burgess Shale type faunas</span> </div> </a> <ul id="toc-Burgess_Shale_type_faunas-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Stages" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Stages"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Stages</span> </div> </a> <ul id="toc-Stages-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Validity" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Validity"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Validity</span> </div> </a> <button aria-controls="toc-Validity-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Validity subsection</span> </button> <ul id="toc-Validity-sublist" class="vector-toc-list"> <li id="toc-The_Cambrian_explosion_as_survivorship_bias" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#The_Cambrian_explosion_as_survivorship_bias"> <div class="vector-toc-text"> <span class="vector-toc-numb">6.1</span> <span>The Cambrian explosion as survivorship bias</span> </div> </a> <ul id="toc-The_Cambrian_explosion_as_survivorship_bias-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Possible_causes" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Possible_causes"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Possible causes</span> </div> </a> <button aria-controls="toc-Possible_causes-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span class="vector-icon mw-ui-icon-wikimedia-expand"></span> <span>Toggle Possible causes subsection</span> </button> <ul id="toc-Possible_causes-sublist" class="vector-toc-list"> <li id="toc-Changes_in_the_environment" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Changes_in_the_environment"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.1</span> <span>Changes in the environment</span> </div> </a> <ul id="toc-Changes_in_the_environment-sublist" class="vector-toc-list"> <li id="toc-Increase_in_oxygen_levels" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Increase_in_oxygen_levels"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.1.1</span> <span>Increase in oxygen levels</span> </div> </a> <ul id="toc-Increase_in_oxygen_levels-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Ozone_formation" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Ozone_formation"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.1.2</span> <span>Ozone formation</span> </div> </a> <ul id="toc-Ozone_formation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Snowball_Earth" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Snowball_Earth"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.1.3</span> <span>Snowball Earth</span> </div> </a> <ul id="toc-Snowball_Earth-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Increase_in_the_calcium_concentration_of_the_Cambrian_seawater" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Increase_in_the_calcium_concentration_of_the_Cambrian_seawater"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.1.4</span> <span>Increase in the calcium concentration of the Cambrian seawater</span> </div> </a> <ul id="toc-Increase_in_the_calcium_concentration_of_the_Cambrian_seawater-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Developmental_explanations" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Developmental_explanations"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.2</span> <span>Developmental explanations</span> </div> </a> <ul id="toc-Developmental_explanations-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Ecological_explanations" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Ecological_explanations"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.3</span> <span>Ecological explanations</span> </div> </a> <ul id="toc-Ecological_explanations-sublist" class="vector-toc-list"> <li id="toc-End-Ediacaran_mass_extinction" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#End-Ediacaran_mass_extinction"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.3.1</span> <span>End-Ediacaran mass extinction</span> </div> </a> <ul id="toc-End-Ediacaran_mass_extinction-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Anoxia" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Anoxia"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.3.2</span> <span>Anoxia</span> </div> </a> <ul id="toc-Anoxia-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Increase_in_sensory_and_cognitive_abilities" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Increase_in_sensory_and_cognitive_abilities"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.3.3</span> <span>Increase in sensory and cognitive abilities</span> </div> </a> <ul id="toc-Increase_in_sensory_and_cognitive_abilities-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Arms_races_between_predators_and_prey" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Arms_races_between_predators_and_prey"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.3.4</span> <span>Arms races between predators and prey</span> </div> </a> <ul id="toc-Arms_races_between_predators_and_prey-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Increase_in_size_and_diversity_of_planktonic_animals" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Increase_in_size_and_diversity_of_planktonic_animals"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.3.5</span> <span>Increase in size and diversity of planktonic animals</span> </div> </a> <ul id="toc-Increase_in_size_and_diversity_of_planktonic_animals-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Ecosystem_engineering" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Ecosystem_engineering"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.4</span> <span>Ecosystem engineering</span> </div> </a> <ul id="toc-Ecosystem_engineering-sublist" class="vector-toc-list"> <li id="toc-Burrowing_2" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Burrowing_2"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.4.1</span> <span>Burrowing</span> </div> </a> <ul id="toc-Burrowing_2-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Complexity_threshold" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Complexity_threshold"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.5</span> <span>Complexity threshold</span> </div> </a> <ul id="toc-Complexity_threshold-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Relationship_with_the_Great_Ordovician_Biodiversification_Event" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Relationship_with_the_Great_Ordovician_Biodiversification_Event"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>Relationship with the Great Ordovician Biodiversification Event</span> </div> </a> <ul id="toc-Relationship_with_the_Great_Ordovician_Biodiversification_Event-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Uniqueness_of_the_early_Cambrian_biodiversification" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Uniqueness_of_the_early_Cambrian_biodiversification"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>Uniqueness of the early Cambrian biodiversification</span> </div> </a> <ul id="toc-Uniqueness_of_the_early_Cambrian_biodiversification-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"> <span class="vector-toc-numb">10</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Footnotes" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Footnotes"> <div class="vector-toc-text"> <span class="vector-toc-numb">11</span> <span>Footnotes</span> </div> </a> <ul id="toc-Footnotes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">12</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Further_reading" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> <span class="vector-toc-numb">13</span> <span>Further reading</span> </div> </a> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> <span class="vector-toc-numb">14</span> <span>External links</span> </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" title="Table of Contents" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" ><span class="vector-icon mw-ui-icon-listBullet mw-ui-icon-wikimedia-listBullet"></span> <span class="vector-dropdown-label-text">Toggle the table of contents</span> </label> <div class="vector-dropdown-content"> <div id="vector-page-titlebar-toc-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Cambrian explosion</span></h1> <div id="p-lang-btn" class="vector-dropdown mw-portlet mw-portlet-lang" > <input type="checkbox" id="p-lang-btn-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-p-lang-btn" class="vector-dropdown-checkbox mw-interlanguage-selector" aria-label="Go to an article in another language. Available in 48 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-48" aria-hidden="true" ><span class="vector-icon mw-ui-icon-language-progressive mw-ui-icon-wikimedia-language-progressive"></span> <span class="vector-dropdown-label-text">48 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-af badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://af.wikipedia.org/wiki/Kambriese_ontploffing" title="Kambriese ontploffing – Afrikaans" lang="af" hreflang="af" data-title="Kambriese ontploffing" data-language-autonym="Afrikaans" data-language-local-name="Afrikaans" class="interlanguage-link-target"><span>Afrikaans</span></a></li><li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%A7%D9%84%D8%A7%D9%86%D9%81%D8%AC%D8%A7%D8%B1_%D8%A7%D9%84%D9%83%D9%85%D8%A8%D8%B1%D9%8A" title="الانفجار الكمبري – Arabic" lang="ar" hreflang="ar" data-title="الانفجار الكمبري" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-az mw-list-item"><a href="https://az.wikipedia.org/wiki/Kembri_partlay%C4%B1%C5%9F%C4%B1" title="Kembri partlayışı – Azerbaijani" lang="az" hreflang="az" data-title="Kembri partlayışı" data-language-autonym="Azərbaycanca" data-language-local-name="Azerbaijani" class="interlanguage-link-target"><span>Azərbaycanca</span></a></li><li class="interlanguage-link interwiki-zh-min-nan mw-list-item"><a href="https://zh-min-nan.wikipedia.org/wiki/Cambria_po%CC%8Dk-hoat" title="Cambria po̍k-hoat – Minnan" lang="nan" hreflang="nan" data-title="Cambria po̍k-hoat" data-language-autonym="閩南語 / Bân-lâm-gú" data-language-local-name="Minnan" class="interlanguage-link-target"><span>閩南語 / Bân-lâm-gú</span></a></li><li class="interlanguage-link interwiki-be-x-old mw-list-item"><a href="https://be-tarask.wikipedia.org/wiki/%D0%9A%D0%B5%D0%BC%D0%B1%D1%80%D1%8B%D0%B9%D1%81%D0%BA%D1%96_%D0%B2%D1%8B%D0%B1%D1%83%D1%85" title="Кембрыйскі выбух – Belarusian (Taraškievica orthography)" lang="be-tarask" hreflang="be-tarask" data-title="Кембрыйскі выбух" data-language-autonym="Беларуская (тарашкевіца)" data-language-local-name="Belarusian (Taraškievica orthography)" class="interlanguage-link-target"><span>Беларуская (тарашкевіца)</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%9A%D0%B0%D0%BC%D0%B1%D1%80%D0%B8%D0%B9%D1%81%D0%BA%D0%B8_%D0%B2%D0%B7%D1%80%D0%B8%D0%B2" title="Камбрийски взрив – Bulgarian" lang="bg" hreflang="bg" data-title="Камбрийски взрив" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target"><span>Български</span></a></li><li class="interlanguage-link interwiki-br mw-list-item"><a href="https://br.wikipedia.org/wiki/Tarzhadenn_gambrian" title="Tarzhadenn gambrian – Breton" lang="br" hreflang="br" data-title="Tarzhadenn gambrian" data-language-autonym="Brezhoneg" data-language-local-name="Breton" class="interlanguage-link-target"><span>Brezhoneg</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Explosi%C3%B3_cambriana" title="Explosió cambriana – Catalan" lang="ca" hreflang="ca" data-title="Explosió cambriana" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-ceb mw-list-item"><a href="https://ceb.wikipedia.org/wiki/Pagbutong_Cambrian" title="Pagbutong Cambrian – Cebuano" lang="ceb" hreflang="ceb" data-title="Pagbutong Cambrian" data-language-autonym="Cebuano" data-language-local-name="Cebuano" class="interlanguage-link-target"><span>Cebuano</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Kambrick%C3%A1_exploze" title="Kambrická exploze – Czech" lang="cs" hreflang="cs" data-title="Kambrická exploze" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Kambriske_Eksplosion" title="Kambriske Eksplosion – Danish" lang="da" hreflang="da" data-title="Kambriske Eksplosion" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target"><span>Dansk</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Kambrische_Explosion" title="Kambrische Explosion – German" lang="de" hreflang="de" data-title="Kambrische Explosion" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%9A%CE%AC%CE%BC%CE%B2%CF%81%CE%B9%CE%B1_%CE%AD%CE%BA%CF%81%CE%B7%CE%BE%CE%B7" title="Κάμβρια έκρηξη – Greek" lang="el" hreflang="el" data-title="Κάμβρια έκρηξη" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target"><span>Ελληνικά</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Explosi%C3%B3n_c%C3%A1mbrica" title="Explosión cámbrica – Spanish" lang="es" hreflang="es" data-title="Explosión cámbrica" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Kanbriarreko_leherketa" title="Kanbriarreko leherketa – Basque" lang="eu" hreflang="eu" data-title="Kanbriarreko leherketa" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target"><span>Euskara</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%A7%D9%86%D9%81%D8%AC%D8%A7%D8%B1_%DA%A9%D8%A7%D9%85%D8%A8%D8%B1%DB%8C%D9%86" title="انفجار کامبرین – Persian" lang="fa" hreflang="fa" data-title="انفجار کامبرین" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Explosion_cambrienne" title="Explosion cambrienne – French" lang="fr" hreflang="fr" data-title="Explosion cambrienne" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Explosi%C3%B3n_c%C3%A1mbrica" title="Explosión cámbrica – Galician" lang="gl" hreflang="gl" data-title="Explosión cámbrica" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EC%BA%84%EB%B8%8C%EB%A6%AC%EC%95%84%EA%B8%B0_%ED%8F%AD%EB%B0%9C" title="캄브리아기 폭발 – Korean" lang="ko" hreflang="ko" data-title="캄브리아기 폭발" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-hi mw-list-item"><a href="https://hi.wikipedia.org/wiki/%E0%A4%95%E0%A5%88%E0%A4%AE%E0%A5%8D%E0%A4%AC%E0%A5%8D%E0%A4%B0%E0%A4%BF%E0%A4%AF%E0%A4%BE%E0%A4%88_%E0%A4%B5%E0%A4%BF%E0%A4%B8%E0%A5%8D%E0%A4%A4%E0%A4%BE%E0%A4%B0" title="कैम्ब्रियाई विस्तार – Hindi" lang="hi" hreflang="hi" data-title="कैम्ब्रियाई विस्तार" data-language-autonym="हिन्दी" data-language-local-name="Hindi" class="interlanguage-link-target"><span>हिन्दी</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Letusan_Kambrium" title="Letusan Kambrium – Indonesian" lang="id" hreflang="id" data-title="Letusan Kambrium" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target"><span>Bahasa Indonesia</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Esplosione_cambriana" title="Esplosione cambriana – Italian" lang="it" hreflang="it" data-title="Esplosione cambriana" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%94%D7%A4%D7%99%D7%A6%D7%95%D7%A5_%D7%94%D7%A7%D7%9E%D7%91%D7%A8%D7%99%D7%95%D7%A0%D7%99" title="הפיצוץ הקמבריוני – Hebrew" lang="he" hreflang="he" data-title="הפיצוץ הקמבריוני" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/Kambro_sprogimas" title="Kambro sprogimas – Lithuanian" lang="lt" hreflang="lt" data-title="Kambro sprogimas" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target"><span>Lietuvių</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Kambriumi_robban%C3%A1s" title="Kambriumi robbanás – Hungarian" lang="hu" hreflang="hu" data-title="Kambriumi robbanás" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-ml mw-list-item"><a href="https://ml.wikipedia.org/wiki/%E0%B4%95%E0%B5%87%E0%B4%82%E0%B4%AC%E0%B5%8D%E0%B4%B0%E0%B4%BF%E0%B4%AF%E0%B5%BB_%E0%B4%B5%E0%B4%BF%E0%B4%B8%E0%B5%8D%E0%B4%AB%E0%B5%8B%E0%B4%9F%E0%B4%A8%E0%B4%82" title="കേംബ്രിയൻ വിസ്ഫോടനം – Malayalam" lang="ml" hreflang="ml" data-title="കേംബ്രിയൻ വിസ്ഫോടനം" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target"><span>മലയാളം</span></a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Letupan_Kambria" title="Letupan Kambria – Malay" lang="ms" hreflang="ms" data-title="Letupan Kambria" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target"><span>Bahasa Melayu</span></a></li><li class="interlanguage-link interwiki-mn mw-list-item"><a href="https://mn.wikipedia.org/wiki/%D0%9A%D0%B5%D0%BC%D0%B1%D1%80%D0%B8%D0%B9%D0%BD_%D1%82%D1%8D%D1%81%D1%80%D1%8D%D0%BB%D1%82" title="Кембрийн тэсрэлт – Mongolian" lang="mn" hreflang="mn" data-title="Кембрийн тэсрэлт" data-language-autonym="Монгол" data-language-local-name="Mongolian" class="interlanguage-link-target"><span>Монгол</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Cambrische_explosie" title="Cambrische explosie – Dutch" lang="nl" hreflang="nl" data-title="Cambrische explosie" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E3%82%AB%E3%83%B3%E3%83%96%E3%83%AA%E3%82%A2%E7%88%86%E7%99%BA" title="カンブリア爆発 – Japanese" lang="ja" hreflang="ja" data-title="カンブリア爆発" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target"><span>日本語</span></a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Den_kambriske_eksplosjon" title="Den kambriske eksplosjon – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Den kambriske eksplosjon" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target"><span>Norsk bokmål</span></a></li><li class="interlanguage-link interwiki-nn mw-list-item"><a href="https://nn.wikipedia.org/wiki/Den_kambriske_eksplosjonen" title="Den kambriske eksplosjonen – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="Den kambriske eksplosjonen" data-language-autonym="Norsk nynorsk" data-language-local-name="Norwegian Nynorsk" class="interlanguage-link-target"><span>Norsk nynorsk</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Eksplozja_kambryjska" title="Eksplozja kambryjska – Polish" lang="pl" hreflang="pl" data-title="Eksplozja kambryjska" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target"><span>Polski</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Explos%C3%A3o_Cambriana" title="Explosão Cambriana – Portuguese" lang="pt" hreflang="pt" data-title="Explosão Cambriana" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/Explozia_cambrian%C4%83" title="Explozia cambriană – Romanian" lang="ro" hreflang="ro" data-title="Explozia cambriană" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target"><span>Română</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%9A%D0%B5%D0%BC%D0%B1%D1%80%D0%B8%D0%B9%D1%81%D0%BA%D0%B8%D0%B9_%D0%B2%D0%B7%D1%80%D1%8B%D0%B2" title="Кембрийский взрыв – Russian" lang="ru" hreflang="ru" data-title="Кембрийский взрыв" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Cambrian_explosion" title="Cambrian explosion – Simple English" lang="en-simple" hreflang="en-simple" data-title="Cambrian explosion" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target"><span>Simple English</span></a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/Kambrick%C3%A1_expl%C3%B3zia" title="Kambrická explózia – Slovak" lang="sk" hreflang="sk" data-title="Kambrická explózia" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target"><span>Slovenčina</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%9A%D0%B0%D0%BC%D0%B1%D1%80%D0%B8%D1%98%D1%83%D0%BC%D1%81%D0%BA%D0%B0_%D0%B5%D0%BA%D1%81%D0%BF%D0%BB%D0%BE%D0%B7%D0%B8%D1%98%D0%B0" title="Камбријумска експлозија – Serbian" lang="sr" hreflang="sr" data-title="Камбријумска експлозија" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target"><span>Српски / srpski</span></a></li><li class="interlanguage-link interwiki-sh mw-list-item"><a href="https://sh.wikipedia.org/wiki/Kambrijska_eksplozija" title="Kambrijska eksplozija – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Kambrijska eksplozija" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target"><span>Srpskohrvatski / српскохрватски</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Kambrikauden_lajir%C3%A4j%C3%A4hdys" title="Kambrikauden lajiräjähdys – Finnish" lang="fi" hreflang="fi" data-title="Kambrikauden lajiräjähdys" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Kambriska_explosionen" title="Kambriska explosionen – Swedish" lang="sv" hreflang="sv" data-title="Kambriska explosionen" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/Kambriyen_Patlamas%C4%B1" title="Kambriyen Patlaması – Turkish" lang="tr" hreflang="tr" data-title="Kambriyen Patlaması" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%9A%D0%B5%D0%BC%D0%B1%D1%80%D1%96%D0%B9%D1%81%D1%8C%D0%BA%D0%B8%D0%B9_%D0%B2%D0%B8%D0%B1%D1%83%D1%85" title="Кембрійський вибух – Ukrainian" lang="uk" hreflang="uk" data-title="Кембрійський вибух" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-ur mw-list-item"><a href="https://ur.wikipedia.org/wiki/%DA%A9%DB%8C%D9%85%D8%A8%D8%B1%DB%8C%D9%86_%D8%A7%DB%8C%DA%A9%D9%BE%D9%84%D9%88%DA%98%D9%86" title="کیمبرین ایکپلوژن – Urdu" lang="ur" hreflang="ur" data-title="کیمبرین ایکپلوژن" data-language-autonym="اردو" data-language-local-name="Urdu" class="interlanguage-link-target"><span>اردو</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/S%E1%BB%B1_b%C3%B9ng_n%E1%BB%95_k%E1%BB%B7_Cambri" title="Sự bùng nổ kỷ Cambri – Vietnamese" lang="vi" hreflang="vi" data-title="Sự bùng nổ kỷ Cambri" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target"><span>Tiếng Việt</span></a></li><li class="interlanguage-link interwiki-war mw-list-item"><a href="https://war.wikipedia.org/wiki/Cambrian_explosion" title="Cambrian explosion – Waray" lang="war" hreflang="war" data-title="Cambrian explosion" data-language-autonym="Winaray" data-language-local-name="Waray" class="interlanguage-link-target"><span>Winaray</span></a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E5%AF%92%E6%AD%A6%E7%BA%AA%E5%A4%A7%E7%88%86%E5%8F%91" title="寒武纪大爆发 – Chinese" lang="zh" hreflang="zh" data-title="寒武纪大爆发" data-language-autonym="中文" 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.mw-parser-output .sidebar:not(.notheme) .sidebar-title-with-pretitle{background:transparent!important}html.skin-theme-clientpref-os .mw-parser-output .sidebar:not(.notheme) .sidebar-title-with-pretitle a{color:var(--color-progressive)!important}}@media print{body.ns-0 .mw-parser-output .sidebar{display:none!important}}</style><table class="sidebar sidebar-collapse nomobile nowraplinks" style="width: 22em;"><tbody><tr><td class="sidebar-pretitle"><i>Part of a series on</i></td></tr><tr><th class="sidebar-title-with-pretitle" style="font-size: 115%; background: rgb(127,160,86); padding: 0.4em;">The <a class="mw-selflink selflink">Cambrian explosion</a></th></tr><tr><td class="sidebar-image"><figure class="mw-halign-center" typeof="mw:File"><a href="/wiki/File:Cambrian_explosion_taskforce_logo.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b0/Cambrian_explosion_taskforce_logo.svg/90px-Cambrian_explosion_taskforce_logo.svg.png" decoding="async" width="90" height="55" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b0/Cambrian_explosion_taskforce_logo.svg/135px-Cambrian_explosion_taskforce_logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b0/Cambrian_explosion_taskforce_logo.svg/180px-Cambrian_explosion_taskforce_logo.svg.png 2x" data-file-width="150" data-file-height="92" /></a><figcaption></figcaption></figure></td></tr><tr><td class="sidebar-content" style="text-align: left;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="border-top: 1px #aaa solid;;color: var(--color-base)">Fossil localities</div><div class="sidebar-list-content mw-collapsible-content" style="text-align: left;"> <ul><li><a href="/wiki/Burgess_Shale" title="Burgess Shale">Burgess Shale</a></li> <li><a href="/wiki/Maotianshan_Shales" title="Maotianshan Shales">Chengjiang</a></li> <li><a href="/wiki/Sirius_Passet" title="Sirius Passet">Sirius Passet</a></li> <li><a href="/wiki/Doushantuo_Formation" title="Doushantuo Formation">Doushantuo</a></li></ul></div></div></td> </tr><tr><td class="sidebar-content" style="text-align: left;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="border-top: 1px #aaa solid;;color: var(--color-base)">Key organisms</div><div class="sidebar-list-content mw-collapsible-content" style="text-align: left;"> <dl><dt><a href="/wiki/Ediacaran_biota" title="Ediacaran biota">Ediacaran biota</a></dt></dl> <ul><li><i><a href="/wiki/Dickinsonia" title="Dickinsonia">Dickinsonia</a></i></li> <li><i><a href="/wiki/Kimberella" title="Kimberella">Kimberella</a></i> <ul><li><i><a href="/wiki/Kimberichnus" title="Kimberichnus">Kimberichnus</a></i></li></ul></li> <li><i><a href="/wiki/Vernanimalcula" title="Vernanimalcula">Vernanimalcula</a></i></li></ul> <dl><dt><a href="/wiki/Burgess_Shale_type_fauna" class="mw-redirect" title="Burgess Shale type fauna">Burgess-type</a></dt></dl> <ul><li><i><a href="/wiki/Marrella" title="Marrella">Marrella</a></i></li> <li><a href="/wiki/Radiodont" class="mw-redirect" title="Radiodont">Radiodonts</a></li> <li><a href="/wiki/Halwaxiida" title="Halwaxiida">Halwaxiids</a></li> <li><i><a href="/wiki/Opabinia" title="Opabinia">Opabinia</a></i></li> <li><i><a href="/wiki/Odontogriphus" title="Odontogriphus">Odontogriphus</a></i></li></ul> <dl><dt><a href="/wiki/Small_shelly_fauna" title="Small shelly fauna">Small shelly fauna</a></dt></dl> <ul><li><a href="/wiki/Helcionellid" title="Helcionellid">Helcionellids</a></li></ul></div></div></td> </tr><tr><td class="sidebar-content" style="text-align: left;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="border-top: 1px #aaa solid;;color: var(--color-base)">Evolutionary concepts</div><div class="sidebar-list-content mw-collapsible-content" style="text-align: left;"> <dl><dt>Trends</dt> <dd></dd></dl> <ul><li><a href="/wiki/Cambrian_substrate_revolution" title="Cambrian substrate revolution">Cambrian substrate revolution</a></li></ul> <dl><dt>Themes</dt> <dd></dd></dl> <ul><li><a href="/wiki/Cladistics" title="Cladistics">Cladistics</a></li> <li><a href="/wiki/Convergent_evolution" title="Convergent evolution">Convergent evolution</a></li> <li>Stem and <a href="/wiki/Crown_group" title="Crown group">crown groups</a></li></ul></div></div></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:CEXNAV" title="Template:CEXNAV"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:CEXNAV" title="Template talk:CEXNAV"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:CEXNAV" title="Special:EditPage/Template:CEXNAV"><abbr title="Edit this template">e</abbr></a></li></ul></div></td></tr></tbody></table> <p>The <b>Cambrian explosion</b> (also known as <b>Cambrian radiation</b><sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> or <b>Cambrian diversification</b>) is an interval of time beginning approximately <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=538.8">538.8</a> million years ago</span> in the <a href="/wiki/Cambrian" title="Cambrian">Cambrian</a> period of the early <a href="/wiki/Paleozoic" title="Paleozoic">Paleozoic</a>, when a sudden <a href="/wiki/Evolutionary_radiation" title="Evolutionary radiation">radiation</a> of <a href="/wiki/Complex_life" class="mw-redirect" title="Complex life">complex life</a> occurred and practically all major <a href="/wiki/Animal" title="Animal">animal</a> <a href="/wiki/Phylum" title="Phylum">phyla</a> started appearing in the <a href="/wiki/Fossil_record" class="mw-redirect" title="Fossil record">fossil record</a>.<sup id="cite_ref-StratChart_2022_2-0" class="reference"><a href="#cite_note-StratChart_2022-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> It lasted for about 13<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-dev.biologists.org_6-0" class="reference"><a href="#cite_note-dev.biologists.org-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Budd_7-0" class="reference"><a href="#cite_note-Budd-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> to 25<sup id="cite_ref-The_Cambrian_conundrum:_early_diver_8-0" class="reference"><a href="#cite_note-The_Cambrian_conundrum:_early_diver-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> million years and resulted in the <a href="/wiki/Divergent_evolution" title="Divergent evolution">divergence</a> of most modern <a href="/wiki/Metazoan" class="mw-redirect" title="Metazoan">metazoan</a> phyla.<sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> The event was accompanied by major diversification in other groups of organisms as well.<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>a<span class="cite-bracket">]</span></a></sup> </p><p>Before early Cambrian diversification,<sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>b<span class="cite-bracket">]</span></a></sup> most organisms were relatively simple, composed of individual cells or small multicellular organisms, occasionally organized into <a href="/wiki/Colony_(biology)" title="Colony (biology)">colonies</a>. As the rate of diversification subsequently accelerated, the variety of life became much more complex and began to resemble that of today.<sup id="cite_ref-Bambach2007_14-0" class="reference"><a href="#cite_note-Bambach2007-14"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> Almost all present-day animal phyla appeared during this period,<sup id="cite_ref-Budd2000_15-0" class="reference"><a href="#cite_note-Budd2000-15"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Budd2003_16-0" class="reference"><a href="#cite_note-Budd2003-16"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> including the <a href="/wiki/Cambrian_chordates" title="Cambrian chordates">earliest chordates</a>.<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p> <table id="Container" role="presentation" class="toccolours searchaux" style="text-align:left;padding:0 0.5em;border-style:solid;float:right;clear:right;margin:0.3em 0 0.8em 1.4em;overflow:hidden;min-width:26.100em"><tbody><tr><td colspan="3" id="Title" style="padding:1em 1em 0 1em"><div style="background-color:#77bb77;padding:0 0.2em 0 0.2em;font-weight:bold;text-align:center">Key Cambrian explosion events</div></td></tr><tr><td colspan="3" id="Navbox" style="padding:0;margin:0 1em 0 0;text-align:right"><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"><span>This box: </span><ul><li class="nv-view"><a href="/wiki/Template:Cambrian_explosion_graphical_timeline" title="Template:Cambrian explosion graphical timeline"><span title="View this template">view</span></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Cambrian_explosion_graphical_timeline" title="Template talk:Cambrian explosion graphical timeline"><span title="Discuss this template">talk</span></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Cambrian_explosion_graphical_timeline" title="Special:EditPage/Template:Cambrian explosion graphical timeline"><span title="Edit this template">edit</span></a></li></ul></div></td></tr><tr style="vertical-align:top"><td id="Scale" style="padding:0;margin:0.7em 0 0.7em 0;width:4.2em;position:relative;float:left;font-size:100%;height:65em;border-right:1px solid #242020"><div style="float:right;position:absolute;right:-1px;top:65.00em;transform:translateY(-50%)"><span style="font-size:90%">−590 —</span></div><div style="float:right;position:absolute;right:-1px;top:61.96em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:58.93em;transform:translateY(-50%)"><span style="font-size:90%">−580 —</span></div><div style="float:right;position:absolute;right:-1px;top:55.89em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:52.85em;transform:translateY(-50%)"><span style="font-size:90%">−570 —</span></div><div style="float:right;position:absolute;right:-1px;top:49.81em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:46.78em;transform:translateY(-50%)"><span style="font-size:90%">−560 —</span></div><div style="float:right;position:absolute;right:-1px;top:43.74em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:40.70em;transform:translateY(-50%)"><span style="font-size:90%">−550 —</span></div><div style="float:right;position:absolute;right:-1px;top:37.66em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:34.63em;transform:translateY(-50%)"><span style="font-size:90%">−540 —</span></div><div style="float:right;position:absolute;right:-1px;top:31.59em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:28.55em;transform:translateY(-50%)"><span style="font-size:90%">−530 —</span></div><div style="float:right;position:absolute;right:-1px;top:25.51em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:22.48em;transform:translateY(-50%)"><span style="font-size:90%">−520 —</span></div><div style="float:right;position:absolute;right:-1px;top:19.44em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:16.40em;transform:translateY(-50%)"><span style="font-size:90%">−510 —</span></div><div style="float:right;position:absolute;right:-1px;top:13.36em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:10.33em;transform:translateY(-50%)"><span style="font-size:90%">−500 —</span></div><div style="float:right;position:absolute;right:-1px;top:7.29em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:4.25em;transform:translateY(-50%)"><span style="font-size:90%">−490 —</span></div><div style="float:right;position:absolute;right:-1px;top:1.21em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div></td><td id="Timeline" class="toccolours notheme" style="padding:0px;margin:0.7em 0 0.7em 0;position:relative;font-size:100%;width:10em;height:65em;float:left;border:none;background-color:#000000"><div class="notheme" style="font-size:100%;background-color:rgb(254,179,66);border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:33.897em;height:31.103em;left:0.000em;width:1.250em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-15.551em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="line-height:10px;display:block;"><b><a href="/wiki/Neoproterozoic" title="Neoproterozoic">N<br />e<br />o<br />p<br />r<br />o<br />t<br />e<br />r<br />o<br />z<br />o<br />i<br />c</a></b></span></div></div><div class="notheme" style="font-size:100%;background-color:rgb(153,192,141);border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:0.000em;height:33.897em;left:0.000em;width:1.250em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-16.949em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="line-height:10px;display:block;"><b><a href="/wiki/Paleozoic" title="Paleozoic">P<br />a<br />l<br />e<br />o<br />z<br />o<br />i<br />c</a></b></span></div></div><div class="notheme" style="font-size:100%;background-color:rgb(254,217,106);border-width:0em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:33.897em;height:31.103em;left:1.310em;width:8.690em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-15.551em;transform:translateY(-50%);z-index:5;pointer-events:initial"><b><a href="/wiki/Ediacaran" title="Ediacaran">Ediacaran</a></b></div></div><div class="notheme" style="font-size:100%;background-color:rgb(127,160,86);border-width:0em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:1.458em;height:32.439em;left:1.310em;width:1.190em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-16.220em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="line-height:10px;display:block;"><b><a href="/wiki/Cambrian" title="Cambrian">C<br />a<br />m<br />b<br />r<br />i<br />a<br />n</a></b></span></div></div><div class="notheme" style="font-size:100%;background-color:rgb(0,146,112);border-width:0em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:0.000em;height:1.458em;left:1.310em;width:8.690em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.729em;transform:translateY(-50%);z-index:5;pointer-events:initial"><b><a href="/wiki/Ordovician" title="Ordovician">Ordovician</a></b></div></div><div class="notheme" style="font-size:100%;background-color:rgb(140,176,108);border-width:0em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:23.084em;height:10.813em;left:2.570em;width:1.180em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-5.407em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="font-size:85%;line-height:8px;display:block;"><a href="/wiki/Terreneuvian" title="Terreneuvian">T<br />e<br />r<br />r<br />e<br />n<br />e<br />u<br />v<br />i<br />a<br />n</a><br /><br /></span></div></div><div class="notheme" style="font-size:100%;background-color:rgb(153,192,120);border-width:0.05em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:15.794em;height:7.190em;left:2.570em;width:1.180em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-3.595em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="font-size:85%;line-height:8px;display:block;"><a href="/wiki/Cambrian_Series_2" title="Cambrian Series 2">S<br />e<br />r<br />i<br />e<br />s<br /><br />2</a></span></div></div><div class="notheme" style="font-size:100%;background-color:rgb(166,207,134);border-width:0.05em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:8.505em;height:7.190em;left:2.570em;width:1.180em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-3.595em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="font-size:85%;line-height:8px;display:block;"><a href="/wiki/Cambrian_Series_3" class="mw-redirect" title="Cambrian Series 3">S<br />e<br />r<br />i<br />e<br />s<br /><br />3</a></span></div></div><div class="notheme" style="font-size:100%;background-color:rgb(179,224,149);border-width:0.05em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:1.458em;height:6.947em;left:2.570em;width:1.180em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-3.473em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="font-size:85%;line-height:8px;display:block;"><a href="/wiki/Furongian" title="Furongian">F<br />u<br />r<br />o<br />n<br />g<br />i<br />a<br />n</a></span></div></div><div class="notheme" style="font-size:100%;background-color:rgb(179,224,149);border-width:0.05em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:1.458em;height:6.947em;left:3.810em;width:1.170em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-3.473em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="line-height:10px;display:block;"><a href="/wiki/Upper_Cambrian" class="mw-redirect" title="Upper Cambrian">U<br />p<br />p<br />e<br />r</a></span></div></div><div class="notheme" style="font-size:100%;background-color:rgb(166,207,134);border-width:0.05em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:8.505em;height:7.190em;left:3.810em;width:1.170em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-3.595em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="line-height:10px;display:block;"><a href="/wiki/Middle_Cambrian" class="mw-redirect" title="Middle Cambrian">M<br />i<br />d<br />d<br />l<br />e</a></span></div></div><div class="notheme" style="font-size:100%;background-color:rgb(140,176,108);border-width:0.05em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:15.794em;height:18.003em;left:3.810em;width:1.170em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-9.001em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="line-height:10px;display:block;"><a href="/wiki/Lower_Cambrian" class="mw-redirect" title="Lower Cambrian">L<br />o<br />w<br />e<br />r</a></span></div></div><div class="notheme" style="font-size:100%;background-color:rgb(153,181,117);border-width:0em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:27.944em;height:5.953em;left:5.060em;width:4.940em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-2.977em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="font-size:90%"><a href="/wiki/Fortunian" title="Fortunian">Fortunian</a><br /></span></div></div><div class="notheme" style="font-size:100%;background-color:rgb(166,186,128);border-width:0.05em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:23.084em;height:4.760em;left:5.060em;width:4.940em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-2.380em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="font-size:90%">"<a href="/wiki/Cambrian_Stage_2" title="Cambrian Stage 2">Stage 2</a>"</span></div></div><div class="notheme" style="font-size:100%;background-color:rgb(166,197,131);border-width:0.05em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:18.832em;height:4.152em;left:5.060em;width:4.940em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-2.076em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="font-size:90%">"<a href="/wiki/Cambrian_Stage_3" title="Cambrian Stage 3">Stage 3</a>"</span></div></div><div class="notheme" style="font-size:100%;background-color:rgb(179,202,142);border-width:0.05em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:15.794em;height:2.937em;left:5.060em;width:4.940em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-1.469em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="font-size:90%">"<a href="/wiki/Cambrian_Stage_4" title="Cambrian Stage 4">Stage 4</a>"</span></div></div><div class="notheme" style="font-size:100%;background-color:rgb(179,212,146);border-width:0.05em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:13.061em;height:2.634em;left:5.060em;width:4.940em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-1.317em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="font-size:90%"><a href="/wiki/Wuliuan" title="Wuliuan">Wuliuan</a></span></div></div><div class="notheme" style="font-size:100%;background-color:rgb(191,217,157);border-width:0.05em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:10.631em;height:2.330em;left:5.060em;width:4.940em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-1.165em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="font-size:90%"><a href="/wiki/Drumian" title="Drumian">Drumian</a></span></div></div><div class="notheme" style="font-size:100%;background-color:rgb(204,223,170);border-width:0.05em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:8.505em;height:2.026em;left:5.060em;width:4.940em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-1.013em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="font-size:90%"><a href="/wiki/Guzhangian" title="Guzhangian">Guzhangian</a></span></div></div><div class="notheme" style="font-size:100%;background-color:rgb(204,235,174);border-width:0.05em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:6.682em;height:1.722em;left:5.060em;width:4.940em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.861em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="font-size:90%"><a href="/wiki/Paibian" title="Paibian">Paibian</a></span></div></div><div class="notheme" style="font-size:100%;background-color:rgb(217,240,187);border-width:0.05em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:3.949em;height:2.634em;left:5.060em;width:4.940em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-1.317em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="font-size:75%"><a href="/wiki/Jiangshanian" title="Jiangshanian">Jiangshanian</a></span></div></div><div class="notheme" style="font-size:100%;background-color:rgb(230,245,201);border-width:0.05em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:1.458em;height:2.391em;left:5.060em;width:4.940em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-1.195em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="font-size:90%">"<a href="/wiki/Cambrian_Stage_10" title="Cambrian Stage 10">Stage 10</a>"</span></div></div><div class="notheme" style="font-size:100%;background-color:black;border-width:0.05em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:1.358em;height:0.000em;left:1.310em;width:8.690em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.000em;transform:translateY(-50%);z-index:5;pointer-events:initial"> </div></div><div class="notheme" style="font-size:100%;background-color:black;border-width:0.05em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:33.797em;height:0.000em;left:0.000em;width:10.000em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.000em;transform:translateY(-50%);z-index:5;pointer-events:initial"> </div></div><div class="notheme" style="font-size:100%;background-color:#FEB757;border-width:0.05em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:33.897em;height:24.321em;left:4.000em;width:5.500em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-12.160em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="line-height:12px;display:block;"><a href="/wiki/Ediacaran_Biota" class="mw-redirect" title="Ediacaran Biota">Ediacaran <br />Biota</a></span></div></div><div class="notheme" style="font-size:100%;border-width:0em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:33.897em;height:2.551em;left:1.310em;width:8.690em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-1.276em;transform:translateY(-50%);z-index:5;pointer-events:initial"><b><span style="color:#FFFFFF;line-height:5px;display:block;">* * * * * * * * * * * * * * * * * * * * * * * * * </span></b></div></div><div class="notheme" style="font-size:100%;border-width:0em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:33.897em;height:4.981em;left:1.310em;width:8.690em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-2.491em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="font-size:85%;line-height:10px;display:block;">Baykonur <br />glaciation</span></div></div><div class="notheme" style="font-size:100%;border-width:0em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:37.967em;height:1.215em;left:1.310em;width:8.690em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.607em;transform:translateY(-50%);z-index:5;pointer-events:initial"><b><span style="color:#FFFFFF;line-height:5px;display:block;">* * * * * * * * * * * * * * * * * * * * * * * * * </span></b></div></div><div class="notheme" style="font-size:100%;border-width:0em;border-style:solid none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:60.140em;height:1.822em;left:1.310em;width:8.690em"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-0.911em;transform:translateY(-50%);z-index:5;pointer-events:initial"><b><span style="color:#FFFFFF;line-height:5px;display:block;">* * * * * * * * * * * *<br />* * * * * * * * * * * * * * * * * * * * * * * * *</span></b></div></div></td><td id="Annotations" style="padding:0;margin:0.7em 0 0.7em 0;float:left;position:relative;width:10.200em;height:65em;pointer-events:none"><table role="presentation" style="position:absolute;z-index:15;top:8.505em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div style="z-index:10;font-size:75%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Orsten" title="Orsten">Orsten</a> Fauna</div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:13.061em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div style="z-index:10;font-size:75%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Burgess_Shale" title="Burgess Shale">Burgess Shale</a></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:15.491em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div style="z-index:10;font-size:75%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Kaili_biota" class="mw-redirect" title="Kaili biota">Kaili biota</a></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:16.098em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div style="z-index:10;font-size:75%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><span style="line-height:10px;display:block;"><a href="/wiki/Archaeocyatha" title="Archaeocyatha">Archaeocyatha</a> extinction</span></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:18.528em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div style="z-index:10;font-size:75%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Emu_Bay_shale" class="mw-redirect" title="Emu Bay shale">Emu Bay Shale</a></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:20.654em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div style="z-index:10;font-size:75%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Sirius_Passet" title="Sirius Passet">Sirius Passet</a> biota</div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:21.262em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div style="z-index:10;font-size:75%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Chengjiang_biota" class="mw-redirect" title="Chengjiang biota">Chengjiang</a> biota</div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:23.084em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div style="z-index:10;font-size:75%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><span style="line-height:10px;display:block;">First arthropods with mineralized <a href="/wiki/Carapace" title="Carapace">carapace</a> (<a href="/wiki/Trilobites" class="mw-redirect" title="Trilobites">Trilobites</a>)</span></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:27.944em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div style="z-index:10;font-size:75%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><span style="line-height:10px;display:block;"><a href="/wiki/Small_shelly_fauna" title="Small shelly fauna">SSF</a> diversification, first <a href="/wiki/Brachiopod" title="Brachiopod">brachiopods</a> & <a href="/wiki/Archaeocyatha" title="Archaeocyatha">archaeocyatha</a></span></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:32.196em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div style="z-index:10;font-size:75%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><span style="line-height:10px;display:block;">First <a href="/wiki/Halkieriid" title="Halkieriid">halkieriids</a>, <a href="/wiki/Helcionelloida" title="Helcionelloida">mollusсs</a>, <a href="/wiki/Hyolith" class="mw-redirect" title="Hyolith">hyoliths</a> <a href="/wiki/Small_shelly_fauna" title="Small shelly fauna">SSF</a></span></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:33.897em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div style="z-index:10;font-size:75%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><span style="line-height:10px;display:block;"><i><a href="/wiki/Treptichnus_pedum" class="mw-redirect" title="Treptichnus pedum">Treptichnus pedum</a></i><br />Large negative <a href="/wiki/%CE%9413C" title="Δ13C">δ<sup>13</sup>C</a> peak</span></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:40.701em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div style="z-index:10;font-size:75%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><span style="line-height:10px;display:block;">First <i><a href="/wiki/Cloudina" class="mw-redirect" title="Cloudina">Cloudina</a></i> & <i><a href="/wiki/Namacalathus" title="Namacalathus">Namacalathus</a></i> mineral tubular fossils</span></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:43.738em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div style="z-index:10;font-size:75%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><span style="line-height:10px;display:block;"><a href="/wiki/Mollusc" class="mw-redirect" title="Mollusc">Mollusc</a>-like <i><a href="/wiki/Kimberella" title="Kimberella">Kimberella</a></i> and its trace fossils</span></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:61.355em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div style="z-index:10;font-size:75%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><span style="line-height:10px;display:block;">Gaskiers glaciation<br /><i>Archaeonassa</i>-type trace fossils</span></div></td></tr></tbody></table></td></tr><tr><td id="Caption" colspan="3" style="padding:0;margin:0 0.2em 0.7em 0.2em"><div id="Caption" class="toccolours" style="margin:0 0.5em;border-style:none;clear:both;text-align:center;width:23.000em"></div></td></tr></tbody></table> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="History_and_significance">History and significance</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=1" title="Edit section: History and significance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <table id="Container" role="presentation" class="nomobile toccolours searchaux" style="text-align:left;padding:0 0.5em;border-style:solid;float:right;clear:right;margin:0.3em 0 0.8em 1.4em;overflow:hidden;min-width:29.760em"><tbody><tr><td colspan="3" id="Title" style="padding:1em 1em 0 1em"><div style="background-color:#77bb77;padding:0 0.2em 0 0.2em;font-weight:bold;text-align:center"><a href="/wiki/Timeline_of_the_evolutionary_history_of_life" title="Timeline of the evolutionary history of life"><span style="color:#ffffff;">Life timeline</span></a></div></td></tr><tr><td colspan="3" id="Navbox" style="padding:0;margin:0 1em 0 0;text-align:right"><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"><span>This box: </span><ul><li class="nv-view"><a href="/wiki/Template:Life_timeline" title="Template:Life timeline"><span title="View this template">view</span></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Life_timeline" title="Template talk:Life timeline"><span title="Discuss this template">talk</span></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Life_timeline" title="Special:EditPage/Template:Life timeline"><span title="Edit this template">edit</span></a></li></ul></div></td></tr><tr style="vertical-align:top"><td id="Scale" style="padding:0;margin:0.7em 0 0.7em 0;width:4.2em;position:relative;float:left;font-size:100%;height:50em;border-right:1px solid #242020"><div style="float:right;position:absolute;right:-1px;top:49.67em;transform:translateY(-50%)"><span style="font-size:90%">−4500 —</span></div><div style="float:right;position:absolute;right:-1px;top:48.63em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:47.58em;transform:translateY(-50%)"><span style="font-size:90%">—</span></div><div style="float:right;position:absolute;right:-1px;top:46.53em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:45.47em;transform:translateY(-50%)"><span style="font-size:90%">−4000 —</span></div><div style="float:right;position:absolute;right:-1px;top:44.40em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:43.32em;transform:translateY(-50%)"><span style="font-size:90%">—</span></div><div style="float:right;position:absolute;right:-1px;top:42.23em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:41.14em;transform:translateY(-50%)"><span style="font-size:90%">−3500 —</span></div><div style="float:right;position:absolute;right:-1px;top:40.03em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:38.91em;transform:translateY(-50%)"><span style="font-size:90%">—</span></div><div style="float:right;position:absolute;right:-1px;top:37.78em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:36.65em;transform:translateY(-50%)"><span style="font-size:90%">−3000 —</span></div><div style="float:right;position:absolute;right:-1px;top:35.49em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:34.33em;transform:translateY(-50%)"><span style="font-size:90%">—</span></div><div style="float:right;position:absolute;right:-1px;top:33.15em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:31.96em;transform:translateY(-50%)"><span style="font-size:90%">−2500 —</span></div><div style="float:right;position:absolute;right:-1px;top:30.76em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:29.53em;transform:translateY(-50%)"><span style="font-size:90%">—</span></div><div style="float:right;position:absolute;right:-1px;top:28.29em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:27.04em;transform:translateY(-50%)"><span style="font-size:90%">−2000 —</span></div><div style="float:right;position:absolute;right:-1px;top:25.76em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:24.46em;transform:translateY(-50%)"><span style="font-size:90%">—</span></div><div style="float:right;position:absolute;right:-1px;top:23.14em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:21.79em;transform:translateY(-50%)"><span style="font-size:90%">−1500 —</span></div><div style="float:right;position:absolute;right:-1px;top:20.41em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:19.00em;transform:translateY(-50%)"><span style="font-size:90%">—</span></div><div style="float:right;position:absolute;right:-1px;top:17.56em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:16.08em;transform:translateY(-50%)"><span style="font-size:90%">−1000 —</span></div><div style="float:right;position:absolute;right:-1px;top:14.54em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:12.96em;transform:translateY(-50%)"><span style="font-size:90%">—</span></div><div style="float:right;position:absolute;right:-1px;top:11.30em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:9.56em;transform:translateY(-50%)"><span style="font-size:90%">−500 —</span></div><div style="float:right;position:absolute;right:-1px;top:7.70em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:5.68em;transform:translateY(-50%)"><span style="font-size:90%">—</span></div><div style="float:right;position:absolute;right:-1px;top:3.38em;transform:translateY(-50%)"><span style="font-size:90%">–</span></div><div style="float:right;position:absolute;right:-1px;top:0.00em;transform:translateY(-50%)"><span style="font-size:90%">0 —</span></div></td><td id="Timeline" class="toccolours notheme" style="padding:0px;margin:0.7em 0 0.7em 0;position:relative;font-size:100%;width:13em;height:50em;float:left;border:none"><div class="notheme" style="font-size:100%;background-color:#e5cc7f;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:0.000em;height:50.000em;left:0.910em;width:12.090em"><div style="font-size:100%;position:relative;top:0em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-25.000em;transform:translateY(-50%);z-index:5;pointer-events:initial"> </div></div></div><div class="notheme" style="font-size:100%;background-color:#aebfd1;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:45.470em;height:3.469em;left:1.040em;width:11.960em"><div style="font-size:100%;position:relative;top:0em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-1.735em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="/wiki/Origin_of_water_on_Earth" title="Origin of water on Earth">Water</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#aebfd1;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:0.000em;height:45.470em;left:1.040em;width:0.130em"><div style="font-size:100%;position:relative;top:0em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-22.735em;transform:translateY(-50%);z-index:5;pointer-events:initial"> </div></div></div><div class="notheme" style="font-size:100%;background-color:#b3d9ff;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:41.137em;height:4.333em;left:1.170em;width:11.830em"><div style="font-size:100%;position:relative;top:0em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-2.167em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="/wiki/Unicellular_organism" title="Unicellular organism">Single-celled life</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#b3d9ff;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:0.000em;height:41.137em;left:1.170em;width:0.260em"><div style="font-size:100%;position:relative;top:0em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-20.568em;transform:translateY(-50%);z-index:5;pointer-events:initial"> </div></div></div><div class="notheme" style="font-size:100%;background-color:#99ffff;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:28.044em;height:13.093em;left:1.430em;width:11.570em"><div style="font-size:100%;position:relative;top:0em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-6.546em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="/wiki/Evolution_of_photosynthesis" title="Evolution of photosynthesis">Photosynthesis</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#99ffff;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:0.000em;height:28.044em;left:1.430em;width:0.260em"><div style="font-size:100%;position:relative;top:0em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-14.022em;transform:translateY(-50%);z-index:5;pointer-events:initial"> </div></div></div><div class="notheme" style="font-size:100%;background-color:#b3ffea;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:22.870em;height:5.174em;left:1.690em;width:11.310em"><div style="font-size:100%;position:relative;top:0em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-2.587em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="/wiki/Eukaryote" title="Eukaryote">Eukaryotes</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#b3ffea;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:0.000em;height:22.870em;left:1.690em;width:0.260em"><div style="font-size:100%;position:relative;top:0em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-11.435em;transform:translateY(-50%);z-index:5;pointer-events:initial"> </div></div></div><div class="notheme" style="font-size:100%;background-color:#ccffe7;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:10.141em;height:12.729em;left:1.950em;width:11.050em"><div style="font-size:100%;position:relative;top:0em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-6.365em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="/wiki/Multicellular_organism" title="Multicellular organism">Multicellular life</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#ccffe7;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:0.000em;height:10.141em;left:1.950em;width:0.260em"><div style="font-size:100%;position:relative;top:0em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-5.070em;transform:translateY(-50%);z-index:5;pointer-events:initial"> </div></div></div><div class="notheme" style="font-size:100%;background-color:#77dd00;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:4.443em;height:9.789em;left:2.210em;width:1.170em"><div style="font-size:100%;position:relative;top:0em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;font-size:90%;vertical-align:middle;display:block;bottom:-4.894em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="/wiki/Plant" title="Plant"><span style="display:block; text-align:center; line-height: 0.85em;">P<br />l<br />a<br />n<br />t<br />s</span></a></div></div></div><div class="notheme" style="font-size:100%;background-color:#77dd00;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:0.000em;height:4.443em;left:2.210em;width:0.260em"><div style="font-size:100%;position:relative;top:0em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-2.221em;transform:translateY(-50%);z-index:5;pointer-events:initial"> </div></div></div><div class="notheme" style="font-size:100%;background-color:#ccbbaa;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:4.808em;height:5.333em;left:3.380em;width:9.620em"><div style="font-size:100%;position:relative;top:0.5em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;font-size:90%;vertical-align:middle;display:block;bottom:-2.667em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="/wiki/Arthropod" title="Arthropod">Arthropods</a> <a href="/wiki/Mollusca" title="Mollusca">Molluscs</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#ffff00;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:0.000em;height:4.443em;left:2.470em;width:3.510em"><div style="font-size:100%;position:relative;top:-0.53em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;font-size:90%;vertical-align:middle;display:block;bottom:-2.221em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="/wiki/Flowering_plant#Evolutionary_history" title="Flowering plant">Flowers</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#e6b880;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:2.081em;height:4.435em;left:3.380em;width:5.070em"><div style="font-size:100%;position:relative;top:0.5em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;font-size:90%;vertical-align:middle;display:block;bottom:-2.218em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="/wiki/Dinosaur" title="Dinosaur">Dinosaurs</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#ffcc00;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:0.000em;height:4.808em;left:8.580em;width:4.420em"><div style="font-size:100%;position:relative;top:0em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-2.404em;transform:translateY(-50%);z-index:5;pointer-events:initial"> </div></div></div><div class="notheme" style="font-size:100%;background-color:#ffcc00;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:1.826em;height:2.982em;left:8.450em;width:4.290em"><div style="font-size:100%;position:relative;top:0.15em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;font-size:87%;vertical-align:middle;display:block;bottom:-1.491em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="/wiki/Mammal" title="Mammal">Mammals</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#e6b880;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:0.000em;height:3.298em;left:5.980em;width:2.600em"><div style="font-size:100%;position:relative;top:-0.225em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;font-size:90%;vertical-align:middle;display:block;bottom:-1.649em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="/wiki/Bird" title="Bird">Birds</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#ffbbbb;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:0.000em;height:1.826em;left:8.710em;width:4.030em"><div style="font-size:100%;position:relative;top:0.1em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;font-size:90%;vertical-align:middle;display:block;bottom:-0.913em;transform:translateY(-50%);z-index:5;pointer-events:initial"><a href="/wiki/Primate" title="Primate">Primates</a></div></div></div><div class="notheme" style="font-size:100%;background-color:#993c00;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:45.470em;height:4.530em;left:0.000em;width:0.910em"><div style="font-size:100%;position:relative;top:0em;left:-0.05em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;font-size:100%;vertical-align:middle;display:block;bottom:-2.265em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="display:block; text-align:center; line-height: 0.80em;"><a href="/wiki/Hadean" title="Hadean"><span style="color:#ffffff;">H<br />a<br />d<br />e<br />a<br />n</span></a></span></div></div></div><div class="notheme" style="font-size:100%;background-color:#936a00;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:31.962em;height:13.508em;left:0.000em;width:0.910em"><div style="font-size:100%;position:relative;top:-1.2em;left:-0.05em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;font-size:100%;vertical-align:middle;display:block;bottom:-6.754em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="display:block; text-align:center; line-height: 0.85em;"><a href="/wiki/Archean" title="Archean"><span style="color:#ffffff;"><br /><br /><br />A<br />r<br />c<br />h<br />e<br />a<br />n</span></a></span></div></div></div><div class="notheme" style="font-size:100%;background-color:#f1c309;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:10.141em;height:21.821em;left:0.000em;width:0.910em"><div style="font-size:100%;position:relative;top:0em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;font-size:100%;vertical-align:middle;display:block;bottom:-10.911em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="display:block; text-align:center; line-height: 0.85em;"><a href="/wiki/Proterozoic" title="Proterozoic"><br /><br /><br /><br /><br /><br /><br />P<br />r<br />o<br />t<br />e<br />r<br />o<br />z<br />o<br />i<br />c</a></span></div></div></div><div class="notheme" style="font-size:100%;background-color:#c1e0c1;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:0.000em;height:10.141em;left:0.000em;width:0.910em"><div style="font-size:100%;position:relative;top:0.25em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;font-size:100%;vertical-align:middle;display:block;bottom:-5.070em;transform:translateY(-50%);z-index:5;pointer-events:initial"><span style="display:block; text-align:center; line-height: 0.85em;"><a href="/wiki/Phanerozoic" title="Phanerozoic">P<br />h<br />a<br />n<br />e<br />r<br />o<br />z<br />o<br />i<br />c</a></span></div></div></div><div class="notheme" style="font-size:100%;background-color:#ccbbaa;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:0.000em;height:12.956em;left:12.714em;width:0.286em"><div style="font-size:100%;position:relative;top:0em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-6.478em;transform:translateY(-50%);z-index:5;pointer-events:initial"> </div></div></div><div class="notheme" style="font-size:100%;background-color:#ccbbaa;border-style:none none;position:absolute;text-align:center;margin:0;padding:0;pointer-events:none;top:0.000em;height:4.808em;left:12.714em;width:0.286em"><div style="font-size:100%;position:relative;top:0em;left:0em;pointer-events:none"><div class="notheme" style="position:relative;text-align:center;vertical-align:middle;display:block;bottom:-2.404em;transform:translateY(-50%);z-index:5;pointer-events:initial"> </div></div></div></td><td id="Annotations" style="padding:0;margin:0.7em 0 0.7em 0;float:left;position:relative;width:10.860em;height:50em;pointer-events:none"><table role="presentation" style="position:absolute;z-index:15;top:50.000em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Age_of_Earth" title="Age of Earth">Earth formed</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:48.922em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0.2em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Origin_of_water_on_Earth" title="Origin of water on Earth">Earliest water</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:48.422em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:-0.2em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Last_universal_common_ancestor" title="Last universal common ancestor">LUCA</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:45.470em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0.1em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Earliest_known_life_forms" title="Earliest known life forms">Earliest fossils</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:44.615em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:-0.1em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Late_Heavy_Bombardment" title="Late Heavy Bombardment">LHB meteorites</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:41.137em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Geological_history_of_oxygen" title="Geological history of oxygen">Earliest oxygen</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:35.170em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Pongola_glaciation" title="Pongola glaciation">Pongola glaciation</a>*</div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:30.024em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0.2em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Great_Oxidation_Event" title="Great Oxidation Event">Atmospheric oxygen</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:29.534em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:-0.2em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Huronian_glaciation" title="Huronian glaciation">Huronian glaciation</a>*</div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:27.037em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Evolution_of_sexual_reproduction#Origin_of_sexual_reproduction" title="Evolution of sexual reproduction">Sexual reproduction</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:28.044em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Francevillian_biota" title="Francevillian biota">Multicellular life</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:21.789em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Evolution_of_fungi" title="Evolution of fungi">Earliest fungi</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:14.231em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0.25em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Evolutionary_history_of_plants" title="Evolutionary history of plants">Earliest plants</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:13.085em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0.4em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Otavia" title="Otavia">Earliest animals</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:12.038em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0.2em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Cryogenian" title="Cryogenian">Cryogenian ice age</a>*</div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:10.546em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0.4em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Ediacaran_biota" title="Ediacaran biota">Ediacaran biota</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:10.141em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:-0.25em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a class="mw-selflink selflink">Cambrian explosion</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:8.833em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:-0.1em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Hirnantian_glaciation" title="Hirnantian glaciation">Hirnantian ice age</a>*</div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:7.627em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:-0.1em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Tetrapod" title="Tetrapod">Earliest tetrapods</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:6.517em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:-0.2em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Late_Paleozoic_icehouse" title="Late Paleozoic icehouse">Karoo ice age</a>*</div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:0.887em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:0.1em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Hominoid" class="mw-redirect" title="Hominoid">Earliest hominoid</a></div></div></td></tr></tbody></table><table role="presentation" style="position:absolute;z-index:15;top:0.129em;left:0em;transform:translateY(-50%);padding:0;margin:0;font-size:100%"><tbody><tr><td style="padding:0;text-align:left;vertical-align:middle"><span style="color:var( --color-base, #000)">←</span></td><td style="padding:0;text-align:left;vertical-align:middle"><div class="annot-nudge" style="font-size:100%;float:left;position:relative;top:-0.2em"><div style="z-index:10;font-size:90%;color:var( --color-base, #000);display:block;line-height:105%;bottom:0;pointer-events:initial"><a href="/wiki/Quaternary_glaciation" title="Quaternary glaciation">Quaternary ice age</a>*</div></div></td></tr></tbody></table></td></tr><tr><td id="Caption" colspan="3" style="padding:0;margin:0 0.2em 0.7em 0.2em"><div id="Caption" class="toccolours" style="margin:0 0.5em;border-style:none;clear:both;text-align:center;width:26.000em"><div style="float:left;font-size:85%;">(<a href="/wiki/Myr" class="mw-redirect" title="Myr">million years ago</a>)</div><div style="float:right;font-size:95%;">*<i><a href="/wiki/Timeline_of_glaciation" title="Timeline of glaciation">Ice Ages</a></i></div></div></td></tr></tbody></table> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/History_of_life" title="History of life">History of life</a></div> <p>The seemingly rapid appearance of fossils in the "Primordial Strata" was noted by <a href="/wiki/William_Buckland" title="William Buckland">William Buckland</a> in the 1840s,<sup id="cite_ref-Buckland1841_18-0" class="reference"><a href="#cite_note-Buckland1841-18"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> and in his 1859 book <i><a href="/wiki/On_the_Origin_of_Species" title="On the Origin of Species">On the Origin of Species</a></i>, <a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a> discussed the then-inexplicable lack of earlier fossils as one of the main difficulties for his theory of descent with slow modification through <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a>.<sup id="cite_ref-OriginOfSpecies_19-0" class="reference"><a href="#cite_note-OriginOfSpecies-19"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> The long-running puzzlement about the seemingly-sudden appearance of the Cambrian <a href="/wiki/Fauna" title="Fauna">fauna</a> without evident precursor(s) centers on three key points: whether there really was a mass diversification of complex organisms over a relatively short period during the early Cambrian, what might have caused such rapid change, and what it would imply about the origin of animal life. Interpretation is difficult, owing to a limited supply of evidence based mainly on an incomplete fossil record and chemical signatures remaining in Cambrian rocks. </p><p>The first discovered Cambrian fossils were <a href="/wiki/Trilobites" class="mw-redirect" title="Trilobites">trilobites</a>, described by <a href="/wiki/Edward_Lhuyd" title="Edward Lhuyd">Edward Lhuyd</a>, the curator of <a href="/wiki/Oxford_Museum" class="mw-redirect" title="Oxford Museum">Oxford Museum</a>, in 1698.<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> Although their evolutionary importance was not known, on the basis of their old age, William Buckland (1784–1856) realized that a dramatic step-change in the fossil record had occurred around the base of what we now call the Cambrian.<sup id="cite_ref-Buckland1841_18-1" class="reference"><a href="#cite_note-Buckland1841-18"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> Nineteenth-century geologists such as <a href="/wiki/Adam_Sedgwick" title="Adam Sedgwick">Adam Sedgwick</a> and <a href="/wiki/Roderick_Murchison" title="Roderick Murchison">Roderick Murchison</a> used the fossils for dating rock strata, specifically for establishing the <a href="/wiki/Cambrian" title="Cambrian">Cambrian</a> and <a href="/wiki/Silurian" title="Silurian">Silurian</a> periods.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> By 1859, leading geologists including Roderick Murchison were convinced that what was then called the lowest Silurian stratum showed the origin of life on Earth, though others, including <a href="/wiki/Charles_Lyell" title="Charles Lyell">Charles Lyell</a>, differed. In <i>On the Origin of Species</i>, Darwin considered this sudden appearance of a solitary group of trilobites, with no apparent antecedents, and absent other fossils, to be "undoubtedly of the gravest nature" among the difficulties in his theory of natural selection. He reasoned that earlier seas had swarmed with living creatures, but that their fossils had not been found because of the imperfections of the fossil record.<sup id="cite_ref-OriginOfSpecies_19-1" class="reference"><a href="#cite_note-OriginOfSpecies-19"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> In the sixth edition of his book, he stressed his problem further as:<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> </p> <style data-mw-deduplicate="TemplateStyles:r1244412712">.mw-parser-output .templatequote{overflow:hidden;margin:1em 0;padding:0 32px}.mw-parser-output .templatequotecite{line-height:1.5em;text-align:left;margin-top:0}@media(min-width:500px){.mw-parser-output .templatequotecite{padding-left:1.6em}}</style><blockquote class="templatequote"><p>To the question why we do not find rich fossiliferous deposits belonging to these assumed earliest periods prior to the Cambrian system, I can give no satisfactory answer.</p></blockquote> <p>American paleontologist <a href="/wiki/Charles_Doolittle_Walcott" title="Charles Doolittle Walcott">Charles Walcott</a>, who studied the <a href="/wiki/Fossils_of_the_Burgess_Shale" title="Fossils of the Burgess Shale">Burgess Shale fauna</a>, proposed that an interval of time, the "Lipalian", was not represented in the fossil record or did not preserve fossils, and that the ancestors of the Cambrian animals evolved during this time.<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> </p><p>Earlier fossil evidence has since been found. The earliest claim is that the history of life on Earth goes back <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=3,850">3,850</a> million years</span>:<sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> Rocks of that age at <a href="/wiki/Warrawoona,_Australia" class="mw-redirect" title="Warrawoona, Australia">Warrawoona, Australia</a>, were claimed to contain fossil <a href="/wiki/Stromatolite" title="Stromatolite">stromatolites</a>, stubby pillars formed by colonies of <a href="/wiki/Microorganism" title="Microorganism">microorganisms</a>. Fossils (<i><a href="/wiki/Grypania" title="Grypania">Grypania</a></i>) of more complex <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotic</a> cells, from which all animals, plants and fungi are built, have been found in rocks from <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=1,400">1,400</a> million years ago</span>, in <a href="/wiki/China" title="China">China</a> and <a href="/wiki/Montana" title="Montana">Montana</a>. Rocks dating from <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=580–543">580 to 543</a> million years ago</span> contain fossils of the <a href="/wiki/Ediacara_biota" class="mw-redirect" title="Ediacara biota">Ediacara biota</a>, organisms so large that they are likely multicelled, but very unlike any modern organism.<sup id="cite_ref-CowenHistLife_25-0" class="reference"><a href="#cite_note-CowenHistLife-25"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> In 1948, <a href="/wiki/Preston_Cloud" title="Preston Cloud">Preston Cloud</a> argued that a period of "eruptive" evolution occurred in the Early Cambrian,<sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> but as recently as the 1970s, no sign was seen of how the 'relatively' modern-looking organisms of the Middle and Late <a href="/wiki/Cambrian" title="Cambrian">Cambrian</a> arose.<sup id="cite_ref-CowenHistLife_25-1" class="reference"><a href="#cite_note-CowenHistLife-25"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:20191108_Opabinia_regalis.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/21/20191108_Opabinia_regalis.png/220px-20191108_Opabinia_regalis.png" decoding="async" width="220" height="93" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/21/20191108_Opabinia_regalis.png/330px-20191108_Opabinia_regalis.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/21/20191108_Opabinia_regalis.png/440px-20191108_Opabinia_regalis.png 2x" data-file-width="1900" data-file-height="800" /></a><figcaption><i><a href="/wiki/Opabinia" title="Opabinia">Opabinia</a></i> made the largest single contribution to modern interest in the Cambrian explosion.</figcaption></figure> <p>The intense modern interest in this "Cambrian explosion" was sparked by the work of <a href="/wiki/Harry_B._Whittington" title="Harry B. Whittington">Harry B. Whittington</a> and colleagues, who, in the 1970s, reanalysed many fossils from the Burgess Shale and concluded that several were as complex as, but different from, any living animals.<sup id="cite_ref-Whittington1979Lawn_27-0" class="reference"><a href="#cite_note-Whittington1979Lawn-27"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Whittington1985BurgessShale_28-0" class="reference"><a href="#cite_note-Whittington1985BurgessShale-28"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> The most common organism, <i><a href="/wiki/Marrella" title="Marrella">Marrella</a></i>, was clearly an <a href="/wiki/Arthropod" title="Arthropod">arthropod</a>, but not a member of any known arthropod <a href="/wiki/Class_(biology)" title="Class (biology)">class</a>. Organisms such as the five-eyed <i><a href="/wiki/Opabinia" title="Opabinia">Opabinia</a></i> and spiny slug-like <i><a href="/wiki/Wiwaxia" title="Wiwaxia">Wiwaxia</a></i> were so different from anything else known that Whittington's team assumed they must represent different phyla, seemingly unrelated to anything known today. <a href="/wiki/Stephen_Jay_Gould" title="Stephen Jay Gould">Stephen Jay Gould</a>'s popular 1989 account of this work, <i><a href="/wiki/Wonderful_Life_(book)" title="Wonderful Life (book)">Wonderful Life</a></i>,<sup id="cite_ref-WonderfulLife_29-0" class="reference"><a href="#cite_note-WonderfulLife-29"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> brought the matter into the public eye and raised questions about what the explosion represented. While differing significantly in details, both Whittington and Gould proposed that all modern animal phyla had appeared almost simultaneously in a rather short span of geological period. This view led to the modernization of Darwin's tree of life and the theory of <a href="/wiki/Punctuated_equilibrium" title="Punctuated equilibrium">punctuated equilibrium</a>, which <a href="/wiki/Niles_Eldredge" title="Niles Eldredge">Eldredge</a> and Gould developed in the early 1970s and which views evolution as long intervals of near-stasis "punctuated" by short periods of rapid change.<sup id="cite_ref-Bengtson2004EarlySkeletalFossils_30-0" class="reference"><a href="#cite_note-Bengtson2004EarlySkeletalFossils-30"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p><p>Other analyses, some more recent and some dating back to the 1970s, argue that complex animals similar to modern types evolved well before the start of the Cambrian.<sup id="cite_ref-McNamara1996DatingOriginAnimals_31-0" class="reference"><a href="#cite_note-McNamara1996DatingOriginAnimals-31"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-AwramikStromatoliteDiversityMetazoanAppearance_32-0" class="reference"><a href="#cite_note-AwramikStromatoliteDiversityMetazoanAppearance-32"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FedonkinWaggoner1997KimberellaMollusc_33-0" class="reference"><a href="#cite_note-FedonkinWaggoner1997KimberellaMollusc-33"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Dating_the_Cambrian">Dating the Cambrian</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=2" title="Edit section: Dating the Cambrian"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Radiometric_dating" title="Radiometric dating">Radiometric</a> dates for much of the Cambrian, obtained by analysis of radioactive elements contained within rocks, have only recently become available, and for only a few regions. </p><p>Relative dating (<i>A</i> was before <i>B</i>) is often assumed sufficient for studying processes of evolution, but this, too, has been difficult, because of the problems involved in matching up rocks of the same age across different <a href="/wiki/Continent" title="Continent">continents</a>.<sup id="cite_ref-DatingProblems_34-0" class="reference"><a href="#cite_note-DatingProblems-34"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> </p><p>Therefore, dates or descriptions of sequences of events should be regarded with some caution until better data become available. In 2004, the start of the Cambrian was dated to 542 <a href="/wiki/Year#SI_prefix_multipliers" title="Year">Ma</a>.<sup id="cite_ref-USGS_2007_35-0" class="reference"><a href="#cite_note-USGS_2007-35"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> In 2012, it was revised to 541 Ma<sup id="cite_ref-Gradstein_et-al_2012_36-0" class="reference"><a href="#cite_note-Gradstein_et-al_2012-36"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> then in 2022 it was changed again to 538.8 Ma.<sup id="cite_ref-StratChart_2022_2-1" class="reference"><a href="#cite_note-StratChart_2022-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p><p>Some theory suggest Cambrian explosion occurred during the last stages of <a href="/wiki/Gondwana" title="Gondwana">Gondwanan</a> assembly, which is formed following <a href="/wiki/Rodinia" title="Rodinia">Rodinia</a> splitting, overlapped with the opening of the <a href="/wiki/Iapetus_Ocean" title="Iapetus Ocean">Iapetus Ocean</a> between <a href="/wiki/Laurentia" title="Laurentia">Laurentia</a> and western Gondwana.<sup id="cite_ref-37" class="reference"><a href="#cite_note-37"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-38" class="reference"><a href="#cite_note-38"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> The largest Cambrian faunal province is located around Gondwana, which extended from the low northern latitudes to the high southern latitudes, just short of the South Pole. By the middle and later parts of the Cambrian, continued rifting had sent the paleocontinents of Laurentia, <a href="/wiki/Baltica" title="Baltica">Baltica</a> and <a href="/wiki/Siberia_(continent)" title="Siberia (continent)">Siberia</a> on their separate ways.<sup id="cite_ref-39" class="reference"><a href="#cite_note-39"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Body_fossils">Body fossils</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=3" title="Edit section: Body fossils"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Fossils of organisms' bodies are usually the most informative type of evidence. Fossilization is a rare event, and most fossils are destroyed by <a href="/wiki/Erosion" title="Erosion">erosion</a> or <a href="/wiki/Metamorphism" title="Metamorphism">metamorphism</a> before they can be observed. Hence, the fossil record is very incomplete, increasingly so as earlier times are considered. Despite this, they are often adequate to illustrate the broader patterns of life's history.<sup id="cite_ref-BentonQualityFossilRecord_40-0" class="reference"><a href="#cite_note-BentonQualityFossilRecord-40"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> Also, biases exist in the fossil record: different environments are more favourable to the preservation of different types of organism or parts of organisms.<sup id="cite_ref-Butterfield2003ExceptionalFossilPreservation_41-0" class="reference"><a href="#cite_note-Butterfield2003ExceptionalFossilPreservation-41"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> Further, only the parts of organisms that were already <a href="/wiki/Mineralization_(biology)" class="mw-redirect" title="Mineralization (biology)">mineralised</a> are usually preserved, such as the shells of <a href="/wiki/Mollusca" title="Mollusca">molluscs</a>. Since most animal species are soft-bodied, they decay before they can become fossilised. As a result, although 30-plus phyla of living animals are known, two-thirds have never been found as fossils.<sup id="cite_ref-CowenHistLife_25-2" class="reference"><a href="#cite_note-CowenHistLife-25"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Marrella_(fossil).png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/dd/Marrella_%28fossil%29.png/250px-Marrella_%28fossil%29.png" decoding="async" width="220" height="171" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/dd/Marrella_%28fossil%29.png/330px-Marrella_%28fossil%29.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/dd/Marrella_%28fossil%29.png/500px-Marrella_%28fossil%29.png 2x" data-file-width="516" data-file-height="401" /></a><figcaption> This <i><a href="/wiki/Marrella" title="Marrella">Marrella</a></i> specimen illustrates how clear and detailed the fossils from the <a href="/wiki/Burgess_Shale" title="Burgess Shale">Burgess Shale</a> <a href="/wiki/Lagerst%C3%A4tte" title="Lagerstätte">Lagerstätte</a> actually are as well as the oldest evidence for liquid <a href="/wiki/Blood" title="Blood">blood</a> in an animal.</figcaption></figure> <p>The Cambrian fossil record includes an unusually high number of <a href="/wiki/Lagerst%C3%A4tte" title="Lagerstätte">lagerstätten</a>, which preserve soft tissues. These allow <a href="/wiki/Paleontology" title="Paleontology">paleontologists</a> to examine the internal anatomy of animals, which in other sediments are only represented by shells, spines, claws, etc.—if they are preserved at all. The most significant Cambrian lagerstätten are the early Cambrian <a href="/wiki/Maotianshan_shale" class="mw-redirect" title="Maotianshan shale">Maotianshan shale</a> beds of Chengjiang (<a href="/wiki/Yunnan" title="Yunnan">Yunnan</a>, <a href="/wiki/China" title="China">China</a>) and <a href="/wiki/Sirius_Passet" title="Sirius Passet">Sirius Passet</a> (<a href="/wiki/Greenland" title="Greenland">Greenland</a>),<sup id="cite_ref-ConwayMorris1979_42-0" class="reference"><a href="#cite_note-ConwayMorris1979-42"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> the middle Cambrian <a href="/wiki/Burgess_Shale" title="Burgess Shale">Burgess Shale</a> (<a href="/wiki/British_Columbia" title="British Columbia">British Columbia</a>, <a href="/wiki/Canada" title="Canada">Canada</a>)<sup id="cite_ref-Yochelson1996_43-0" class="reference"><a href="#cite_note-Yochelson1996-43"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> and the late Cambrian <a href="/wiki/Orsten" title="Orsten">Orsten</a> (<a href="/wiki/Sweden" title="Sweden">Sweden</a>) fossil beds. </p><p>While lagerstätten preserve far more than the conventional fossil record, they are far from complete. Because lagerstätten are restricted to a narrow range of environments (where soft-bodied organisms can be preserved very quickly, e.g. by mudslides), most animals are probably not represented; further, the exceptional conditions that create lagerstätten probably do not represent normal living conditions.<sup id="cite_ref-Butterfield2001_44-0" class="reference"><a href="#cite_note-Butterfield2001-44"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> In addition, the known Cambrian lagerstätten are rare and difficult to date, while <a href="/wiki/Precambrian" title="Precambrian">Precambrian</a> lagerstätten have yet to be studied in detail. </p><p>The sparseness of the fossil record means that organisms usually exist long before they are found in the fossil record—this is known as the <a href="/wiki/Signor%E2%80%93Lipps_effect" title="Signor–Lipps effect">Signor–Lipps effect</a>.<sup id="cite_ref-Signor1982_45-0" class="reference"><a href="#cite_note-Signor1982-45"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> </p><p>In 2019, a "stunning" find of lagerstätten, known as the <a href="/wiki/Qingjiang_biota" title="Qingjiang biota">Qingjiang biota</a>, was reported from the <a href="/w/index.php?title=Danshui_river&action=edit&redlink=1" class="new" title="Danshui river (page does not exist)">Danshui river</a> in <a href="/wiki/Hubei" title="Hubei">Hubei</a> province, <a href="/wiki/China" title="China">China</a>. More than 20,000 fossil specimens were collected, including many soft bodied animals such as jellyfish, <a href="/wiki/Sea_anemone" title="Sea anemone">sea anemones</a> and worms, as well as sponges, arthropods and algae. In some specimens the internal body structures were sufficiently preserved that soft tissues, including muscles, gills, mouths, guts and eyes, can be seen. The remains were dated to around 518 Mya and around half of the species identified at the time of reporting were previously unknown.<sup id="cite_ref-46" class="reference"><a href="#cite_note-46"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-47" class="reference"><a href="#cite_note-47"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-48" class="reference"><a href="#cite_note-48"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Trace_fossils">Trace fossils</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=4" title="Edit section: Trace fossils"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:CambrianRusophycus.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a3/CambrianRusophycus.jpg/250px-CambrianRusophycus.jpg" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a3/CambrianRusophycus.jpg/330px-CambrianRusophycus.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/a3/CambrianRusophycus.jpg/500px-CambrianRusophycus.jpg 2x" data-file-width="4000" data-file-height="3000" /></a><figcaption><i>Rusophycus</i> and other trace fossils from the <a href="/wiki/Gog_Group" title="Gog Group">Gog Group</a>, <a href="/wiki/Middle_Cambrian" class="mw-redirect" title="Middle Cambrian">Middle Cambrian</a>, <a href="/wiki/Lake_Louise_(Alberta)" title="Lake Louise (Alberta)">Lake Louise</a>, Alberta, Canada</figcaption></figure> <p><a href="/wiki/Trace_fossil" title="Trace fossil">Trace fossils</a> consist mainly of tracks and burrows, but also include <a href="/wiki/Coprolite" title="Coprolite">coprolites</a> (fossil <a href="/wiki/Feces" title="Feces">feces</a>) and marks left by feeding.<sup id="cite_ref-UCMPWhatIsPaleo_49-0" class="reference"><a href="#cite_note-UCMPWhatIsPaleo-49"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FedonkinGehlingEtAl2007RiseOfAnimals_50-0" class="reference"><a href="#cite_note-FedonkinGehlingEtAl2007RiseOfAnimals-50"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> Trace fossils are particularly significant because they represent a data source that is not limited to animals with easily fossilized hard parts, and reflects organisms' behaviour. Also, many traces date from significantly earlier than the body fossils of animals that are thought to have been capable of making them.<sup id="cite_ref-Seilacher1994_51-0" class="reference"><a href="#cite_note-Seilacher1994-51"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> While exact assignment of trace fossils to their makers is generally impossible, traces may, for example, provide the earliest physical evidence of the appearance of moderately complex animals (comparable to <a href="/wiki/Earthworm" title="Earthworm">earthworms</a>).<sup id="cite_ref-FedonkinGehlingEtAl2007RiseOfAnimals_50-1" class="reference"><a href="#cite_note-FedonkinGehlingEtAl2007RiseOfAnimals-50"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Geochemical_observations">Geochemical observations</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=5" title="Edit section: Geochemical observations"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Early_Cambrian_geochemical_fluctuations" title="Early Cambrian geochemical fluctuations">Early Cambrian geochemical fluctuations</a></div> <p>Several <a href="/wiki/Geochemistry" title="Geochemistry">chemical markers</a> indicate a drastic change in the environment around the start of the Cambrian. The markers are consistent with a mass extinction,<sup id="cite_ref-Knoll1999_52-0" class="reference"><a href="#cite_note-Knoll1999-52"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Amthor2003_53-0" class="reference"><a href="#cite_note-Amthor2003-53"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> or with a massive warming resulting from the release of <a href="/wiki/Methane_clathrate" title="Methane clathrate">methane ice</a>.<sup id="cite_ref-Marshall2006Explaining_54-0" class="reference"><a href="#cite_note-Marshall2006Explaining-54"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> Such changes may reflect a cause of the Cambrian explosion, although they may also have resulted from an increased level of biological activity—a possible result of the explosion.<sup id="cite_ref-Marshall2006Explaining_54-1" class="reference"><a href="#cite_note-Marshall2006Explaining-54"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> Despite these uncertainties, the geochemical evidence helps by making scientists focus on theories that are consistent with at least one of the likely environmental changes. </p> <div class="mw-heading mw-heading3"><h3 id="Phylogenetic_techniques">Phylogenetic techniques</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=6" title="Edit section: Phylogenetic techniques"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Cladistics" title="Cladistics">Cladistics</a> is a technique for working out the "family tree" of a set of organisms. It works by the logic that, if groups B and C have more similarities to each other than either has to group A, then B and C are more closely related to each other than either is to A. Characteristics that are compared may be <a href="/wiki/Anatomy" title="Anatomy">anatomical</a>, such as the presence of a <a href="/wiki/Notochord" title="Notochord">notochord</a>, or <a href="/wiki/Molecular_phylogeny" class="mw-redirect" title="Molecular phylogeny">molecular</a>, by comparing sequences of <a href="/wiki/DNA" title="DNA">DNA</a> or <a href="/wiki/Protein" title="Protein">protein</a>. The result of a successful analysis is a hierarchy of <a href="/wiki/Clade" title="Clade">clades</a>—groups whose members are believed to share a common ancestor. The cladistic technique is sometimes problematic, as some features, such as wings or <a href="/wiki/Evolution_of_the_eye" title="Evolution of the eye">camera eyes</a>, evolved more than once, <a href="/wiki/Convergent_evolution" title="Convergent evolution">convergently</a>—this must be taken into account in analyses. </p><p>From the relationships, it may be possible to constrain the date that lineages first appeared. For instance, if fossils of B or C date to X million years ago and the calculated "family tree" says A was an ancestor of B and C, then A must have evolved more than X million years ago. </p><p>It is also possible to estimate how long ago two living clades diverged—i.e. about how long ago their last common ancestor must have lived—by assuming that DNA <a href="/wiki/Mutation" title="Mutation">mutations</a> accumulate at a constant rate. These "<a href="/wiki/Molecular_clock" title="Molecular clock">molecular clocks</a>", however, are fallible, and provide only a very approximate timing: they are not sufficiently precise and reliable for estimating when the groups that feature in the Cambrian explosion first evolved,<sup id="cite_ref-55" class="reference"><a href="#cite_note-55"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> and estimates produced by different techniques vary by a factor of two.<sup id="cite_ref-PetersonEtAl2005_56-0" class="reference"><a href="#cite_note-PetersonEtAl2005-56"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> However, the clocks can give an indication of branching rate, and when combined with the constraints of the fossil record, recent clocks suggest a sustained period of diversification through the Ediacaran and Cambrian.<sup id="cite_ref-57" class="reference"><a href="#cite_note-57"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Explanation_of_key_scientific_terms">Explanation of key scientific terms</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=7" title="Edit section: Explanation of key scientific terms"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Crown_n_Stem_Groups.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/1a/Crown_n_Stem_Groups.svg/330px-Crown_n_Stem_Groups.svg.png" decoding="async" width="260" height="181" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/1a/Crown_n_Stem_Groups.svg/390px-Crown_n_Stem_Groups.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/1a/Crown_n_Stem_Groups.svg/520px-Crown_n_Stem_Groups.svg.png 2x" data-file-width="479" data-file-height="334" /></a><figcaption><a href="/wiki/Stem_group" class="mw-redirect" title="Stem group">Stem groups</a><sup id="cite_ref-CraskeJefferies1989_58-0" class="reference"><a href="#cite_note-CraskeJefferies1989-58"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup><style data-mw-deduplicate="TemplateStyles:r1184024115">.mw-parser-output .div-col{margin-top:0.3em;column-width:30em}.mw-parser-output .div-col-small{font-size:90%}.mw-parser-output .div-col-rules{column-rule:1px solid #aaa}.mw-parser-output .div-col dl,.mw-parser-output .div-col ol,.mw-parser-output .div-col ul{margin-top:0}.mw-parser-output .div-col li,.mw-parser-output .div-col dd{page-break-inside:avoid;break-inside:avoid-column}</style><div class="div-col"><style data-mw-deduplicate="TemplateStyles:r1126788409">.mw-parser-output .plainlist ol,.mw-parser-output .plainlist ul{line-height:inherit;list-style:none;margin:0;padding:0}.mw-parser-output .plainlist ol li,.mw-parser-output .plainlist ul li{margin-bottom:0}</style><div class="plainlist"><ul><li><style data-mw-deduplicate="TemplateStyles:r981673959">.mw-parser-output .legend{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .legend-color{display:inline-block;min-width:1.25em;height:1.25em;line-height:1.25;margin:1px 0;text-align:center;border:1px solid black;background-color:transparent;color:black}.mw-parser-output .legend-text{}</style><span class="legend nowrap"><span class="legend-color mw-no-invert" style=""><span class="legend-text" style="font-family: monospace, monospace;"><span style="font-size:120%;"><b>—</b></span></span></span> <span class="nowrap">= Lines of descent</span></span></li><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r981673959" /><span class="legend nowrap"><span class="legend-color mw-no-invert" style="border: 1px solid gray;background-color:black; color:white;"> </span> = Basal node</span></li><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r981673959" /><span class="legend nowrap"><span class="legend-color mw-no-invert" style="border: 1px solid gray;background-color:white; color:black;"> </span> = Crown node</span></li><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r981673959" /><span class="legend nowrap"><span class="legend-color mw-no-invert" style="border: 1px solid gray;background-color:#8080ff; color:black;"> </span> = Total group</span></li><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r981673959" /><span class="legend nowrap"><span class="legend-color mw-no-invert" style="border: 1px solid gray;background-color:#faada7; color:black;"> </span> = Crown group</span></li><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r981673959" /><span class="legend nowrap"><span class="legend-color mw-no-invert" style="border: 1px solid gray;background-color:#fdefa4; color:black;"> </span> = Stem group</span></li></ul></div></div></figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Phylum">Phylum</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=8" title="Edit section: Phylum"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A <a href="/wiki/Phylum" title="Phylum">phylum</a> is the highest level in the <a href="/wiki/Linnaean_taxonomy" title="Linnaean taxonomy">Linnaean system for classifying organisms</a>. Phyla can be thought of as groupings of animals based on general body plan.<sup id="cite_ref-59" class="reference"><a href="#cite_note-59"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> Despite the seemingly different external appearances of organisms, they are classified into phyla based on their internal and developmental organizations.<sup id="cite_ref-60" class="reference"><a href="#cite_note-60"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup> For example, despite their obvious differences, <a href="/wiki/Spider" title="Spider">spiders</a> and <a href="/wiki/Barnacle" title="Barnacle">barnacles</a> both belong to the phylum Arthropoda, but <a href="/wiki/Earthworm" title="Earthworm">earthworms</a> and <a href="/wiki/Tapeworm" class="mw-redirect" title="Tapeworm">tapeworms</a>, although similar in shape, belong to different phyla. As chemical and genetic testing becomes more accurate, previously hypothesised phyla are often entirely reworked. </p><p>A phylum is not a fundamental division of nature, such as the difference between <a href="/wiki/Electron" title="Electron">electrons</a> and <a href="/wiki/Proton" title="Proton">protons</a>. It is simply a very high-level grouping in a <a href="/wiki/Linnaean_taxonomy" title="Linnaean taxonomy">classification system</a> created to describe all currently living organisms. This system is imperfect, even for modern animals: different books quote different numbers of phyla, mainly because they disagree about the classification of a huge number of worm-like species. As it is based on living organisms, it accommodates extinct organisms poorly, if at all.<sup id="cite_ref-CowenHistLife_25-3" class="reference"><a href="#cite_note-CowenHistLife-25"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Jefferies1979OriginOfChordates_61-0" class="reference"><a href="#cite_note-Jefferies1979OriginOfChordates-61"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Stem_group">Stem group</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=9" title="Edit section: Stem group"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The concept of <a href="/wiki/Stem_group" class="mw-redirect" title="Stem group">stem groups</a> was introduced to cover evolutionary "aunts" and "cousins" of living groups, and have been hypothesized based on this scientific theory. A <a href="/wiki/Crown_group" title="Crown group">crown group</a> is a group of closely related living animals plus their last common ancestor plus all its descendants. A stem group is a set of offshoots from the lineage at a point earlier than the last common ancestor of the crown group; it is a relative concept, for example <a href="/wiki/Tardigrade" title="Tardigrade">tardigrades</a> are living animals that form a crown group in their own right, but Budd (1996) regarded them as also being a stem group relative to the arthropods.<sup id="cite_ref-CraskeJefferies1989_58-1" class="reference"><a href="#cite_note-CraskeJefferies1989-58"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Budd1996_62-0" class="reference"><a href="#cite_note-Budd1996-62"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> </p> <div style="clear:both;" class=""></div> <div class="noresize thumb tright" style=";"> <div class="thumbinner" style="overflow:hidden;width:245px;"> <div class="thumbimage" style="height:256px; overflow:hidden; position:relative; background-color:white;"> <div style=";left:0px; top:0px; width:250px; position:absolute;"> <span typeof="mw:File"><a href="/wiki/File:Coelomate.svg" class="mw-file-description" title="A coelomate animal is basically a set of concentric tubes, with a gap between the gut and the outer tubes."><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c3/Coelomate.svg/250px-Coelomate.svg.png" decoding="async" width="250" height="250" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c3/Coelomate.svg/375px-Coelomate.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c3/Coelomate.svg/500px-Coelomate.svg.png 2x" data-file-width="250" data-file-height="250" /></a></span></div> <div style="text-align:left; background-color:transparent; font-size:15px; line-height:17px;"> <div id="annotation_30x5" style="position:absolute; left:30px; top:5px; line-height:110%;"><span style="background-color:transparent; color:inherit;"><a href="/wiki/Ectoderm" title="Ectoderm">Skin<br />(ectoderm)</a></span></div> <div id="annotation_10x60" style="position:absolute; left:10px; top:60px; line-height:110%;"><span style="background-color:transparent; color:inherit;"><a href="/wiki/Mesoderm" title="Mesoderm">Muscle<br />(mesoderm)</a></span></div> <div id="annotation_10x110" style="position:absolute; left:10px; top:110px; line-height:110%;"><span style="background-color:transparent; color:inherit;"><a href="/wiki/Coelom" title="Coelom">Coelom</a></span></div> <div id="annotation_10x150" style="position:absolute; left:10px; top:150px; line-height:110%;"><span style="background-color:transparent; color:inherit;"><a href="/wiki/Organ_(anatomy)" class="mw-redirect" title="Organ (anatomy)">Internal<br />organ</a></span></div> <div id="annotation_10x200" style="position:absolute; left:10px; top:200px; line-height:110%;"><span style="background-color:transparent; color:inherit;"><a href="/wiki/Peritoneum" title="Peritoneum">Membrane<br />(peritoneum)</a></span></div> <div id="annotation_160x210" style="position:absolute; left:160px; top:210px; line-height:110%;"><span style="background-color:transparent; color:inherit;"><a href="/wiki/Endoderm" title="Endoderm">Gut<br />(endoderm)</a></span></div> </div> <div style="visibility:hidden"><span class="noviewer" typeof="mw:File/Frameless"><a href="/wiki/File:Coelomate.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c3/Coelomate.svg/250px-Coelomate.svg.png" decoding="async" width="250" height="250" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c3/Coelomate.svg/375px-Coelomate.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c3/Coelomate.svg/500px-Coelomate.svg.png 2x" data-file-width="250" data-file-height="250" /></a></span></div> </div> <div class="thumbcaption"><div class="magnify"><a href="/wiki/File:Coelomate.svg" title="File:Coelomate.svg"> </a></div>A coelomate animal is basically a set of concentric tubes, with a gap between the gut and the outer tubes.</div> </div></div> <div class="mw-heading mw-heading3"><h3 id="Triploblastic">Triploblastic</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=10" title="Edit section: Triploblastic"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The term <i><a href="/wiki/Triploblastic" class="mw-redirect" title="Triploblastic">Triploblastic</a></i> means consisting of three layers, which are formed in the <a href="/wiki/Embryo" title="Embryo">embryo</a>, quite early in the animal's development from a single-celled egg to a larva or juvenile form. The innermost layer forms the <a href="/wiki/Digestive_tract" class="mw-redirect" title="Digestive tract">digestive tract</a> (gut); the outermost forms skin; and the middle one forms muscles and all the internal organs except the digestive system. Most types of living animal are triploblastic—the best-known exceptions are <a href="/wiki/Porifera" class="mw-redirect" title="Porifera">Porifera</a> (sponges) and <a href="/wiki/Cnidaria" title="Cnidaria">Cnidaria</a> (jellyfish, sea anemones, etc.). </p> <div class="mw-heading mw-heading3"><h3 id="Bilaterian">Bilaterian</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=11" title="Edit section: Bilaterian"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The <a href="/wiki/Bilateria" title="Bilateria">bilaterians</a> are animals that have right and left sides at some point in their life histories. This implies that they have top and bottom surfaces and, importantly, distinct front and back ends. All known bilaterian animals are triploblastic, and all known triploblastic animals are bilaterian. Living <a href="/wiki/Echinoderm" title="Echinoderm">echinoderms</a> (<a href="/wiki/Sea_star" class="mw-redirect" title="Sea star">sea stars</a>, <a href="/wiki/Sea_urchin" title="Sea urchin">sea urchins</a>, <a href="/wiki/Holothuroidea" class="mw-redirect" title="Holothuroidea">sea cucumbers</a>, etc.) 'look' radially symmetrical (like wheels) rather than bilaterian, but their larvae exhibit bilateral symmetry and some of the earliest echinoderms may have been bilaterally symmetrical.<sup id="cite_ref-63" class="reference"><a href="#cite_note-63"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup> Porifera and Cnidaria are radially symmetrical, not bilaterian, and not triploblastic (but the common Bilateria-Cnidaria ancestor's <a href="/wiki/Planula" title="Planula">planula</a> larva is suspected to be bilaterally symmetric). </p> <div class="mw-heading mw-heading3"><h3 id="Coelomate">Coelomate</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=12" title="Edit section: Coelomate"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The term <i><a href="/wiki/Coelomate" class="mw-redirect" title="Coelomate">Coelomate</a></i> means having a body cavity (coelom) containing the internal organs. Most of the phyla featured in the debate about the Cambrian explosion<sup class="noprint Inline-Template" style="margin-left:0.1em; white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Please_clarify" title="Wikipedia:Please clarify"><span title="The text near this tag may need clarification or removal of jargon. (January 2019)">clarification needed</span></a></i>]</sup> are coelomates: arthropods, <a href="/wiki/Annelid" title="Annelid">annelid</a> worms, molluscs, echinoderms and <a href="/wiki/Chordate" title="Chordate">chordates</a>—the noncoelomate <a href="/wiki/Priapulid" class="mw-redirect" title="Priapulid">priapulids</a> are an important exception. All known coelomate animals are triploblastic bilaterians, but some triploblastic bilaterian animals do not have a coelom—for example <a href="/wiki/Flatworm" title="Flatworm">flatworms</a>, whose organs are surrounded by <a href="/wiki/Parenchyma" title="Parenchyma">unspecialized tissues</a>. </p> <div class="mw-heading mw-heading2"><h2 id="Precambrian_life">Precambrian life</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=13" title="Edit section: Precambrian life"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Evidence_of_animals_around_1_billion_years_ago">Evidence of animals around 1 billion years ago</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=14" title="Edit section: Evidence of animals around 1 billion years ago"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Acritarch" title="Acritarch">Acritarch</a> and <a href="/wiki/Stromatolite" title="Stromatolite">Stromatolite</a></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:CambrianStromatolites.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f0/CambrianStromatolites.jpg/220px-CambrianStromatolites.jpg" decoding="async" width="220" height="79" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f0/CambrianStromatolites.jpg/330px-CambrianStromatolites.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f0/CambrianStromatolites.jpg/440px-CambrianStromatolites.jpg 2x" data-file-width="3929" data-file-height="1416" /></a><figcaption><a href="/wiki/Stromatolite" title="Stromatolite">Stromatolites</a> (Pika Formation, Middle Cambrian) near Helen Lake, <a href="/wiki/Banff_National_Park" title="Banff National Park">Banff National Park</a>, Canada</figcaption></figure> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Stromatolites_in_Sharkbay.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Stromatolites_in_Sharkbay.jpg/250px-Stromatolites_in_Sharkbay.jpg" decoding="async" width="220" height="164" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Stromatolites_in_Sharkbay.jpg/330px-Stromatolites_in_Sharkbay.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/1b/Stromatolites_in_Sharkbay.jpg/500px-Stromatolites_in_Sharkbay.jpg 2x" data-file-width="2440" data-file-height="1816" /></a><figcaption>Modern stromatolites in <a href="/wiki/Hamelin_Pool_Marine_Nature_Reserve" title="Hamelin Pool Marine Nature Reserve">Hamelin Pool Marine Nature Reserve</a>, Western Australia</figcaption></figure> <p>Changes in the abundance and diversity of some types of fossil have been interpreted as evidence for "attacks" by animals or other organisms. Stromatolites, stubby pillars built by colonies of <a href="/wiki/Microorganism" title="Microorganism">microorganisms</a>, are a major constituent of the fossil record from about <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=2,700">2,700</a> million years ago</span>, but their abundance and diversity declined steeply after about <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=1,250">1,250</a> million years ago</span>. This decline has been attributed to disruption by grazing and burrowing animals.<sup id="cite_ref-McNamara1996DatingOriginAnimals_31-1" class="reference"><a href="#cite_note-McNamara1996DatingOriginAnimals-31"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-AwramikStromatoliteDiversityMetazoanAppearance_32-1" class="reference"><a href="#cite_note-AwramikStromatoliteDiversityMetazoanAppearance-32"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bengtson2002OriginsOfPredation_64-0" class="reference"><a href="#cite_note-Bengtson2002OriginsOfPredation-64"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup> </p><p>Precambrian marine diversity was dominated by small fossils known as <a href="/wiki/Acritarch" title="Acritarch">acritarchs</a>. This term describes almost any small organic-walled fossil—from the egg cases of small <a href="/wiki/Animal" title="Animal">metazoans</a> to resting <a href="/wiki/Cyst" title="Cyst">cysts</a> of many different kinds of <a href="/wiki/Chlorophyta" title="Chlorophyta">green algae</a>. After appearing around <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=2,000">2,000</a> million years ago</span>, acritarchs underwent a boom around <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=1,000">1,000</a> million years ago</span>, increasing in abundance, diversity, size, complexity of shape, and especially size and number of spines. Their increasingly spiny forms in the last 1 billion years may indicate an increased need for defence against predation. Other groups of small organisms from the <a href="/wiki/Neoproterozoic" title="Neoproterozoic">Neoproterozoic</a> era also show signs of antipredator defenses.<sup id="cite_ref-Bengtson2002OriginsOfPredation_64-1" class="reference"><a href="#cite_note-Bengtson2002OriginsOfPredation-64"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup> A consideration of taxon longevity appears to support an increase in predation pressure around this time.<sup id="cite_ref-Stanley2008_65-0" class="reference"><a href="#cite_note-Stanley2008-65"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup> </p><p> In general, the fossil record shows a very slow appearance of these lifeforms in the Precambrian, with many <a href="/wiki/Cyanobacteria" title="Cyanobacteria">cyanobacterial</a> species making up much of the underlying sediment.<sup id="cite_ref-Butterfield2007_66-0" class="reference"><a href="#cite_note-Butterfield2007-66"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup></p><figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Ediacaran_trace_fossil.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8f/Ediacaran_trace_fossil.jpg/220px-Ediacaran_trace_fossil.jpg" decoding="async" width="220" height="231" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8f/Ediacaran_trace_fossil.jpg/330px-Ediacaran_trace_fossil.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8f/Ediacaran_trace_fossil.jpg/440px-Ediacaran_trace_fossil.jpg 2x" data-file-width="1194" data-file-height="1251" /></a><figcaption>An Ediacaran trace fossil, made when an organism burrowed below a <a href="/wiki/Microbial_mat" title="Microbial mat">microbial mat</a></figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Ediacaran_organisms">Ediacaran organisms</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=15" title="Edit section: Ediacaran organisms"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:DickinsoniaCostata.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fb/DickinsoniaCostata.jpg/220px-DickinsoniaCostata.jpg" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fb/DickinsoniaCostata.jpg/330px-DickinsoniaCostata.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fb/DickinsoniaCostata.jpg/440px-DickinsoniaCostata.jpg 2x" data-file-width="2272" data-file-height="1704" /></a><figcaption><i><a href="/wiki/Dickinsonia" title="Dickinsonia">Dickinsonia costata</a></i>, an Ediacaran organism of unknown affinity, with a quilted appearance</figcaption></figure> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main articles: <a href="/wiki/Ediacaran_biota" title="Ediacaran biota">Ediacaran biota</a>, <a href="/wiki/Cloudinidae" title="Cloudinidae">Cloudinidae</a>, <a href="/wiki/Kimberella" title="Kimberella">Kimberella</a>, and <a href="/wiki/Spriggina" title="Spriggina">Spriggina</a></div> <p>At the start of the Ediacaran period, much of the <a href="/wiki/Acritarch" title="Acritarch">acritarch</a> fauna, which had remained relatively unchanged for hundreds of millions of years, became extinct, to be replaced with a range of new, larger species, which would prove far more ephemeral.<sup id="cite_ref-Butterfield2007_66-1" class="reference"><a href="#cite_note-Butterfield2007-66"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup> This radiation, the first in the fossil record,<sup id="cite_ref-Butterfield2007_66-2" class="reference"><a href="#cite_note-Butterfield2007-66"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup> is followed soon after by an array of unfamiliar, large fossils dubbed the Ediacara biota,<sup id="cite_ref-67" class="reference"><a href="#cite_note-67"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup> which flourished for 40 million years until the start of the Cambrian.<sup id="cite_ref-Grazhdankin2004_68-0" class="reference"><a href="#cite_note-Grazhdankin2004-68"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup> Most of this "Ediacara biota" were at least a few centimeters long, significantly larger than any earlier fossils. The organisms form three distinct assemblages, increasing in size and complexity as time progressed.<sup id="cite_ref-Erwin1999OriginOfBodyplans_69-0" class="reference"><a href="#cite_note-Erwin1999OriginOfBodyplans-69"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> </p><p>Many of these organisms were quite unlike anything that appeared before or since, resembling discs, mud-filled bags, or quilted mattresses—one paleontologist proposed that the strangest organisms should be classified as a separate <a href="/wiki/Kingdom_(biology)" title="Kingdom (biology)">kingdom</a>, Vendozoa.<sup id="cite_ref-Seilacher1992_70-0" class="reference"><a href="#cite_note-Seilacher1992-70"><span class="cite-bracket">[</span>68<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Kimberella_blue.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b1/Kimberella_blue.jpg/220px-Kimberella_blue.jpg" decoding="async" width="220" height="161" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b1/Kimberella_blue.jpg/330px-Kimberella_blue.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b1/Kimberella_blue.jpg/440px-Kimberella_blue.jpg 2x" data-file-width="795" data-file-height="580" /></a><figcaption>Fossil of <i><a href="/wiki/Kimberella" title="Kimberella">Kimberella</a></i>, a triploblastic bilaterian, and possibly a mollusc</figcaption></figure> <p>At least some may have been early forms of the phyla at the heart of the "Cambrian explosion" debate,<sup class="noprint Inline-Template" style="margin-left:0.1em; white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Please_clarify" title="Wikipedia:Please clarify"><span title="The text near this tag may need clarification or removal of jargon. (January 2019)">clarification needed</span></a></i>]</sup> having been interpreted as early molluscs (<i><a href="/wiki/Kimberella" title="Kimberella">Kimberella</a></i>),<sup id="cite_ref-FedonkinWaggoner1997KimberellaMollusc_33-1" class="reference"><a href="#cite_note-FedonkinWaggoner1997KimberellaMollusc-33"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Martin2000_71-0" class="reference"><a href="#cite_note-Martin2000-71"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup> echinoderms (<i><a href="/wiki/Arkarua" title="Arkarua">Arkarua</a></i>)<sup id="cite_ref-72" class="reference"><a href="#cite_note-72"><span class="cite-bracket">[</span>70<span class="cite-bracket">]</span></a></sup> and arthropods (<i><a href="/wiki/Spriggina" title="Spriggina">Spriggina</a></i>,<sup id="cite_ref-73" class="reference"><a href="#cite_note-73"><span class="cite-bracket">[</span>71<span class="cite-bracket">]</span></a></sup> <i><a href="/wiki/Parvancorina" title="Parvancorina">Parvancorina</a></i>,<sup id="cite_ref-74" class="reference"><a href="#cite_note-74"><span class="cite-bracket">[</span>72<span class="cite-bracket">]</span></a></sup> <i><a href="/wiki/Yilingia" title="Yilingia">Yilingia</a></i>). Still, debate exists about the classification of these specimens, mainly because the diagnostic features that allow taxonomists to classify more recent organisms, such as similarities to living organisms, are generally absent in the ediacarans.<sup id="cite_ref-Butterfield2006_75-0" class="reference"><a href="#cite_note-Butterfield2006-75"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup> However, there seems little doubt that <i>Kimberella</i> was at least a triploblastic bilaterian animal.<sup id="cite_ref-Butterfield2006_75-1" class="reference"><a href="#cite_note-Butterfield2006-75"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup> These organisms are central to the debate about how abrupt the Cambrian explosion was.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2019)">citation needed</span></a></i>]</sup> If some were early members of the animal phyla seen today, the "explosion" looks a lot less sudden than if all these organisms represent an unrelated "experiment", and were replaced by the animal kingdom fairly soon thereafter (40 million years is "soon" by evolutionary and geological standards). </p><p>The traces of organisms moving on and directly underneath the microbial mats that covered the Ediacaran sea floor are preserved from the Ediacaran period, about <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=565">565</a> million years ago</span>.<sup id="cite_ref-78" class="reference"><a href="#cite_note-78"><span class="cite-bracket">[</span>c<span class="cite-bracket">]</span></a></sup> They were probably made by organisms resembling <a href="/wiki/Earthworm" title="Earthworm">earthworms</a> in shape, size and how they moved. The burrow-makers have never been found preserved, but, because they would need a head and a tail, the burrowers probably had bilateral symmetry—which would in all probability make them bilaterian animals.<sup id="cite_ref-79" class="reference"><a href="#cite_note-79"><span class="cite-bracket">[</span>76<span class="cite-bracket">]</span></a></sup> They fed above the sediment surface, but were forced to burrow to avoid predators.<sup id="cite_ref-Dzik2007_80-0" class="reference"><a href="#cite_note-Dzik2007-80"><span class="cite-bracket">[</span>77<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Cambrian_life">Cambrian life</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=16" title="Edit section: Cambrian life"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Trace_fossils_2">Trace fossils</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=17" title="Edit section: Trace fossils"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Trace fossils (burrows, etc.) are a reliable indicator of what life was around, and indicate a diversification of life around the start of the Cambrian, with the freshwater realm colonized by animals almost as quickly as the oceans.<sup id="cite_ref-Kennedy2011_81-0" class="reference"><a href="#cite_note-Kennedy2011-81"><span class="cite-bracket">[</span>78<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Small_shelly_fauna">Small shelly fauna</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=18" title="Edit section: Small shelly fauna"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Small_shelly_fauna" title="Small shelly fauna">Small shelly fauna</a></div> <p>Fossils known as "<a href="/wiki/Small_shelly_fauna" title="Small shelly fauna">small shelly fauna</a>" have been found in many parts on the world, and date from just before the Cambrian to about 10 million years after the start of the Cambrian (the <a href="/wiki/Nemakit-Daldynian" class="mw-redirect" title="Nemakit-Daldynian">Nemakit-Daldynian</a> and <a href="/wiki/Tommotian" class="mw-redirect" title="Tommotian">Tommotian</a> ages; see <a href="#Timeline">timeline</a>). These are a very mixed collection of fossils: spines, <a href="/wiki/Sclerite" title="Sclerite">sclerites</a> (armor plates), tubes, <a href="/wiki/Archeocyathid" class="mw-redirect" title="Archeocyathid">archeocyathids</a> (sponge-like animals) and small shells very like those of <a href="/wiki/Brachiopod" title="Brachiopod">brachiopods</a> and snail-like molluscs—but all tiny, mostly 1 to 2 mm long.<sup id="cite_ref-Matthews1975_82-0" class="reference"><a href="#cite_note-Matthews1975-82"><span class="cite-bracket">[</span>79<span class="cite-bracket">]</span></a></sup> </p> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Artistic_reconstruction_of_the_Cambrian_(Drumian)_Marjum_biota.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d6/Artistic_reconstruction_of_the_Cambrian_%28Drumian%29_Marjum_biota.png/250px-Artistic_reconstruction_of_the_Cambrian_%28Drumian%29_Marjum_biota.png" decoding="async" width="250" height="162" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d6/Artistic_reconstruction_of_the_Cambrian_%28Drumian%29_Marjum_biota.png/500px-Artistic_reconstruction_of_the_Cambrian_%28Drumian%29_Marjum_biota.png 1.5x" data-file-width="3238" data-file-height="2095" /></a><figcaption>Artistic reconstruction of Cambrian life</figcaption></figure> <p>While small, these fossils are far more common than complete fossils of the organisms that produced them; crucially, they cover the window from the start of the Cambrian to the first lagerstätten: a period of time otherwise lacking in fossils. Hence, they supplement the conventional fossil record and allow the fossil ranges of many groups to be extended. </p> <div class="mw-heading mw-heading3"><h3 id="Cnidarians">Cnidarians</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=19" title="Edit section: Cnidarians"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The first cnidarian larvae, represented by the genus <i>Eolarva</i>, appeared in the Cambrian, although the identity of <i>Eolarva</i> as such is controversial. If it does represent a cnidarian larva, <i>Eolarva</i> would represent the first fossil evidence of indirect development in metazoans in the earliest Cambrian.<sup id="cite_ref-83" class="reference"><a href="#cite_note-83"><span class="cite-bracket">[</span>80<span class="cite-bracket">]</span></a></sup> </p><p>Medusozoans developed complex life cycles with a medusa stage during the Cambrian explosion, as evidenced by the discovery of <i><a href="/wiki/Burgessomedusa" title="Burgessomedusa">Burgessomedusa phasmiformis</a>.</i><sup id="cite_ref-84" class="reference"><a href="#cite_note-84"><span class="cite-bracket">[</span>81<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Trilobites">Trilobites</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=20" title="Edit section: Trilobites"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Olenoides_serratus_oblique_with_antennas.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d7/Olenoides_serratus_oblique_with_antennas.jpg/220px-Olenoides_serratus_oblique_with_antennas.jpg" decoding="async" width="220" height="227" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d7/Olenoides_serratus_oblique_with_antennas.jpg/330px-Olenoides_serratus_oblique_with_antennas.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d7/Olenoides_serratus_oblique_with_antennas.jpg/440px-Olenoides_serratus_oblique_with_antennas.jpg 2x" data-file-width="1114" data-file-height="1150" /></a><figcaption>A fossilized trilobite, an ancient type of <a href="/wiki/Arthropod" title="Arthropod">arthropod</a>: This specimen, from the Burgess Shale, preserves "soft parts"—the antennae and legs. </figcaption></figure> <p>The earliest <a href="/wiki/Trilobite" title="Trilobite">trilobite</a> fossils are about 530 million years old, but the class was already quite diverse and <a href="/wiki/Cosmopolitan_distribution" title="Cosmopolitan distribution">cosmopolitan</a>, suggesting they had been around for quite some time.<sup id="cite_ref-Lieberman1999Trilobites_85-0" class="reference"><a href="#cite_note-Lieberman1999Trilobites-85"><span class="cite-bracket">[</span>82<span class="cite-bracket">]</span></a></sup> The fossil record of trilobites began with the appearance of trilobites with mineral exoskeletons—not from the time of their origin. </p> <div class="mw-heading mw-heading3"><h3 id="Crustaceans">Crustaceans</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=21" title="Edit section: Crustaceans"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Orsten" title="Orsten">Orsten</a></div> <p>Crustaceans, one of the four great modern groups of arthropods, are very rare throughout the Cambrian. Convincing <a href="/wiki/Crustacean" title="Crustacean">crustaceans</a> were once thought to be common in Burgess Shale-type biotas, but none of these individuals can be shown to fall into the crown group of "true crustaceans".<sup id="cite_ref-Harvey2008_86-0" class="reference"><a href="#cite_note-Harvey2008-86"><span class="cite-bracket">[</span>83<span class="cite-bracket">]</span></a></sup> The Cambrian record of crown-group crustaceans comes from <a href="/wiki/Microfossil" title="Microfossil">microfossils</a>. The Swedish <a href="/wiki/Orsten" title="Orsten">Orsten</a> horizons contain later Cambrian crustaceans, but only organisms smaller than 2 mm are preserved. This restricts the data set to juveniles and miniaturised adults. </p><p>A more informative data source is the organic microfossils of the <a href="/wiki/Mount_Cap_formation" title="Mount Cap formation">Mount Cap formation</a>, Mackenzie Mountains, Canada. This late Early Cambrian assemblage (<span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=510–515">510 to 515</a> million years ago</span>) consists of microscopic fragments of arthropods' cuticle, which is left behind when the rock is dissolved with <a href="/wiki/Hydrofluoric_acid" title="Hydrofluoric acid">hydrofluoric acid</a>. The diversity of this assemblage is similar to that of modern crustacean faunas. Analysis of fragments of feeding machinery found in the formation shows that it was adapted to feed in a very precise and refined fashion. This contrasts with most other early Cambrian arthropods, which fed messily by shovelling anything they could get their feeding appendages on into their mouths. This sophisticated and specialised feeding machinery belonged to a large (about 30 cm)<sup id="cite_ref-87" class="reference"><a href="#cite_note-87"><span class="cite-bracket">[</span>84<span class="cite-bracket">]</span></a></sup> organism, and would have provided great potential for diversification: Specialised feeding apparatus allows a number of different approaches to feeding and development, and creates a number of different approaches to avoid being eaten.<sup id="cite_ref-Harvey2008_86-1" class="reference"><a href="#cite_note-Harvey2008-86"><span class="cite-bracket">[</span>83<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Echinoderms">Echinoderms</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=22" title="Edit section: Echinoderms"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The earliest generally accepted echinoderm fossils appeared in the Late <a href="/wiki/Atdabanian" class="mw-redirect" title="Atdabanian">Atdabanian</a>; unlike modern echinoderms, these early Cambrian echinoderms were not all radially symmetrical.<sup id="cite_ref-DornbosBottjer2000Helicoplacoids_88-0" class="reference"><a href="#cite_note-DornbosBottjer2000Helicoplacoids-88"><span class="cite-bracket">[</span>85<span class="cite-bracket">]</span></a></sup> These provide firm data points for the "end" of the explosion, or at least indications that the crown groups of modern phyla were represented. </p> <div class="mw-heading mw-heading3"><h3 id="Burrowing">Burrowing</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=23" title="Edit section: Burrowing"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Cambrian_substrate_revolution" title="Cambrian substrate revolution">Cambrian substrate revolution</a></div> <p>Around the start of the Cambrian (about <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=539">539</a> million years ago</span>), many new types of traces first appear, including well-known vertical burrows such as <i><a href="/wiki/Diplocraterion" title="Diplocraterion">Diplocraterion</a></i> and <i><a href="/wiki/Skolithos" title="Skolithos">Skolithos</a></i>, and traces normally attributed to arthropods, such as <i><a href="/wiki/Cruziana" title="Cruziana">Cruziana</a></i> and <i><a href="/wiki/Rusophycus" title="Rusophycus">Rusophycus</a></i>. The vertical burrows indicate that worm-like animals acquired new behaviours, and possibly new physical capabilities. Some Cambrian trace fossils indicate that their makers possessed hard <a href="/wiki/Exoskeleton" title="Exoskeleton">exoskeletons</a>, although they were not necessarily mineralised.<sup id="cite_ref-Jensen2003_77-1" class="reference"><a href="#cite_note-Jensen2003-77"><span class="cite-bracket">[</span>75<span class="cite-bracket">]</span></a></sup> Meiofaunal as well as macrofaunal bilaterians participated in this invasion of infaunal niches.<sup id="cite_ref-89" class="reference"><a href="#cite_note-89"><span class="cite-bracket">[</span>86<span class="cite-bracket">]</span></a></sup> </p><p>Burrows provide firm evidence of complex organisms; they are also much more readily preserved than body fossils, to the extent that the absence of trace fossils has been used to imply the genuine absence of large, motile, bottom-dwelling organisms.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="try Budd (June 2008)">citation needed</span></a></i>]</sup> They provide a further line of evidence to show that the Cambrian explosion represents a real diversification, and is not a preservational artefact.<sup id="cite_ref-Seilacher2005_90-0" class="reference"><a href="#cite_note-Seilacher2005-90"><span class="cite-bracket">[</span>87<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Skeletonisation">Skeletonisation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=24" title="Edit section: Skeletonisation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The first Ediacaran and lowest Cambrian (<a href="/wiki/Nemakit-Daldynian" class="mw-redirect" title="Nemakit-Daldynian">Nemakit-Daldynian</a>) skeletal fossils represent tubes and problematic sponge spicules.<sup id="cite_ref-Li1998_91-0" class="reference"><a href="#cite_note-Li1998-91"><span class="cite-bracket">[</span>88<span class="cite-bracket">]</span></a></sup> The oldest sponge spicules are monaxon siliceous, aged around <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=580">580</a> million years ago</span>, known from the <a href="/wiki/Doushantuo_Formation" title="Doushantuo Formation">Doushantuo Formation</a> in China and from deposits of the same age in <a href="/wiki/Mongolia" title="Mongolia">Mongolia</a>, although the interpretation of these fossils as spicules has been challenged.<sup id="cite_ref-Yin2001_92-0" class="reference"><a href="#cite_note-Yin2001-92"><span class="cite-bracket">[</span>89<span class="cite-bracket">]</span></a></sup> In the late Ediacaran-lowest Cambrian, numerous tube dwellings of enigmatic organisms appeared. It was organic-walled tubes (e.g. <i><a href="/wiki/Saarina" title="Saarina">Saarina</a></i>) and chitinous tubes of the sabelliditids (e.g. <i>Sokoloviina</i>, <i>Sabellidites</i>, <i>Paleolina</i>)<sup id="cite_ref-Gnilovskaya1996_93-0" class="reference"><a href="#cite_note-Gnilovskaya1996-93"><span class="cite-bracket">[</span>90<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Fedonkin2003_94-0" class="reference"><a href="#cite_note-Fedonkin2003-94"><span class="cite-bracket">[</span>91<span class="cite-bracket">]</span></a></sup> that prospered up to the beginning of the <a href="/wiki/Tommotian" class="mw-redirect" title="Tommotian">Tommotian</a>. The mineralized tubes of <i><a href="/wiki/Cloudina" class="mw-redirect" title="Cloudina">Cloudina</a></i>, <i><a href="/wiki/Namacalathus" title="Namacalathus">Namacalathus</a></i>, <i><a href="/wiki/Sinotubulites" title="Sinotubulites">Sinotubulites</a></i> and a dozen more of the other organisms from carbonate rocks formed near the end of the Ediacaran period from <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=549–542">549 to 542</a> million years ago</span>, as well as the triradially symmetrical mineralized tubes of anabaritids (e.g. <i><a href="/wiki/Anabarites" title="Anabarites">Anabarites</a></i>, <i>Cambrotubulus</i>) from uppermost Ediacaran and lower Cambrian.<sup id="cite_ref-Zhuravlev_2009_95-0" class="reference"><a href="#cite_note-Zhuravlev_2009-95"><span class="cite-bracket">[</span>92<span class="cite-bracket">]</span></a></sup> Ediacaran mineralized tubes are often found in carbonates of the stromatolite reefs and <a href="/wiki/Thrombolite" title="Thrombolite">thrombolites</a>,<sup id="cite_ref-Hofmann2001_96-0" class="reference"><a href="#cite_note-Hofmann2001-96"><span class="cite-bracket">[</span>93<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Grotzinger2000_97-0" class="reference"><a href="#cite_note-Grotzinger2000-97"><span class="cite-bracket">[</span>94<span class="cite-bracket">]</span></a></sup> i.e. they could live in an environment adverse to the majority of animals. </p><p>Although they are as hard to classify as most other Ediacaran organisms, they are important in two other ways. First, they are the earliest known calcifying organisms (organisms that built shells from <a href="/wiki/Calcium_carbonate" title="Calcium carbonate">calcium carbonate</a>).<sup id="cite_ref-Grotzinger2000_97-1" class="reference"><a href="#cite_note-Grotzinger2000-97"><span class="cite-bracket">[</span>94<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Hua2005_98-0" class="reference"><a href="#cite_note-Hua2005-98"><span class="cite-bracket">[</span>95<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Miller2004_99-0" class="reference"><a href="#cite_note-Miller2004-99"><span class="cite-bracket">[</span>96<span class="cite-bracket">]</span></a></sup> Secondly, these tubes are a device to rise over a substrate and competitors for effective feeding and, to a lesser degree, they serve as armor for protection against predators and adverse conditions of environment. Some <i>Cloudina</i> fossils show small holes in shells. The holes possibly are evidence of boring by predators sufficiently advanced to penetrate shells.<sup id="cite_ref-Hua2003_100-0" class="reference"><a href="#cite_note-Hua2003-100"><span class="cite-bracket">[</span>97<span class="cite-bracket">]</span></a></sup> A possible "<a href="/wiki/Evolutionary_arms_race" title="Evolutionary arms race">evolutionary arms race</a>" between predators and prey is one of the hypotheses that attempt to explain the Cambrian explosion.<sup id="cite_ref-Bengtson2002OriginsOfPredation_64-2" class="reference"><a href="#cite_note-Bengtson2002OriginsOfPredation-64"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup> </p><p>In the lowest Cambrian, the stromatolites were decimated. This allowed animals to begin colonization of warm-water pools with carbonate sedimentation. At first, it was <a href="/wiki/Anabaritid" title="Anabaritid">anabaritids</a> and <i><a href="/wiki/Protohertzina" title="Protohertzina">Protohertzina</a></i> (the fossilized grasping spines of <a href="/wiki/Chaetognatha" title="Chaetognatha">chaetognaths</a>) fossils. Such mineral skeletons as shells, sclerites, thorns and plates appeared in uppermost <a href="/wiki/Nemakit-Daldynian" class="mw-redirect" title="Nemakit-Daldynian">Nemakit-Daldynian</a>; they were the earliest species of <a href="/wiki/Halkieria" class="mw-redirect" title="Halkieria">halkierids</a>, <a href="/wiki/Helcionelloida" title="Helcionelloida">gastropods</a>, <a href="/wiki/Hyolitha" title="Hyolitha">hyoliths</a> and other rare organisms. The beginning of the <a href="/wiki/Tommotian" class="mw-redirect" title="Tommotian">Tommotian</a> has historically been understood to mark an explosive increase of the number and variety of fossils of molluscs, <a href="/wiki/Hyolith" class="mw-redirect" title="Hyolith">hyoliths</a> and <a href="/wiki/Sponge" title="Sponge">sponges</a>, along with a rich complex of skeletal elements of unknown animals, the first <a href="/wiki/Archaeocyathid" class="mw-redirect" title="Archaeocyathid">archaeocyathids</a>, <a href="/wiki/Brachiopod" title="Brachiopod">brachiopods</a>, <a href="/wiki/Tommotiid" title="Tommotiid">tommotiids</a> and others.<sup id="cite_ref-Steiner2007_101-0" class="reference"><a href="#cite_note-Steiner2007-101"><span class="cite-bracket">[</span>98<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Rozanov2008_102-0" class="reference"><a href="#cite_note-Rozanov2008-102"><span class="cite-bracket">[</span>99<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Khomentovsky2005_103-0" class="reference"><a href="#cite_note-Khomentovsky2005-103"><span class="cite-bracket">[</span>100<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Khomentovsky2002_104-0" class="reference"><a href="#cite_note-Khomentovsky2002-104"><span class="cite-bracket">[</span>101<span class="cite-bracket">]</span></a></sup> Also <a href="/wiki/Soft-bodied_organism" title="Soft-bodied organism">soft-bodied</a> extant phyla such as <a href="/wiki/Ctenophora" title="Ctenophora">comb jellies</a>, <a href="/wiki/Scalidophora" title="Scalidophora">scalidophorans</a>, <a href="/wiki/Entoprocta" title="Entoprocta">entoproctans</a>, <a href="/wiki/Phoronid" title="Phoronid">horseshoe worms</a> and <a href="/wiki/Lobopodia" title="Lobopodia">lobopodians</a> had armored forms.<sup id="cite_ref-105" class="reference"><a href="#cite_note-105"><span class="cite-bracket">[</span>102<span class="cite-bracket">]</span></a></sup> This sudden increase is partially an artefact of missing strata at the Tommotian-type section, and most of this fauna in fact began to diversify in a series of pulses through the Nemakit-Daldynian and into the Tommotian.<sup id="cite_ref-Maloof2010_106-0" class="reference"><a href="#cite_note-Maloof2010-106"><span class="cite-bracket">[</span>103<span class="cite-bracket">]</span></a></sup> </p><p>Some animals may already have had sclerites, thorns, and plates in the Ediacaran (e.g. <i>Kimberella</i> had hard sclerites, probably of carbonate), but thin carbonate skeletons cannot be fossilized in <a href="/wiki/Siliciclastic" title="Siliciclastic">siliciclastic</a> deposits.<sup id="cite_ref-Ivantsov2009_107-0" class="reference"><a href="#cite_note-Ivantsov2009-107"><span class="cite-bracket">[</span>104<span class="cite-bracket">]</span></a></sup> Older (~750 Ma) fossils indicate that mineralization long preceded the Cambrian, probably defending small photosynthetic algae from single-celled eukaryotic predators.<sup id="cite_ref-Porter2011_108-0" class="reference"><a href="#cite_note-Porter2011-108"><span class="cite-bracket">[</span>105<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Cohen2011_109-0" class="reference"><a href="#cite_note-Cohen2011-109"><span class="cite-bracket">[</span>106<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Burgess_Shale_type_faunas">Burgess Shale type faunas</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=25" title="Edit section: Burgess Shale type faunas"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Burgess_Shale-type_preservation" title="Burgess Shale-type preservation">Burgess Shale-type preservation</a></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Diorama_of_the_Burgess_Shale_Biota_(Middle_Cambrian)_-_arthropods,_sponges,_worms.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/30/Diorama_of_the_Burgess_Shale_Biota_%28Middle_Cambrian%29_-_arthropods%2C_sponges%2C_worms.jpg/260px-Diorama_of_the_Burgess_Shale_Biota_%28Middle_Cambrian%29_-_arthropods%2C_sponges%2C_worms.jpg" decoding="async" width="260" height="119" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/30/Diorama_of_the_Burgess_Shale_Biota_%28Middle_Cambrian%29_-_arthropods%2C_sponges%2C_worms.jpg/390px-Diorama_of_the_Burgess_Shale_Biota_%28Middle_Cambrian%29_-_arthropods%2C_sponges%2C_worms.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/30/Diorama_of_the_Burgess_Shale_Biota_%28Middle_Cambrian%29_-_arthropods%2C_sponges%2C_worms.jpg/520px-Diorama_of_the_Burgess_Shale_Biota_%28Middle_Cambrian%29_-_arthropods%2C_sponges%2C_worms.jpg 2x" data-file-width="3648" data-file-height="1671" /></a><figcaption>Diorama of the <a href="/wiki/Paleobiota_of_the_Burgess_Shale" title="Paleobiota of the Burgess Shale">Burgess Shale Biota</a></figcaption></figure> <p>The Burgess Shale and similar lagerstätten preserve the soft parts of organisms, which provide a wealth of data to aid in the classification of enigmatic fossils. It often preserved complete specimens of organisms only otherwise known from dispersed parts, such as loose scales or isolated mouthparts. Further, the majority of organisms and taxa in these horizons are entirely soft-bodied, hence absent from the rest of the fossil record.<sup id="cite_ref-Butterfield2003_110-0" class="reference"><a href="#cite_note-Butterfield2003-110"><span class="cite-bracket">[</span>107<span class="cite-bracket">]</span></a></sup> Since a large part of the ecosystem is preserved, the ecology of the community can also be tentatively reconstructed.<sup class="noprint Inline-Template" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Verifiability" title="Wikipedia:Verifiability"><span title="The material near this tag needs to be fact-checked with the cited source(s). (July 2008)">verification needed</span></a></i>]</sup> However, the assemblages may represent a "museum": a deep-water ecosystem that is evolutionarily "behind" the rapidly diversifying fauna of shallower waters.<sup id="cite_ref-ConwayMorris2008_111-0" class="reference"><a href="#cite_note-ConwayMorris2008-111"><span class="cite-bracket">[</span>108<span class="cite-bracket">]</span></a></sup> </p><p>Because the lagerstätten provide a mode and quality of preservation that is virtually absent outside of the Cambrian, many organisms appear completely different from anything known from the conventional fossil record. This led early workers in the field to attempt to shoehorn the organisms into extant phyla; the shortcomings of this approach led later workers to erect a multitude of new phyla to accommodate all the oddballs. It has since been realised that most oddballs diverged from <a href="/wiki/Stem_group" class="mw-redirect" title="Stem group">lineages</a> before they established the phyla known today<sup class="noprint Inline-Template" style="margin-left:0.1em; white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Please_clarify" title="Wikipedia:Please clarify"><span title="The text near this tag may need clarification or removal of jargon. (August 2008)">clarification needed</span></a></i>]</sup>—slightly different designs, which were fated to perish rather than flourish into phyla, as their cousin lineages did. </p><p>The preservational mode is rare in the preceding Ediacaran period, but those assemblages known show no trace of animal life—perhaps implying a genuine absence of macroscopic metazoans.<sup id="cite_ref-Xiao2002_112-0" class="reference"><a href="#cite_note-Xiao2002-112"><span class="cite-bracket">[</span>109<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Stages">Stages</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=26" title="Edit section: Stages"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The early Cambrian interval of diversification lasted for about the next 20<sup id="cite_ref-dev.biologists.org_6-1" class="reference"><a href="#cite_note-dev.biologists.org-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Budd_7-1" class="reference"><a href="#cite_note-Budd-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup>–25<sup id="cite_ref-The_Cambrian_conundrum:_early_diver_8-1" class="reference"><a href="#cite_note-The_Cambrian_conundrum:_early_diver-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-113" class="reference"><a href="#cite_note-113"><span class="cite-bracket">[</span>110<span class="cite-bracket">]</span></a></sup> million years, and its elevated rates of evolution had ended by the base of <a href="/wiki/Cambrian_Series_2" title="Cambrian Series 2">Cambrian Series 2</a>, <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=521">521</a> million years ago</span>, coincident with the first trilobites in the fossil record.<sup id="cite_ref-114" class="reference"><a href="#cite_note-114"><span class="cite-bracket">[</span>111<span class="cite-bracket">]</span></a></sup> Different authors define intervals of diversification during the early Cambrian different ways: </p><p><a href="/wiki/Ed_Landing" title="Ed Landing">Ed Landing</a> recognizes three stages: Stage 1, spanning the Ediacaran-Cambrian boundary, corresponds to a diversification of biomineralizing animals and of deep and complex burrows; Stage 2, corresponding to the radiation of molluscs and stem-group <a href="/wiki/Brachiopod" title="Brachiopod">Brachiopods</a> (<a href="/wiki/Hyolith" class="mw-redirect" title="Hyolith">hyoliths</a> and <a href="/wiki/Tommotiid" title="Tommotiid">tommotiids</a>), which apparently arose in intertidal waters; and Stage 3, seeing the Atdabanian diversification of trilobites in deeper waters, but little change in the intertidal realm.<sup id="cite_ref-115" class="reference"><a href="#cite_note-115"><span class="cite-bracket">[</span>112<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Graham_Budd" title="Graham Budd">Graham Budd</a> synthesises various schemes to produce a compatible view of the SSF record of the Cambrian explosion, divided slightly differently into four intervals: a "Tube world", lasting from <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=550–536">550 to 536</a> million years ago</span>, spanning the Ediacaran-Cambrian boundary, dominated by Cloudina, Namacalathus and pseudoconodont-type elements; a "Sclerite world", seeing the rise of halkieriids, tommotiids and hyoliths, lasting to the end of the Fortunian (c. 525 Ma); a brachiopod world, perhaps corresponding to the as yet unratified Cambrian Stage 2; and Trilobite World, kicking off in Stage 3.<sup id="cite_ref-Budd,_G.E._2016_116-0" class="reference"><a href="#cite_note-Budd,_G.E._2016-116"><span class="cite-bracket">[</span>113<span class="cite-bracket">]</span></a></sup> </p><p>Complementary to the shelly fossil record, trace fossils can be divided into five subdivisions: "Flat world" (late Ediacaran), with traces restricted to the sediment surface; Protreozoic III (after Jensen), with increasing complexity; <i>pedum</i> world, initiated at the base of the Cambrian with the base of the <i>T.pedum</i> zone (see <a href="/wiki/Cambrian#Dating_the_Cambrian" title="Cambrian">Cambrian#Dating the Cambrian</a>); <i>Rusophycus</i> world, spanning <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=536–521">536 to 521</a> million years ago</span> and thus corresponding exactly to the periods of Sclerite World and Brachiopod World under the SSF paradigm; and <i>Cruziana</i> world, with an obvious correspondence to Trilobite World.<sup id="cite_ref-Budd,_G.E._2016_116-1" class="reference"><a href="#cite_note-Budd,_G.E._2016-116"><span class="cite-bracket">[</span>113<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Validity">Validity</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=27" title="Edit section: Validity"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>There is strong evidence for species of <a href="/wiki/Cnidaria" title="Cnidaria">Cnidaria</a> and <a href="/wiki/Sponge" title="Sponge">Porifera</a> existing in the <a href="/wiki/Ediacaran" title="Ediacaran">Ediacaran</a><sup id="cite_ref-117" class="reference"><a href="#cite_note-117"><span class="cite-bracket">[</span>114<span class="cite-bracket">]</span></a></sup> and possible members of Porifera even before that during the <a href="/wiki/Cryogenian" title="Cryogenian">Cryogenian</a>.<sup id="cite_ref-118" class="reference"><a href="#cite_note-118"><span class="cite-bracket">[</span>115<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Bryozoa" title="Bryozoa">Bryozoans, once thought to</a> not appear in the fossil record until after the Cambrian, are now known from strata of Cambrian Age 3 from Australia and South China.<sup id="cite_ref-Taylor2013_119-0" class="reference"><a href="#cite_note-Taylor2013-119"><span class="cite-bracket">[</span>116<span class="cite-bracket">]</span></a></sup> </p><p>The fossil record as Darwin knew it seemed to suggest that the major metazoan groups appeared in a few million years of the early to mid-Cambrian, and even in the 1980s, this still appeared to be the case.<sup id="cite_ref-Whittington1985BurgessShale_28-1" class="reference"><a href="#cite_note-Whittington1985BurgessShale-28"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-WonderfulLife_29-1" class="reference"><a href="#cite_note-WonderfulLife-29"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> </p><p>However, evidence of Precambrian Metazoa is gradually accumulating. If the Ediacaran <i>Kimberella</i> was a mollusc-like <a href="/wiki/Protostome" title="Protostome">protostome</a> (one of the two main groups of <a href="/wiki/Coelomate" class="mw-redirect" title="Coelomate">coelomates</a>),<sup id="cite_ref-FedonkinWaggoner1997KimberellaMollusc_33-2" class="reference"><a href="#cite_note-FedonkinWaggoner1997KimberellaMollusc-33"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Martin2000_71-1" class="reference"><a href="#cite_note-Martin2000-71"><span class="cite-bracket">[</span>69<span class="cite-bracket">]</span></a></sup> the protostome and <a href="/wiki/Deuterostome" title="Deuterostome">deuterostome</a> lineages must have split significantly before <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=550">550</a> million years ago</span> (deuterostomes are the other main group of coelomates).<sup id="cite_ref-ErwinDavidson2002LastCommonBilaterianAncestor_120-0" class="reference"><a href="#cite_note-ErwinDavidson2002LastCommonBilaterianAncestor-120"><span class="cite-bracket">[</span>117<span class="cite-bracket">]</span></a></sup> Even if it is not a protostome, it is widely accepted as a bilaterian.<sup id="cite_ref-Butterfield2006_75-2" class="reference"><a href="#cite_note-Butterfield2006-75"><span class="cite-bracket">[</span>73<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-ErwinDavidson2002LastCommonBilaterianAncestor_120-1" class="reference"><a href="#cite_note-ErwinDavidson2002LastCommonBilaterianAncestor-120"><span class="cite-bracket">[</span>117<span class="cite-bracket">]</span></a></sup> Since fossils of rather modern-looking <a href="/wiki/Cnidaria" title="Cnidaria">cnidarians</a> (<a href="/wiki/Jellyfish" title="Jellyfish">jellyfish</a>-like organisms) have been found in the <a href="/wiki/Doushantuo_formation" class="mw-redirect" title="Doushantuo formation">Doushantuo</a> <a href="/wiki/Lagerst%C3%A4tte" title="Lagerstätte">lagerstätte</a>, the cnidarian and bilaterian lineages must have diverged well over <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=580">580</a> million years ago</span>.<sup id="cite_ref-ErwinDavidson2002LastCommonBilaterianAncestor_120-2" class="reference"><a href="#cite_note-ErwinDavidson2002LastCommonBilaterianAncestor-120"><span class="cite-bracket">[</span>117<span class="cite-bracket">]</span></a></sup> </p><p>Trace fossils<sup id="cite_ref-Erwin1999OriginOfBodyplans_69-1" class="reference"><a href="#cite_note-Erwin1999OriginOfBodyplans-69"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> and predatory borings in <i>Cloudina</i> shells provide further evidence of Ediacaran animals.<sup id="cite_ref-BengtsonZhao1992PredatorialBorings_121-0" class="reference"><a href="#cite_note-BengtsonZhao1992PredatorialBorings-121"><span class="cite-bracket">[</span>118<span class="cite-bracket">]</span></a></sup> Some fossils from the Doushantuo formation have been interpreted as embryos and one (<i><a href="/wiki/Vernanimalcula" title="Vernanimalcula">Vernanimalcula</a></i>) as a bilaterian coelomate, although these interpretations are not universally accepted.<sup id="cite_ref-Chen2004_122-0" class="reference"><a href="#cite_note-Chen2004-122"><span class="cite-bracket">[</span>119<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bengtson2004_123-0" class="reference"><a href="#cite_note-Bengtson2004-123"><span class="cite-bracket">[</span>120<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-ChenDefendVernanimacula_124-0" class="reference"><a href="#cite_note-ChenDefendVernanimacula-124"><span class="cite-bracket">[</span>121<span class="cite-bracket">]</span></a></sup> Earlier still, predatory pressure has acted on stromatolites and acritarchs since around <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=1,250">1,250</a> million years ago</span>.<sup id="cite_ref-Bengtson2002OriginsOfPredation_64-3" class="reference"><a href="#cite_note-Bengtson2002OriginsOfPredation-64"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup> </p><p>Some say that the evolutionary change was accelerated by an <a href="/wiki/Order_of_magnitude" title="Order of magnitude">order of magnitude</a>,<sup id="cite_ref-125" class="reference"><a href="#cite_note-125"><span class="cite-bracket">[</span>d<span class="cite-bracket">]</span></a></sup> but the presence of Precambrian animals somewhat dampens the "bang" of the explosion; not only was the appearance of animals gradual, but their <a href="/wiki/Evolutionary_radiation" title="Evolutionary radiation">evolutionary radiation</a> ("diversification") may also not have been as rapid as once thought. Indeed, statistical analysis shows that the Cambrian explosion was no faster than any of the other radiations in animals' history.<sup id="cite_ref-127" class="reference"><a href="#cite_note-127"><span class="cite-bracket">[</span>e<span class="cite-bracket">]</span></a></sup> However, it does seem that some innovations linked to the explosion—such as resistant armour—only evolved once in the animal lineage; this makes a lengthy Precambrian animal lineage harder to defend.<sup id="cite_ref-Jacobs2000_128-0" class="reference"><a href="#cite_note-Jacobs2000-128"><span class="cite-bracket">[</span>123<span class="cite-bracket">]</span></a></sup> Further, the conventional view that all the phyla arose in the Cambrian is flawed; while the phyla may have diversified in this time period, representatives of the crown groups of many phyla do not appear until much later in the Phanerozoic.<sup id="cite_ref-Budd2000_15-2" class="reference"><a href="#cite_note-Budd2000-15"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> Further, the mineralised phyla that form the basis of the fossil record may not be representative of other phyla, since most mineralised phyla originated in a <a href="/wiki/Benthic_zone" title="Benthic zone">benthic</a> setting. The fossil record is consistent with a Cambrian explosion that was limited to the benthos, with pelagic phyla evolving much later.<sup id="cite_ref-Budd2000_15-3" class="reference"><a href="#cite_note-Budd2000-15"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p><p>Ecological complexity among marine animals increased in the Cambrian, as well later in the Ordovician.<sup id="cite_ref-Bambach2007_14-1" class="reference"><a href="#cite_note-Bambach2007-14"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> However, recent research has overthrown the once-popular idea that disparity was exceptionally high throughout the Cambrian, before subsequently decreasing.<sup id="cite_ref-129" class="reference"><a href="#cite_note-129"><span class="cite-bracket">[</span>124<span class="cite-bracket">]</span></a></sup> In fact, disparity remains relatively low throughout the Cambrian, with modern levels of disparity only attained after the early Ordovician radiation.<sup id="cite_ref-Bambach2007_14-2" class="reference"><a href="#cite_note-Bambach2007-14"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p><p>The diversity of many Cambrian assemblages is similar to today's,<sup id="cite_ref-Harvey2008_86-2" class="reference"><a href="#cite_note-Harvey2008-86"><span class="cite-bracket">[</span>83<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-130" class="reference"><a href="#cite_note-130"><span class="cite-bracket">[</span>125<span class="cite-bracket">]</span></a></sup> and at a high (class/phylum) level, diversity is thought by some to have risen relatively smoothly through the Cambrian, stabilizing somewhat in the Ordovician.<sup id="cite_ref-Erwin2011_131-0" class="reference"><a href="#cite_note-Erwin2011-131"><span class="cite-bracket">[</span>126<span class="cite-bracket">]</span></a></sup> This interpretation, however, glosses over the astonishing and fundamental pattern of basal <a href="/wiki/Polytomy" title="Polytomy">polytomy</a> and phylogenetic telescoping at or near the Cambrian boundary, as seen in most major animal lineages.<sup id="cite_ref-132" class="reference"><a href="#cite_note-132"><span class="cite-bracket">[</span>127<span class="cite-bracket">]</span></a></sup> Thus <a href="/wiki/Harry_Blackmore_Whittington" class="mw-redirect" title="Harry Blackmore Whittington">Harry Blackmore Whittington</a>'s questions regarding the abrupt nature of the Cambrian explosion remain, and have yet to be satisfactorily answered.<sup id="cite_ref-133" class="reference"><a href="#cite_note-133"><span class="cite-bracket">[</span>128<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="The_Cambrian_explosion_as_survivorship_bias">The Cambrian explosion as survivorship bias</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=28" title="Edit section: The Cambrian explosion as survivorship bias"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Graham_Budd" title="Graham Budd">Budd</a> and Mann<sup id="cite_ref-POTPa_134-0" class="reference"><a href="#cite_note-POTPa-134"><span class="cite-bracket">[</span>129<span class="cite-bracket">]</span></a></sup> suggested that the Cambrian explosion was the result of a type of <a href="/wiki/Survivorship_bias" title="Survivorship bias">survivorship bias</a> called the "<a href="/wiki/Push_of_the_past" title="Push of the past">Push of the past</a>". As groups at their origin tend to go extinct, it follows that any long-lived group would have experienced an unusually rapid rate of diversification early on, creating the illusion of a general speed-up in diversification rates. However, rates of diversification could remain at background levels and still generate this sort of effect in the surviving lineages. </p> <div class="mw-heading mw-heading2"><h2 id="Possible_causes">Possible causes</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=29" title="Edit section: Possible causes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Despite the evidence that moderately complex animals (<a href="/wiki/Triploblastic" class="mw-redirect" title="Triploblastic">triploblastic</a> <a href="/wiki/Bilateria" title="Bilateria">bilaterians</a>) existed before and possibly long before the start of the Cambrian, it seems that the pace of evolution was exceptionally fast in the early Cambrian. Possible explanations for this fall into three broad categories: environmental, developmental and ecological changes. Any explanation must explain both the timing and magnitude of the explosion. </p> <div class="mw-heading mw-heading3"><h3 id="Changes_in_the_environment">Changes in the environment</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=30" title="Edit section: Changes in the environment"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading4"><h4 id="Increase_in_oxygen_levels">Increase in oxygen levels</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=31" title="Edit section: Increase in oxygen levels"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Atmosphere_of_Earth#Earliest_atmosphere" title="Atmosphere of Earth">Earth's earliest atmosphere</a> contained no free <a href="/wiki/Oxygen" title="Oxygen">oxygen</a> (O<sub>2</sub>); the oxygen that animals breathe today, both in the air and dissolved in water, is the product of billions of years of <a href="/wiki/Photosynthesis" title="Photosynthesis">photosynthesis</a>. Cyanobacteria were the first organisms to evolve the ability to photosynthesize, introducing a steady supply of oxygen into the environment.<sup id="cite_ref-135" class="reference"><a href="#cite_note-135"><span class="cite-bracket">[</span>130<span class="cite-bracket">]</span></a></sup> Initially, oxygen levels did not increase substantially in the atmosphere.<sup id="cite_ref-ReferenceA_136-0" class="reference"><a href="#cite_note-ReferenceA-136"><span class="cite-bracket">[</span>131<span class="cite-bracket">]</span></a></sup> The oxygen quickly reacted with iron and other minerals in the surrounding rock and ocean water. Once a saturation point was reached for the reactions in rock and water, oxygen was able to exist as a gas in its diatomic form. Oxygen levels in the atmosphere increased substantially afterward.<sup id="cite_ref-137" class="reference"><a href="#cite_note-137"><span class="cite-bracket">[</span>132<span class="cite-bracket">]</span></a></sup> As a general trend, the <a href="/wiki/Concentration" title="Concentration">concentration</a> of oxygen in the atmosphere has risen gradually over about the last 2.5 billion years.<sup id="cite_ref-CowenHistLife_25-4" class="reference"><a href="#cite_note-CowenHistLife-25"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> </p><p>Oxygen levels seem to have a positive correlation with diversity in <a href="/wiki/Eukaryotes" class="mw-redirect" title="Eukaryotes">eukaryotes</a> well before the Cambrian period.<sup id="cite_ref-ReferenceB_138-0" class="reference"><a href="#cite_note-ReferenceB-138"><span class="cite-bracket">[</span>133<span class="cite-bracket">]</span></a></sup> The last common ancestor of all extant eukaryotes is thought to have lived around 1.8 billion years ago. Around 800 million years ago, there was a notable increase in the complexity and number of eukaryotes species in the fossil record.<sup id="cite_ref-ReferenceB_138-1" class="reference"><a href="#cite_note-ReferenceB-138"><span class="cite-bracket">[</span>133<span class="cite-bracket">]</span></a></sup> Before the spike in diversity, eukaryotes are thought to have lived in highly sulfuric environments. <a href="/wiki/Sulfide" title="Sulfide">Sulfide</a> interferes with mitochondrial function in aerobic organisms, limiting the amount of oxygen that could be used to drive metabolism. Oceanic sulfide levels decreased around 800 million years ago, which supports the importance of oxygen in eukaryotic diversity.<sup id="cite_ref-ReferenceB_138-2" class="reference"><a href="#cite_note-ReferenceB-138"><span class="cite-bracket">[</span>133<span class="cite-bracket">]</span></a></sup> The increased ventilation of the oceans by sponges, which had already evolved and diversified during the late Neoproterozoic, has been proposed to have increased the availability of oxygen and powered the Cambrian's rapid diversification of multicellular life.<sup id="cite_ref-ChangEtAl2019_139-0" class="reference"><a href="#cite_note-ChangEtAl2019-139"><span class="cite-bracket">[</span>134<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-WangXieWen2022Precambrian_140-0" class="reference"><a href="#cite_note-WangXieWen2022Precambrian-140"><span class="cite-bracket">[</span>135<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Molybdenum" title="Molybdenum">Molybdenum</a> isotopes show that increases in biodiversity were strongly correlated with expansion of oxygenated bottom waters in the Early Cambrian, lending support for oxygen as a driver of the Cambrian evolutionary radiation.<sup id="cite_ref-141" class="reference"><a href="#cite_note-141"><span class="cite-bracket">[</span>136<span class="cite-bracket">]</span></a></sup> </p><p>The shortage of oxygen might well have prevented the rise of large, complex animals. The amount of oxygen an animal can absorb is largely determined by the area of its oxygen-absorbing surfaces (lungs and gills in the most complex animals; the skin in less complex ones), while the amount needed is determined by its volume, which grows faster than the oxygen-absorbing area if an animal's size increases equally in all directions. An increase in the concentration of oxygen in air or water would increase the size to which an organism could grow without its tissues becoming starved of oxygen. However, members of the Ediacara biota reached metres in length tens of millions of years before the Cambrian explosion.<sup id="cite_ref-Knoll1999_52-1" class="reference"><a href="#cite_note-Knoll1999-52"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> Other metabolic functions may have been inhibited by lack of oxygen, for example the construction of tissue such as <a href="/wiki/Collagen" title="Collagen">collagen</a>, which is required for the construction of complex structures,<sup id="cite_ref-Towe1970_142-0" class="reference"><a href="#cite_note-Towe1970-142"><span class="cite-bracket">[</span>137<span class="cite-bracket">]</span></a></sup> or the biosynthesis of molecules for the construction of a hard exoskeleton.<sup id="cite_ref-CatlingEtAl2005_143-0" class="reference"><a href="#cite_note-CatlingEtAl2005-143"><span class="cite-bracket">[</span>138<span class="cite-bracket">]</span></a></sup> However, animals were not affected when similar oceanographic conditions occurred in the Phanerozoic; therefore, some see no forcing role of the oxygen level on evolution.<sup id="cite_ref-Butterfield2009_144-0" class="reference"><a href="#cite_note-Butterfield2009-144"><span class="cite-bracket">[</span>139<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Ozone_formation">Ozone formation</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=32" title="Edit section: Ozone formation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The amount of <a href="/wiki/Ozone" title="Ozone">ozone</a> (O<sub>3</sub>) required to shield Earth from biologically lethal <a href="/wiki/Ultraviolet" title="Ultraviolet">UV radiation</a>, wavelengths from 200 to 300 nanometers (nm), is believed to have been in existence around the Cambrian explosion.<sup id="cite_ref-145" class="reference"><a href="#cite_note-145"><span class="cite-bracket">[</span>140<span class="cite-bracket">]</span></a></sup> The presence of the <a href="/wiki/Ozone_layer" title="Ozone layer">ozone layer</a> may have enabled the development of complex life and life on land, as opposed to life being restricted to the water. </p> <div class="mw-heading mw-heading4"><h4 id="Snowball_Earth">Snowball Earth</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=33" title="Edit section: Snowball Earth"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Snowball_Earth" title="Snowball Earth">Snowball Earth</a></div> <p>In the late <a href="/wiki/Neoproterozoic" title="Neoproterozoic">Neoproterozoic</a> (extending into the early <a href="/wiki/Ediacaran" title="Ediacaran">Ediacaran</a> period), the Earth suffered <a href="/wiki/Snowball_Earth" title="Snowball Earth">massive glaciations</a> in which most of its surface was covered by ice. This may have caused a mass extinction, creating a genetic bottleneck; the resulting diversification may have given rise to the <a href="/wiki/Ediacara_biota" class="mw-redirect" title="Ediacara biota">Ediacara biota</a>, which appears soon after the last "Snowball Earth" episode.<sup id="cite_ref-HoffmanKaufman1998NeoproterozoicSnowball_146-0" class="reference"><a href="#cite_note-HoffmanKaufman1998NeoproterozoicSnowball-146"><span class="cite-bracket">[</span>141<span class="cite-bracket">]</span></a></sup> However, the snowball episodes occurred a long time before the start of the Cambrian, and it is difficult to see how so much diversity could have been caused by even a series of bottlenecks;<sup id="cite_ref-Marshall2006Explaining_54-2" class="reference"><a href="#cite_note-Marshall2006Explaining-54"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> the cold periods may even have <i>delayed</i> the evolution of large size organisms.<sup id="cite_ref-Bengtson2002OriginsOfPredation_64-4" class="reference"><a href="#cite_note-Bengtson2002OriginsOfPredation-64"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup> Massive rock erosion caused by glaciers during the "Snowball Earth" may have deposited nutrient-rich sediments into the oceans, setting the stage for the Cambrian explosion.<sup id="cite_ref-147" class="reference"><a href="#cite_note-147"><span class="cite-bracket">[</span>142<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Increase_in_the_calcium_concentration_of_the_Cambrian_seawater">Increase in the calcium concentration of the Cambrian seawater</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=34" title="Edit section: Increase in the calcium concentration of the Cambrian seawater"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Newer research suggests that volcanically active <a href="/wiki/Mid-ocean_ridge" title="Mid-ocean ridge">mid-ocean ridges</a> caused a massive and sudden surge of the <a href="/wiki/Calcium" title="Calcium">calcium</a> concentration in the oceans, making it possible for marine organisms to build skeletons and hard body parts.<sup id="cite_ref-148" class="reference"><a href="#cite_note-148"><span class="cite-bracket">[</span>143<span class="cite-bracket">]</span></a></sup> Alternatively a high influx of ions could have been provided by the widespread erosion that produced Powell's <a href="/wiki/Great_Unconformity" title="Great Unconformity">Great Unconformity</a>.<sup id="cite_ref-Peters2012_149-0" class="reference"><a href="#cite_note-Peters2012-149"><span class="cite-bracket">[</span>144<span class="cite-bracket">]</span></a></sup> </p><p>An increase of calcium may also have been caused by erosion of the <a href="/wiki/East_African_Orogeny" title="East African Orogeny">Transgondwanan Supermountain</a> that existed at the time of the explosion. The roots of the mountain are preserved in present-day <a href="/wiki/East_Africa" title="East Africa">East Africa</a> as an <a href="/wiki/Orogeny" title="Orogeny">orogen</a>.<sup id="cite_ref-150" class="reference"><a href="#cite_note-150"><span class="cite-bracket">[</span>145<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Developmental_explanations">Developmental explanations</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=35" title="Edit section: Developmental explanations"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evolutionary developmental biology</a></div> <p>A range of theories are based on the concept that minor <a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">modifications to animals' development</a> as they grow from <a href="/wiki/Embryo" title="Embryo">embryo</a> to adult may have been able to cause very large changes in the final adult form. The <a href="/wiki/Hox_gene" title="Hox gene">Hox genes</a>, for example, control which organs individual regions of an embryo will develop into. For instance, if a certain <i>Hox</i> gene is expressed, a region will develop into a limb; if a different Hox gene is expressed in that region (a minor change), it could develop into an eye instead (a phenotypically major change). </p><p>Such a system allows a large range of disparity to appear from a limited set of genes, but such theories linking this with the explosion struggle to explain why the origin of such a development system should by itself lead to increased diversity or disparity. Evidence of Precambrian metazoans<sup id="cite_ref-Marshall2006Explaining_54-3" class="reference"><a href="#cite_note-Marshall2006Explaining-54"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> combines with molecular data<sup id="cite_ref-deRosa1999_151-0" class="reference"><a href="#cite_note-deRosa1999-151"><span class="cite-bracket">[</span>146<span class="cite-bracket">]</span></a></sup> to show that much of the genetic architecture that could feasibly have played a role in the explosion was already well established by the Cambrian. </p><p>This apparent paradox is addressed in a theory that focuses on the <a href="/wiki/Physics" title="Physics">physics</a> of development. It is proposed that the emergence of simple multicellular forms provided a changed context and spatial scale in which novel physical processes and effects were mobilized by the products of genes that had previously evolved to serve unicellular functions. Morphological complexity (layers, segments, <a href="/wiki/Lumen_(anatomy)" title="Lumen (anatomy)">lumens</a>, appendages) arose, in this view, by <a href="/wiki/Self-organization" title="Self-organization">self-organization</a>.<sup id="cite_ref-Newman2008_152-0" class="reference"><a href="#cite_note-Newman2008-152"><span class="cite-bracket">[</span>147<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Horizontal_gene_transfer" title="Horizontal gene transfer">Horizontal gene transfer</a> has also been identified as a possible factor in the rapid acquisition of the biochemical capability of biomineralization among organisms during this period, based on evidence that the gene for a critical protein in the process was originally transferred from a bacterium into sponges.<sup id="cite_ref-153" class="reference"><a href="#cite_note-153"><span class="cite-bracket">[</span>148<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Ecological_explanations">Ecological explanations</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=36" title="Edit section: Ecological explanations"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>These focus on the interactions between different types of organism. Some of these hypotheses deal with changes in the <a href="/wiki/Food_chain" title="Food chain">food chain</a>; some suggest <a href="/wiki/Evolutionary_arms_race" title="Evolutionary arms race">arms races</a> between predators and prey, and others focus on the more general mechanisms of <a href="/wiki/Coevolution" title="Coevolution">coevolution</a>. Such theories are well suited to explaining why there was a rapid increase in both disparity and diversity, but they do not explain why the "explosion" happened when it did.<sup id="cite_ref-Marshall2006Explaining_54-4" class="reference"><a href="#cite_note-Marshall2006Explaining-54"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="End-Ediacaran_mass_extinction">End-Ediacaran mass extinction</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=37" title="Edit section: End-Ediacaran mass extinction"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/End-Ediacaran_extinction" title="End-Ediacaran extinction">End-Ediacaran extinction</a></div> <p>Evidence for such an extinction includes the disappearance from the fossil record of the Ediacara biota and shelly fossils such as <i>Cloudina</i>, and the accompanying perturbation in the <a href="/wiki/%CE%9413C" title="Δ13C"><span class="texhtml">δ</span><sup>13</sup>C</a> record. It is suspected that several global <a href="/wiki/Anoxic_event" title="Anoxic event">anoxic events</a> were responsible for the extinction.<sup id="cite_ref-154" class="reference"><a href="#cite_note-154"><span class="cite-bracket">[</span>149<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-155" class="reference"><a href="#cite_note-155"><span class="cite-bracket">[</span>150<span class="cite-bracket">]</span></a></sup> </p><p>Mass extinctions are often followed by <a href="/wiki/Adaptive_radiation" title="Adaptive radiation">adaptive radiations</a> as existing clades expand to occupy the ecospace emptied by the extinction. However, once the dust had settled, overall disparity and diversity returned to the pre-extinction level in each of the Phanerozoic extinctions.<sup id="cite_ref-Marshall2006Explaining_54-5" class="reference"><a href="#cite_note-Marshall2006Explaining-54"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Anoxia">Anoxia</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=38" title="Edit section: Anoxia"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The late Ediacaran oceans appears to have suffered from an <a href="/wiki/Anoxic_waters" title="Anoxic waters">anoxia</a> that covered much of the seafloor, which would have given mobile animals with the ability to seek out more oxygen-rich environments an advantage over sessile forms of life.<sup id="cite_ref-156" class="reference"><a href="#cite_note-156"><span class="cite-bracket">[</span>151<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Increase_in_sensory_and_cognitive_abilities">Increase in sensory and cognitive abilities</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=39" title="Edit section: Increase in sensory and cognitive abilities"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Evolution_of_the_eye" title="Evolution of the eye">Evolution of the eye</a></div> <p><a href="/wiki/Andrew_Parker_(zoologist)" title="Andrew Parker (zoologist)">Andrew Parker</a> has proposed that predator-prey relationships changed dramatically after eyesight evolved. Prior to that time, hunting and evading were both close-range affairs—smell (chemoreception), vibration and touch were the only senses used. When predators could see their prey from a distance, new defensive strategies were needed. Armor, spines and similar defenses may also have evolved in response to vision. He further observed that, where animals lose vision in unlighted environments such as caves, diversity of animal forms tends to decrease.<sup id="cite_ref-157" class="reference"><a href="#cite_note-157"><span class="cite-bracket">[</span>152<span class="cite-bracket">]</span></a></sup> Nevertheless, many scientists doubt that vision could have caused the explosion. Eyes may well have evolved long before the start of the Cambrian.<sup id="cite_ref-McCall2006_158-0" class="reference"><a href="#cite_note-McCall2006-158"><span class="cite-bracket">[</span>153<span class="cite-bracket">]</span></a></sup> It is also difficult to understand why the evolution of eyesight would have caused an explosion, since other senses, such as smell and pressure detection, can detect things at a greater distance in the sea than sight can, but the appearance of these other senses apparently did not cause an evolutionary explosion.<sup id="cite_ref-Marshall2006Explaining_54-6" class="reference"><a href="#cite_note-Marshall2006Explaining-54"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> </p> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Life_on_the_platform_margin_of_the_Miaolingian_sea,_North_China.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f0/Life_on_the_platform_margin_of_the_Miaolingian_sea%2C_North_China.png/250px-Life_on_the_platform_margin_of_the_Miaolingian_sea%2C_North_China.png" decoding="async" width="250" height="145" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f0/Life_on_the_platform_margin_of_the_Miaolingian_sea%2C_North_China.png/500px-Life_on_the_platform_margin_of_the_Miaolingian_sea%2C_North_China.png 1.5x" data-file-width="1269" data-file-height="738" /></a><figcaption>Life on the platform margin of the <a href="/wiki/Miaolingian" title="Miaolingian">Miaolingian</a> sea</figcaption></figure> <p>One hypothesis posits that the development of increased cognitive abilities during the Cambrian drove diversity increase. This is evidenced by the fact that the novel ecological lifestyles created during the Cambrian required rapid, regular movement, a feature associated with brain-bearing organisms. The increasing complexity of brains, positively correlated with a greater range of motion and sensory abilities, enabled a wider range of novel ecological modes of life to come into being.<sup id="cite_ref-159" class="reference"><a href="#cite_note-159"><span class="cite-bracket">[</span>154<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Arms_races_between_predators_and_prey">Arms races between predators and prey</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=40" title="Edit section: Arms races between predators and prey"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The ability to avoid or recover from <a href="/wiki/Predation" title="Predation">predation</a> often makes the difference between life and death, and is therefore one of the strongest components of <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a>. The pressure to adapt is stronger on the prey than on the predator: if the predator fails to win a contest, it loses a meal; if the prey is the loser, it loses its life.<sup id="cite_ref-DawkinsKrebs1979ArmsRaces_160-0" class="reference"><a href="#cite_note-DawkinsKrebs1979ArmsRaces-160"><span class="cite-bracket">[</span>155<span class="cite-bracket">]</span></a></sup> </p><p>But, there is evidence that predation was rife long before the start of the Cambrian, for example in the increasingly spiny forms of <a href="/wiki/Acritarchs" class="mw-redirect" title="Acritarchs">acritarchs</a>, the holes drilled in <i><a href="/wiki/Cloudina" class="mw-redirect" title="Cloudina">Cloudina</a></i> shells, and traces of burrowing to avoid predators. Hence, it is unlikely that the <i>appearance</i> of predation was the trigger for the Cambrian "explosion", although it may well have exhibited a strong influence on the body forms that the "explosion" produced.<sup id="cite_ref-Bengtson2002OriginsOfPredation_64-5" class="reference"><a href="#cite_note-Bengtson2002OriginsOfPredation-64"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup> However, the intensity of predation does appear to have increased dramatically during the Cambrian<sup id="cite_ref-161" class="reference"><a href="#cite_note-161"><span class="cite-bracket">[</span>156<span class="cite-bracket">]</span></a></sup> as new predatory "tactics" (such as shell-crushing) emerged.<sup id="cite_ref-Zhang2011_162-0" class="reference"><a href="#cite_note-Zhang2011-162"><span class="cite-bracket">[</span>157<span class="cite-bracket">]</span></a></sup> This rise of predation during the Cambrian was confirmed by the temporal pattern of the median predator ratio at the scale of genus, in fossil communities covering the Cambrian and Ordovician periods, but this pattern is not correlated to diversification rate.<sup id="cite_ref-:0_163-0" class="reference"><a href="#cite_note-:0-163"><span class="cite-bracket">[</span>158<span class="cite-bracket">]</span></a></sup> This lack of correlation between predator ratio and diversification over the Cambrian and Ordovician suggests that predators did not trigger the large evolutionary radiation of animals during this interval. Thus the role of predators as triggerers of diversification may have been limited to the very beginning of the "Cambrian explosion".<sup id="cite_ref-:0_163-1" class="reference"><a href="#cite_note-:0-163"><span class="cite-bracket">[</span>158<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Increase_in_size_and_diversity_of_planktonic_animals">Increase in size and diversity of planktonic animals</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=41" title="Edit section: Increase in size and diversity of planktonic animals"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Geochemical" class="mw-redirect" title="Geochemical">Geochemical</a> evidence strongly indicates that the total mass of <a href="/wiki/Plankton" title="Plankton">plankton</a> has been similar to modern levels since early in the Proterozoic. Before the start of the Cambrian, their corpses and droppings were too small to fall quickly towards the seabed, since their <a href="/wiki/Drag_(physics)" title="Drag (physics)">drag</a> was about the same as their weight. This meant they were destroyed by <a href="/wiki/Scavenger" title="Scavenger">scavengers</a> or by chemical processes before they reached the sea floor.<sup id="cite_ref-Butterfield2001_44-1" class="reference"><a href="#cite_note-Butterfield2001-44"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> </p><p>Mesozooplankton are plankton of a larger size. Early Cambrian specimens <a href="/wiki/Filter_feeder" title="Filter feeder">filtered</a> microscopic plankton from the seawater. These larger organisms would have produced droppings and ultimately corpses large enough to fall fairly quickly. This provided a new supply of energy and nutrients to the mid-levels and bottoms of the seas, which opened up a new range of possible ways of life. If any of these remains sank uneaten to the sea floor they could be buried; this would have taken some <a href="/wiki/Carbon" title="Carbon">carbon</a> out of <a href="/wiki/Carbon_cycle" title="Carbon cycle">circulation</a>, resulting in an increase in the <a href="/wiki/Concentration" title="Concentration">concentration</a> of breathable oxygen in the seas (carbon readily <a href="/wiki/Chemical_compound" title="Chemical compound">combines</a> with oxygen).<sup id="cite_ref-Butterfield2001_44-2" class="reference"><a href="#cite_note-Butterfield2001-44"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> </p><p>The initial herbivorous mesozooplankton were probably larvae of benthic (seafloor) animals. A larval stage was probably an evolutionary innovation driven by the increasing level of predation at the seafloor during the <a href="/wiki/Ediacaran" title="Ediacaran">Ediacaran</a> period.<sup id="cite_ref-Butterfield2001ECR_11-1" class="reference"><a href="#cite_note-Butterfield2001ECR-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Peterson2005_164-0" class="reference"><a href="#cite_note-Peterson2005-164"><span class="cite-bracket">[</span>159<span class="cite-bracket">]</span></a></sup> </p><p>Metazoans have an amazing ability to increase diversity through <a href="/wiki/Coevolution" title="Coevolution">coevolution</a>.<sup id="cite_ref-Butterfield2007_66-4" class="reference"><a href="#cite_note-Butterfield2007-66"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup> This means that an organism's traits can lead to traits evolving in other organisms; a number of responses are possible, and a different species can potentially emerge from each one. As a simple example, the evolution of predation may have caused one organism to develop a defence, while another developed motion to flee. This would cause the predator lineage to diverge into two species: one that was good at chasing prey, and another that was good at breaking through defences. Actual coevolution is somewhat more subtle, but, in this fashion, great diversity can arise: three quarters of living species are animals, and most of the rest have formed by coevolution with animals.<sup id="cite_ref-Butterfield2007_66-5" class="reference"><a href="#cite_note-Butterfield2007-66"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Ecosystem_engineering">Ecosystem engineering</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=42" title="Edit section: Ecosystem engineering"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Evolving organisms inevitably change the environment they evolve in. The <a href="/wiki/Devonian" title="Devonian">Devonian</a> <a href="/wiki/Colonization_of_the_land" class="mw-redirect" title="Colonization of the land">colonization of land</a> had planet-wide consequences for sediment cycling and ocean nutrients, and was likely linked to the <a href="/wiki/Devonian_extinction" class="mw-redirect" title="Devonian extinction">Devonian mass extinction</a>. A similar process may have occurred on smaller scales in the oceans, with, for example, the sponges filtering particles from the water and depositing them in the mud in a more digestible form; or burrowing organisms making previously unavailable resources available for other organisms.<sup id="cite_ref-Erwin2011a_165-0" class="reference"><a href="#cite_note-Erwin2011a-165"><span class="cite-bracket">[</span>160<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Burrowing_2">Burrowing</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=43" title="Edit section: Burrowing"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Increases in burrowing changed the seafloor's geochemistry, and led to decreased oxygen in the ocean and increased CO<sub>2</sub> levels in the seas and the atmosphere, resulting in global warming for tens of millions years, and could be responsible for mass extinctions.<sup id="cite_ref-166" class="reference"><a href="#cite_note-166"><span class="cite-bracket">[</span>161<span class="cite-bracket">]</span></a></sup> But as burrowing became established, it allowed an explosion of its own, for as burrowers disturbed the sea floor, they aerated it, mixing oxygen into the toxic muds. This made the bottom sediments more hospitable, and allowed a wider range of organisms to inhabit them—creating new niches and the scope for higher diversity.<sup id="cite_ref-Seilacher2005_90-1" class="reference"><a href="#cite_note-Seilacher2005-90"><span class="cite-bracket">[</span>87<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Complexity_threshold">Complexity threshold</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=44" title="Edit section: Complexity threshold"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The explosion may not have been a significant evolutionary event. It may represent a threshold being crossed: for example a threshold in genetic complexity that allowed a vast range of morphological forms to be employed.<sup id="cite_ref-SoléFernándezKauffman2003AdaptiveWalks_167-0" class="reference"><a href="#cite_note-SoléFernándezKauffman2003AdaptiveWalks-167"><span class="cite-bracket">[</span>162<span class="cite-bracket">]</span></a></sup> This genetic threshold may have a correlation to the amount of oxygen available to organisms. Using oxygen for metabolism produces much more energy than anaerobic processes. Organisms that use more oxygen have the opportunity to produce more complex proteins, providing a template for further evolution.<sup id="cite_ref-ReferenceA_136-1" class="reference"><a href="#cite_note-ReferenceA-136"><span class="cite-bracket">[</span>131<span class="cite-bracket">]</span></a></sup> These proteins translate into larger, more complex structures that allow organisms better to adapt to their environments.<sup id="cite_ref-ReferenceC_168-0" class="reference"><a href="#cite_note-ReferenceC-168"><span class="cite-bracket">[</span>163<span class="cite-bracket">]</span></a></sup> With the help of oxygen, genes that code for these proteins could contribute to the expression of <a href="/wiki/Complex_traits" title="Complex traits">complex traits</a> more efficiently. Access to a wider range of structures and functions would allow organisms to evolve in different directions, increasing the number of niches that could be inhabited. Furthermore, organisms had the opportunity to become more specialized in their own niches.<sup id="cite_ref-ReferenceC_168-1" class="reference"><a href="#cite_note-ReferenceC-168"><span class="cite-bracket">[</span>163<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Relationship_with_the_Great_Ordovician_Biodiversification_Event">Relationship with the Great Ordovician Biodiversification Event</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=45" title="Edit section: Relationship with the Great Ordovician Biodiversification Event"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951" /><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Great_Ordovician_Biodiversification_Event" title="Great Ordovician Biodiversification Event">Great Ordovician Biodiversification Event</a></div> <p>After an <a href="/wiki/Cambrian%E2%80%93Ordovician_extinction_event" title="Cambrian–Ordovician extinction event">extinction</a> at the Cambrian–Ordovician boundary, another radiation occurred, which established the taxa that would dominate the Palaeozoic. This event, known as the Great Ordovician Biodiversification Event (GOBE), has been considered a "follow-up" to the Cambrian explosion.<sup id="cite_ref-Droser2003_169-0" class="reference"><a href="#cite_note-Droser2003-169"><span class="cite-bracket">[</span>164<span class="cite-bracket">]</span></a></sup> Recent studies have suggested that the Cambrian explosion were not two discrete events but one long evolutionary radiation.<sup id="cite_ref-170" class="reference"><a href="#cite_note-170"><span class="cite-bracket">[</span>165<span class="cite-bracket">]</span></a></sup> Analytical study of the Geobiodiversity Database (GBDB) and <a href="/wiki/Paleobiology_Database" title="Paleobiology Database">Paleobiology Database</a> (PBDB) failed to find a statistical basis for separating the two radiations.<sup id="cite_ref-171" class="reference"><a href="#cite_note-171"><span class="cite-bracket">[</span>166<span class="cite-bracket">]</span></a></sup> </p><p>Some researchers have proposed the existence of a biodiversity gap during the <a href="/wiki/Furongian" title="Furongian">Furongian</a> separating the Cambrian explosion and GOBE known as the Furongian Gap.<sup id="cite_ref-172" class="reference"><a href="#cite_note-172"><span class="cite-bracket">[</span>167<span class="cite-bracket">]</span></a></sup> Studies of the Guole Konservat-Lagerstätte and similar fossil sites in South China have instead found the Furongian to instead be a time of rapid biological turnovers though, making the existence of the Furongian Gap highly controversial.<sup id="cite_ref-173" class="reference"><a href="#cite_note-173"><span class="cite-bracket">[</span>168<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Uniqueness_of_the_early_Cambrian_biodiversification">Uniqueness of the early Cambrian biodiversification</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=46" title="Edit section: Uniqueness of the early Cambrian biodiversification"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The "Cambrian explosion" can be viewed as two waves of metazoan expansion into empty niches: first, a <a href="/wiki/Coevolution" title="Coevolution">coevolutionary</a> rise in diversity as animals explored niches on the Ediacaran sea floor, followed by a second expansion in the early Cambrian as they became established in the <a href="/wiki/Water_column" title="Water column">water column</a>.<sup id="cite_ref-Butterfield2007_66-6" class="reference"><a href="#cite_note-Butterfield2007-66"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup> The rate of diversification seen in the Cambrian phase of the explosion is unparalleled among marine animals: it affected all metazoan <a href="/wiki/Clade" title="Clade">clades</a> of which Cambrian fossils have been found. Later <a href="/wiki/Evolutionary_radiation" title="Evolutionary radiation">radiations</a>, such as those of <a href="/wiki/Fish" title="Fish">fish</a> in the <a href="/wiki/Silurian" title="Silurian">Silurian</a> and <a href="/wiki/Devonian" title="Devonian">Devonian</a> periods, involved fewer <a href="/wiki/Taxon" title="Taxon">taxa</a>, mainly with very similar body plans.<sup id="cite_ref-CowenHistLife_25-5" class="reference"><a href="#cite_note-CowenHistLife-25"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> Although the recovery from the <a href="/wiki/Permian%E2%80%93Triassic_extinction_event" title="Permian–Triassic extinction event">Permian-Triassic extinction</a> started with about as few animal species as the Cambrian explosion, the recovery produced far fewer significantly new types of animals.<sup id="cite_ref-174" class="reference"><a href="#cite_note-174"><span class="cite-bracket">[</span>169<span class="cite-bracket">]</span></a></sup> </p><p>Whatever triggered the early Cambrian diversification opened up an exceptionally wide range of previously unavailable <a href="/wiki/Ecological_niche" title="Ecological niche">ecological niches</a>. When these were all occupied, limited space existed for such wide-ranging diversifications to occur again, because strong competition existed in all niches and <a href="https://en.wiktionary.org/wiki/incumbent" class="extiw" title="wikt:incumbent">incumbents</a> usually had the advantage. If a wide range of empty niches had continued, clades would be able to continue diversifying and become disparate enough for us to recognise them as different <a href="/wiki/Phylum" title="Phylum">phyla</a>; when niches are filled, lineages will continue to resemble one another long after they diverge, as limited opportunity exists for them to change their life-styles and forms.<sup id="cite_ref-Valentine1995WhyNoNewPhyla_175-0" class="reference"><a href="#cite_note-Valentine1995WhyNoNewPhyla-175"><span class="cite-bracket">[</span>170<span class="cite-bracket">]</span></a></sup> </p><p>There were two similar explosions in the <a href="/wiki/Evolutionary_history_of_plants#Colonization_of_land" title="Evolutionary history of plants">evolution of land plants</a>: after a cryptic history beginning about <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=450">450</a> million years ago</span>, land plants underwent a uniquely rapid adaptive radiation during the Devonian period, about <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=400">400</a> million years ago</span>.<sup id="cite_ref-CowenHistLife_25-6" class="reference"><a href="#cite_note-CowenHistLife-25"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> Furthermore, angiosperms (<a href="/wiki/Flowering_plant" title="Flowering plant">flowering plants</a>) originated and rapidly diversified during the <a href="/wiki/Cretaceous" title="Cretaceous">Cretaceous</a> period. </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=47" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Avalon_explosion" title="Avalon explosion">Avalon explosion</a>, a similar biodiversity of life that occurred 33 million years prior.</li></ul> <div class="mw-heading mw-heading2"><h2 id="Footnotes">Footnotes</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=48" title="Edit section: Footnotes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width reflist-lower-alpha"> <ol class="references"> <li id="cite_note-12"><span class="mw-cite-backlink"><b><a href="#cite_ref-12">^</a></b></span> <span class="reference-text">This included at least animals, <a href="/wiki/Phytoplankton" title="Phytoplankton">phytoplankton</a> and <a href="/wiki/Calcimicrobe" title="Calcimicrobe">calcimicrobes</a>.<sup id="cite_ref-Butterfield2001ECR_11-0" class="reference"><a href="#cite_note-Butterfield2001ECR-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup></span> </li> <li id="cite_note-13"><span class="mw-cite-backlink"><b><a href="#cite_ref-13">^</a></b></span> <span class="reference-text">At 610 million years ago, <i><a href="/wiki/Aspidella" title="Aspidella">Aspidella</a></i> disks appeared, but it is not clear that these represented complex life forms.</span> </li> <li id="cite_note-78"><span class="mw-cite-backlink"><b><a href="#cite_ref-78">^</a></b></span> <span class="reference-text">Older marks found in billion-year-old rocks<sup id="cite_ref-Seilacher1998_76-0" class="reference"><a href="#cite_note-Seilacher1998-76"><span class="cite-bracket">[</span>74<span class="cite-bracket">]</span></a></sup> have since been recognised as nonbiogenic.<sup id="cite_ref-Budd2000_15-1" class="reference"><a href="#cite_note-Budd2000-15"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Jensen2003_77-0" class="reference"><a href="#cite_note-Jensen2003-77"><span class="cite-bracket">[</span>75<span class="cite-bracket">]</span></a></sup></span> </li> <li id="cite_note-125"><span class="mw-cite-backlink"><b><a href="#cite_ref-125">^</a></b></span> <span class="reference-text">As defined in terms of the extinction and origination rate of species.<sup id="cite_ref-Butterfield2007_66-3" class="reference"><a href="#cite_note-Butterfield2007-66"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup></span> </li> <li id="cite_note-127"><span class="mw-cite-backlink"><b><a href="#cite_ref-127">^</a></b></span> <span class="reference-text">The analysis considered the bioprovinciality of trilobite lineages, as well as their evolutionary rate.<sup id="cite_ref-Lieberman2003_126-0" class="reference"><a href="#cite_note-Lieberman2003-126"><span class="cite-bracket">[</span>122<span class="cite-bracket">]</span></a></sup></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=49" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239543626" /><div class="reflist reflist-columns references-column-width" style="column-width: 25em;"> <ol class="references"> <li id="cite_note-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFZhuravlevRiding2000" class="citation book cs1">Zhuravlev, Andrey; Riding, Robert (2000). <a rel="nofollow" class="external text" href="https://cup.columbia.edu/book/the-ecology-of-the-cambrian-radiation/9780231106139"><i>The Ecology of the Cambrian Radiation</i></a>. <a href="/wiki/Columbia_University_Press" title="Columbia University Press">Columbia University Press</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-231-10613-9" title="Special:BookSources/978-0-231-10613-9"><bdi>978-0-231-10613-9</bdi></a>. <q>The Cambrian radiation was the explosive evolution of marine life that started 550,000,000 years ago. It ranks as one of the most important episodes in Earth history. 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(1989). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20110824010939/http://palaeontology.palass-pubs.org/pdf/Vol%2032/Pages%2069-99.pdf">"A new mitrate from the Upper Ordovician of Norway, and a new approach to subdividing a plesion"</a> <span class="cs1-format">(PDF)</span>. <i>Palaeontology</i>. <b>32</b>: <span class="nowrap">69–</span>99. Archived from <a rel="nofollow" class="external text" href="http://palaeontology.palass-pubs.org/pdf/Vol%2032/Pages%2069-99.pdf">the original</a> <span class="cs1-format">(PDF)</span> on 2011-08-24<span class="reference-accessdate">. Retrieved <span class="nowrap">2010-12-25</span></span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Palaeontology&rft.atitle=A+new+mitrate+from+the+Upper+Ordovician+of+Norway%2C+and+a+new+approach+to+subdividing+a+plesion&rft.volume=32&rft.pages=%3Cspan+class%3D%22nowrap%22%3E69-%3C%2Fspan%3E99&rft.date=1989&rft.au=Craske%2C+A.J.&rft.au=Jefferies%2C+R.P.S.&rft_id=http%3A%2F%2Fpalaeontology.palass-pubs.org%2Fpdf%2FVol%252032%2FPages%252069-99.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACambrian+explosion" class="Z3988"></span></span> </li> <li id="cite_note-59"><span class="mw-cite-backlink"><b><a href="#cite_ref-59">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFValentine2004" class="citation book cs1">Valentine, James W. (2004). <i>On the Origin of Phyla</i>. Chicago: University Of Chicago Press. p. 7. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-226-84548-7" title="Special:BookSources/978-0-226-84548-7"><bdi>978-0-226-84548-7</bdi></a>.</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+Phyla&rft.place=Chicago&rft.pages=7&rft.pub=University+Of+Chicago+Press&rft.date=2004&rft.isbn=978-0-226-84548-7&rft.aulast=Valentine&rft.aufirst=James+W.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACambrian+explosion" class="Z3988"></span>"<cite>Classifications of organisms in hierarchical systems were in use by the 17th and 18th centuries. Usually, organisms were grouped according to their morphological similarities as perceived by those early workers, and those groups were then grouped according to their similarities, and so on, to form a hierarchy.</cite>"</span> </li> <li id="cite_note-60"><span class="mw-cite-backlink"><b><a href="#cite_ref-60">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFParker2003" class="citation book cs1">Parker, Andrew (2003). <i>In the blink of an eye: How vision kick-started the big bang of evolution</i>. 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E.; Jensen, J. (2000). "A critical reappraisal of the fossil record of the bilaterian phyla". <i>Biological Reviews</i>. <b>75</b> (2): <span class="nowrap">253–</span>295. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fj.1469-185X.1999.tb00046.x">10.1111/j.1469-185X.1999.tb00046.x</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/10881389">10881389</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:39772232">39772232</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Biological+Reviews&rft.atitle=A+critical+reappraisal+of+the+fossil+record+of+the+bilaterian+phyla&rft.volume=75&rft.issue=2&rft.pages=%3Cspan+class%3D%22nowrap%22%3E253-%3C%2Fspan%3E295&rft.date=2000&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A39772232%23id-name%3DS2CID&rft_id=info%3Apmid%2F10881389&rft_id=info%3Adoi%2F10.1111%2Fj.1469-185X.1999.tb00046.x&rft.au=Budd%2C+G.+E.&rft.au=Jensen%2C+J.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACambrian+explosion" class="Z3988"></span></li> <li>Collins, Allen G. <a rel="nofollow" class="external text" href="http://www.ucmp.berkeley.edu/phyla/metazoafr.html">"Metazoa: Fossil record"</a>. Retrieved Dec. 14, 2005.</li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFConway_Morris,_S.1997" class="citation book cs1">Conway Morris, S. (1997). <i>The Crucible of Creation: the Burgess Shale and the rise of animals</i>. Oxford University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/0-19-286202-2" title="Special:BookSources/0-19-286202-2"><bdi>0-19-286202-2</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Crucible+of+Creation%3A+the+Burgess+Shale+and+the+rise+of+animals&rft.pub=Oxford+University+Press&rft.date=1997&rft.isbn=0-19-286202-2&rft.au=Conway+Morris%2C+S.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACambrian+explosion" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFConway_Morris,_S.2006" class="citation journal cs1">Conway Morris, S. 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Fedonkin">Fedonkin, M.A</a>; Kirschvink, J.L (2000). "Age of Neoproterozoic Bilaterian Body and Trace Fossils, White Sea, Russia: Implications for Metazoan Evolution". <i>Science</i>. <b>288</b> (5467): <span class="nowrap">841–</span>845. <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/2000Sci...288..841M">2000Sci...288..841M</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1126%2Fscience.288.5467.841">10.1126/science.288.5467.841</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/10797002">10797002</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Science&rft.atitle=Age+of+Neoproterozoic+Bilaterian+Body+and+Trace+Fossils%2C+White+Sea%2C+Russia%3A+Implications+for+Metazoan+Evolution&rft.volume=288&rft.issue=5467&rft.pages=%3Cspan+class%3D%22nowrap%22%3E841-%3C%2Fspan%3E845&rft.date=2000&rft_id=info%3Apmid%2F10797002&rft_id=info%3Adoi%2F10.1126%2Fscience.288.5467.841&rft_id=info%3Abibcode%2F2000Sci...288..841M&rft.au=Martin%2C+M.W&rft.au=Grazhdankin%2C+D.V&rft.au=Bowring%2C+S.A&rft.au=Evans%2C+D.A.D&rft.au=Fedonkin%2C+M.A&rft.au=Kirschvink%2C+J.L&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACambrian+explosion" class="Z3988"></span></li></ul> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cambrian_explosion&action=edit&section=51" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20120213225957/http://www.ijdb.ehu.es/web/contents.php?vol=47">The Cambrian "explosion" of metazoans and molecular biology: would Darwin be satisfied?</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20080407005007/http://genome6.cu-genome.org/andrey/GouldComment.pdf">On embryos and ancestors</a> by <a href="/wiki/Stephen_Jay_Gould" title="Stephen Jay Gould">Stephen Jay Gould</a></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFConway_Morris2000" class="citation journal cs1">Conway Morris, S. (April 2000). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC34314">"The Cambrian "explosion": Slow-fuse or megatonnage?"</a>. <i>Proceedings of the National Academy of Sciences</i>. <b>97</b> (9): <span class="nowrap">4426–</span>4429. <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/2000PNAS...97.4426C">2000PNAS...97.4426C</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.97.9.4426">10.1073/pnas.97.9.4426</a></span>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC34314">34314</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/10781036">10781036</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+National+Academy+of+Sciences&rft.atitle=The+Cambrian+%22explosion%22%3A+Slow-fuse+or+megatonnage%3F&rft.volume=97&rft.issue=9&rft.pages=%3Cspan+class%3D%22nowrap%22%3E4426-%3C%2Fspan%3E4429&rft.date=2000-04&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC34314%23id-name%3DPMC&rft_id=info%3Apmid%2F10781036&rft_id=info%3Adoi%2F10.1073%2Fpnas.97.9.4426&rft_id=info%3Abibcode%2F2000PNAS...97.4426C&rft.aulast=Conway+Morris&rft.aufirst=S.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC34314&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACambrian+explosion" class="Z3988"></span></li> <li><a rel="nofollow" class="external text" href="https://www.bbc.co.uk/programmes/p003k9bg">The Cambrian Explosion</a> – <a rel="nofollow" class="external text" href="https://www.bbc.co.uk/programmes/b006qykl"><i>In Our Time</i></a>, <a href="/wiki/BBC_Radio_4" title="BBC Radio 4">BBC Radio 4</a> broadcast, 17 February 2005</li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://burgess-shale.rom.on.ca/en">"Burgess Shale"</a>. Virtual Museum of Canada. 2011.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Burgess+Shale&rft.pub=Virtual+Museum+of+Canada&rft.date=2011&rft_id=http%3A%2F%2Fburgess-shale.rom.on.ca%2Fen&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACambrian+explosion" class="Z3988"></span>, exhaustive details about the Burgess Shale, its fossils and its significance for the Cambrian explosion</li> <li><a rel="nofollow" class="external text" href="http://www.kumip.ku.edu/cambrianlife/">Utah's Cambrian life</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20120723072953/http://www.kumip.ku.edu/cambrianlife/">Archived</a> 2012-07-23 at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a> – new (2008) website with good images of a range of Burgess-shale-type and other Cambrian fossils</li> <li><a rel="nofollow" class="external text" href="http://paleobiology.si.edu/geotime/main/htmlversion/cambrian2.html">Smithsonian National Museum</a></li></ul>    </div><noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?useformat=desktop&type=1x1&usesul3=0" alt="" width="1" height="1" style="border: none; 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