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Cowles</span> </div> </a> <ul id="toc-H._C._Cowles-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Gleason_and_Clements" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Gleason_and_Clements"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.2</span> <span>Gleason and Clements</span> </div> </a> <ul id="toc-Gleason_and_Clements-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Eugene_Odum" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Eugene_Odum"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.3</span> <span>Eugene Odum</span> </div> </a> <ul id="toc-Eugene_Odum-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Modern_era" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Modern_era"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.4</span> <span>Modern era</span> </div> </a> <ul id="toc-Modern_era-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Factors" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Factors"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Factors</span> </div> </a> <button aria-controls="toc-Factors-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 Factors subsection</span> </button> <ul id="toc-Factors-sublist" class="vector-toc-list"> <li id="toc-Diversity_of_possible_trajectories" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Diversity_of_possible_trajectories"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>Diversity of possible trajectories</span> </div> </a> <ul id="toc-Diversity_of_possible_trajectories-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Stochastic_events" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Stochastic_events"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>Stochastic events</span> </div> </a> <ul id="toc-Stochastic_events-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Dispersal_limitation_vs._environmental_filtering" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Dispersal_limitation_vs._environmental_filtering"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3</span> <span>Dispersal limitation vs. environmental filtering</span> </div> </a> <ul id="toc-Dispersal_limitation_vs._environmental_filtering-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Feedback_loops" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Feedback_loops"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.4</span> <span>Feedback loops</span> </div> </a> <ul id="toc-Feedback_loops-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Patterns" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Patterns"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.5</span> <span>Patterns</span> </div> </a> <ul id="toc-Patterns-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Disruptions" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Disruptions"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.6</span> <span>Disruptions</span> </div> </a> <ul id="toc-Disruptions-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Types_of_succession" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Types_of_succession"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Types of succession</span> </div> </a> <button aria-controls="toc-Types_of_succession-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 Types of succession subsection</span> </button> <ul id="toc-Types_of_succession-sublist" class="vector-toc-list"> <li id="toc-Primary_succession" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Primary_succession"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>Primary succession</span> </div> </a> <ul id="toc-Primary_succession-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Secondary_succession" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Secondary_succession"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Secondary succession</span> </div> </a> <ul id="toc-Secondary_succession-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Seasonal_and_cyclic_dynamics" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Seasonal_and_cyclic_dynamics"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.3</span> <span>Seasonal and cyclic dynamics</span> </div> </a> <ul id="toc-Seasonal_and_cyclic_dynamics-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Causes_of_plant_succession" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Causes_of_plant_succession"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.4</span> <span>Causes of plant succession</span> </div> </a> <ul id="toc-Causes_of_plant_succession-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Mechanisms" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Mechanisms"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.5</span> <span>Mechanisms</span> </div> </a> <ul id="toc-Mechanisms-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Seral_communities" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Seral_communities"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Seral communities</span> </div> </a> <ul id="toc-Seral_communities-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Changes_in_animal_life" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Changes_in_animal_life"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Changes in animal life</span> </div> </a> <ul id="toc-Changes_in_animal_life-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Microsuccession" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Microsuccession"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Microsuccession</span> </div> </a> <ul id="toc-Microsuccession-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Climax_concept" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Climax_concept"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Climax concept</span> </div> </a> <button aria-controls="toc-Climax_concept-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 Climax concept subsection</span> </button> <ul id="toc-Climax_concept-sublist" class="vector-toc-list"> <li id="toc-Climax_community" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Climax_community"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.1</span> <span>Climax community</span> </div> </a> <ul id="toc-Climax_community-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Characteristics" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Characteristics"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.2</span> <span>Characteristics</span> </div> </a> <ul id="toc-Characteristics-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Types_of_climax" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Types_of_climax"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.3</span> <span>Types of climax</span> </div> </a> <ul id="toc-Types_of_climax-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Theories" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Theories"> <div class="vector-toc-text"> <span class="vector-toc-numb">7.4</span> <span>Theories</span> </div> </a> <ul id="toc-Theories-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Succession_by_habitat_type" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Succession_by_habitat_type"> <div class="vector-toc-text"> <span class="vector-toc-numb">8</span> <span>Succession by habitat type</span> </div> </a> <button aria-controls="toc-Succession_by_habitat_type-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 Succession by habitat type subsection</span> </button> <ul id="toc-Succession_by_habitat_type-sublist" class="vector-toc-list"> <li id="toc-Forest_succession" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Forest_succession"> <div class="vector-toc-text"> <span class="vector-toc-numb">8.1</span> <span>Forest succession</span> </div> </a> <ul id="toc-Forest_succession-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Wetland_succession" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Wetland_succession"> <div class="vector-toc-text"> <span class="vector-toc-numb">8.2</span> <span>Wetland succession</span> </div> </a> <ul id="toc-Wetland_succession-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Grassland_succession" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Grassland_succession"> <div class="vector-toc-text"> <span class="vector-toc-numb">8.3</span> <span>Grassland succession</span> </div> </a> <ul id="toc-Grassland_succession-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> <span class="vector-toc-numb">9</span> <span>See also</span> </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">10</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">11</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">12</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">Ecological succession</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 51 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-51" 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">51 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-af mw-list-item"><a href="https://af.wikipedia.org/wiki/Ekologiese_opvolging" title="Ekologiese opvolging – Afrikaans" lang="af" hreflang="af" data-title="Ekologiese opvolging" 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%AA%D8%B9%D8%A7%D9%82%D8%A8_%D8%A8%D9%8A%D8%A6%D9%8A" title="تعاقب بيئي – Arabic" lang="ar" hreflang="ar" data-title="تعاقب بيئي" 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/Suksessiya" title="Suksessiya – Azerbaijani" lang="az" hreflang="az" data-title="Suksessiya" data-language-autonym="Azərbaycanca" data-language-local-name="Azerbaijani" class="interlanguage-link-target"><span>Azərbaycanca</span></a></li><li class="interlanguage-link interwiki-be mw-list-item"><a href="https://be.wikipedia.org/wiki/%D0%A1%D1%83%D0%BA%D1%86%D1%8D%D1%81%D1%96%D1%8F" title="Сукцэсія – Belarusian" lang="be" hreflang="be" data-title="Сукцэсія" data-language-autonym="Беларуская" data-language-local-name="Belarusian" class="interlanguage-link-target"><span>Беларуская</span></a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%A1%D1%83%D0%BA%D1%86%D0%B5%D1%81%D0%B8%D1%8F" title="Сукцесия – Bulgarian" lang="bg" hreflang="bg" data-title="Сукцесия" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target"><span>Български</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Successi%C3%B3_vegetal" title="Successió vegetal – Catalan" lang="ca" hreflang="ca" data-title="Successió vegetal" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Sukcese_(ekologie)" title="Sukcese (ekologie) – Czech" lang="cs" hreflang="cs" data-title="Sukcese (ekologie)" 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/Succession_(%C3%B8kologi)" title="Succession (økologi) – Danish" lang="da" hreflang="da" data-title="Succession (økologi)" 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/Sukzession_(Biologie)" title="Sukzession (Biologie) – German" lang="de" hreflang="de" data-title="Sukzession (Biologie)" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/Suktsessioon" title="Suktsessioon – Estonian" lang="et" hreflang="et" data-title="Suktsessioon" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target"><span>Eesti</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Sucesi%C3%B3n_ecol%C3%B3gica" title="Sucesión ecológica – Spanish" lang="es" hreflang="es" data-title="Sucesión ecológica" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/Sukcesio_(ekologio)" title="Sukcesio (ekologio) – Esperanto" lang="eo" hreflang="eo" data-title="Sukcesio (ekologio)" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target"><span>Esperanto</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Segida_ekologiko" title="Segida ekologiko – Basque" lang="eu" hreflang="eu" data-title="Segida ekologiko" 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%AA%D9%88%D8%A7%D9%84%DB%8C_%D8%A8%D9%88%D9%85%E2%80%8C%D8%B4%D9%86%D8%A7%D8%AE%D8%AA%DB%8C" title="توالی بوم‌شناختی – Persian" lang="fa" hreflang="fa" data-title="توالی بوم‌شناختی" 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/Succession_%C3%A9cologique" title="Succession écologique – French" lang="fr" hreflang="fr" data-title="Succession écologique" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-ga mw-list-item"><a href="https://ga.wikipedia.org/wiki/Comharbas_f%C3%A1sra" title="Comharbas fásra – Irish" lang="ga" hreflang="ga" data-title="Comharbas fásra" data-language-autonym="Gaeilge" data-language-local-name="Irish" class="interlanguage-link-target"><span>Gaeilge</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Sucesi%C3%B3n_ecol%C3%B3xica" title="Sucesión ecolóxica – Galician" lang="gl" hreflang="gl" data-title="Sucesión ecolóxica" 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%B2%9C%EC%9D%B4_(%EC%83%9D%EB%AC%BC%ED%95%99)" title="천이 (생물학) – Korean" lang="ko" hreflang="ko" data-title="천이 (생물학)" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target"><span>한국어</span></a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D5%80%D5%A1%D5%BB%D5%B8%D6%80%D5%A4%D5%A1%D5%AF%D5%A1%D5%B6%D5%B8%D6%82%D5%A9%D5%B5%D5%B8%D6%82%D5%B6_(%D5%A7%D5%AF%D5%B8%D5%AC%D5%B8%D5%A3%D5%AB%D5%A1)" title="Հաջորդականություն (էկոլոգիա) – Armenian" lang="hy" hreflang="hy" data-title="Հաջորդականություն (էկոլոգիա)" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target"><span>Հայերեն</span></a></li><li class="interlanguage-link interwiki-hi mw-list-item"><a href="https://hi.wikipedia.org/wiki/%E0%A4%AA%E0%A4%BE%E0%A4%B0%E0%A4%BF%E0%A4%B8%E0%A5%8D%E0%A4%A5%E0%A4%BF%E0%A4%A4%E0%A4%BF%E0%A4%95_%E0%A4%85%E0%A4%A8%E0%A5%81%E0%A4%95%E0%A5%8D%E0%A4%B0%E0%A4%AE" title="पारिस्थितिक अनुक्रम – Hindi" lang="hi" hreflang="hi" data-title="पारिस्थितिक अनुक्रम" data-language-autonym="हिन्दी" data-language-local-name="Hindi" class="interlanguage-link-target"><span>हिन्दी</span></a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Ekolo%C5%A1ka_sukcesija" title="Ekološka sukcesija – Croatian" lang="hr" hreflang="hr" data-title="Ekološka sukcesija" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target"><span>Hrvatski</span></a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Suksesi_ekologi" title="Suksesi ekologi – Indonesian" lang="id" hreflang="id" data-title="Suksesi ekologi" 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/Successione_(ecologia)" title="Successione (ecologia) – Italian" lang="it" hreflang="it" data-title="Successione (ecologia)" 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%A1%D7%95%D7%A7%D7%A6%D7%A1%D7%99%D7%94_(%D7%90%D7%A7%D7%95%D7%9C%D7%95%D7%92%D7%99%D7%94)" title="סוקצסיה (אקולוגיה) – Hebrew" lang="he" hreflang="he" data-title="סוקצסיה (אקולוגיה)" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-jv mw-list-item"><a href="https://jv.wikipedia.org/wiki/Suks%C3%A8si_(Biologi)" title="Suksèsi (Biologi) – Javanese" lang="jv" hreflang="jv" data-title="Suksèsi (Biologi)" data-language-autonym="Jawa" data-language-local-name="Javanese" class="interlanguage-link-target"><span>Jawa</span></a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D0%AD%D0%BA%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8%D1%8F%D0%BB%D1%8B%D2%9B_%D1%81%D1%83%D0%BA%D1%86%D0%B5%D1%81%D1%81%D0%B8%D1%8F" title="Экологиялық сукцессия – Kazakh" lang="kk" hreflang="kk" data-title="Экологиялық сукцессия" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target"><span>Қазақша</span></a></li><li class="interlanguage-link interwiki-ht mw-list-item"><a href="https://ht.wikipedia.org/wiki/Siksesyon_ekolojik" title="Siksesyon ekolojik – Haitian Creole" lang="ht" hreflang="ht" data-title="Siksesyon ekolojik" data-language-autonym="Kreyòl ayisyen" data-language-local-name="Haitian Creole" class="interlanguage-link-target"><span>Kreyòl ayisyen</span></a></li><li class="interlanguage-link interwiki-ky mw-list-item"><a href="https://ky.wikipedia.org/wiki/%D0%A1%D1%83%D0%BA%D1%86%D0%B5%D1%81%D1%81%D0%B8%D1%8F" title="Сукцессия – Kyrgyz" lang="ky" hreflang="ky" data-title="Сукцессия" data-language-autonym="Кыргызча" data-language-local-name="Kyrgyz" class="interlanguage-link-target"><span>Кыргызча</span></a></li><li class="interlanguage-link interwiki-lv mw-list-item"><a href="https://lv.wikipedia.org/wiki/Ekolo%C4%A3isk%C4%81_sukcesija" title="Ekoloģiskā sukcesija – Latvian" lang="lv" hreflang="lv" data-title="Ekoloģiskā sukcesija" data-language-autonym="Latviešu" data-language-local-name="Latvian" class="interlanguage-link-target"><span>Latviešu</span></a></li><li class="interlanguage-link interwiki-lt mw-list-item"><a href="https://lt.wikipedia.org/wiki/Ekologin%C4%97_sukcesija" title="Ekologinė sukcesija – Lithuanian" lang="lt" hreflang="lt" data-title="Ekologinė sukcesija" 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/Szukcesszi%C3%B3" title="Szukcesszió – Hungarian" lang="hu" hreflang="hu" data-title="Szukcesszió" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-mk mw-list-item"><a href="https://mk.wikipedia.org/wiki/%D0%A1%D1%83%D0%BA%D1%86%D0%B5%D1%81%D0%B8%D1%98%D0%B0" title="Сукцесија – Macedonian" lang="mk" hreflang="mk" data-title="Сукцесија" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target"><span>Македонски</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Successie_(ecologie)" title="Successie (ecologie) – Dutch" lang="nl" hreflang="nl" data-title="Successie (ecologie)" 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/%E9%81%B7%E7%A7%BB_(%E7%94%9F%E7%89%A9%E5%AD%A6)" title="遷移 (生物学) – Japanese" lang="ja" hreflang="ja" data-title="遷移 (生物学)" 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/Suksesjon" title="Suksesjon – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Suksesjon" 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/%C3%98kologisk_suksesjon" title="Økologisk suksesjon – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="Økologisk suksesjon" 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-ps mw-list-item"><a href="https://ps.wikipedia.org/wiki/%D9%86%D8%A8%D8%A7%D8%AA%D9%8A_%D8%AA%D8%B3%D9%84%D8%B3%D9%84" title="نباتي تسلسل – Pashto" lang="ps" hreflang="ps" data-title="نباتي تسلسل" data-language-autonym="پښتو" data-language-local-name="Pashto" class="interlanguage-link-target"><span>پښتو</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Sukcesja_ekologiczna" title="Sukcesja ekologiczna – Polish" lang="pl" hreflang="pl" data-title="Sukcesja ekologiczna" 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/Sucess%C3%A3o_ecol%C3%B3gica" title="Sucessão ecológica – Portuguese" lang="pt" hreflang="pt" data-title="Sucessão ecológica" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%A1%D1%83%D0%BA%D1%86%D0%B5%D1%81%D1%81%D0%B8%D1%8F" title="Сукцессия – Russian" lang="ru" hreflang="ru" data-title="Сукцессия" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-sq mw-list-item"><a href="https://sq.wikipedia.org/wiki/Suksesionet" title="Suksesionet – Albanian" lang="sq" hreflang="sq" data-title="Suksesionet" data-language-autonym="Shqip" data-language-local-name="Albanian" class="interlanguage-link-target"><span>Shqip</span></a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Ecological_succession" title="Ecological succession – Simple English" lang="en-simple" hreflang="en-simple" data-title="Ecological succession" 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/Sukcesia_(ekol%C3%B3gia)" title="Sukcesia (ekológia) – Slovak" lang="sk" hreflang="sk" data-title="Sukcesia (ekológia)" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target"><span>Slovenčina</span></a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Ekolo%C5%A1ka_sukcesija" title="Ekološka sukcesija – Slovenian" lang="sl" hreflang="sl" data-title="Ekološka sukcesija" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target"><span>Slovenščina</span></a></li><li class="interlanguage-link interwiki-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%95%D0%BA%D0%BE%D0%BB%D0%BE%D1%88%D0%BA%D0%B0_%D1%81%D1%83%D0%BA%D1%86%D0%B5%D1%81%D0%B8%D1%98%D0%B0" title="Еколошка сукцесија – Serbian" lang="sr" hreflang="sr" data-title="Еколошка сукцесија" 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-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Ekologinen_sukkessio" title="Ekologinen sukkessio – Finnish" lang="fi" hreflang="fi" data-title="Ekologinen sukkessio" 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/Ekologisk_succession" title="Ekologisk succession – Swedish" lang="sv" hreflang="sv" data-title="Ekologisk succession" 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/S%C3%BCksesyon" title="Süksesyon – Turkish" lang="tr" hreflang="tr" data-title="Süksesyon" 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%A1%D1%83%D0%BA%D1%86%D0%B5%D1%81%D1%96%D1%8F" title="Сукцесія – Ukrainian" lang="uk" hreflang="uk" data-title="Сукцесія" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/Di%E1%BB%85n_th%E1%BA%BF_sinh_th%C3%A1i" title="Diễn thế sinh thái – Vietnamese" lang="vi" hreflang="vi" data-title="Diễn thế sinh 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src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d1/Boreal_pine_forest_after_fire.JPG/300px-Boreal_pine_forest_after_fire.JPG" decoding="async" width="300" height="119" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d1/Boreal_pine_forest_after_fire.JPG/450px-Boreal_pine_forest_after_fire.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d1/Boreal_pine_forest_after_fire.JPG/600px-Boreal_pine_forest_after_fire.JPG 2x" data-file-width="5800" data-file-height="2304" /></a><figcaption>Succession after <a href="/wiki/Disturbance_(ecology)" title="Disturbance (ecology)">disturbance</a>: a <a href="/wiki/Boreal_forest" class="mw-redirect" title="Boreal forest">boreal forest</a> one year (left) and two years (right) after a <a href="/wiki/Wildfire" title="Wildfire">wildfire</a>.</figcaption></figure> <p><b>Ecological succession</b> is the process of change in the <a href="/wiki/Species" title="Species">species</a> that make up an <a href="/wiki/Community_(ecology)" title="Community (ecology)">ecological community</a> over time. </p><p>The process of succession occurs either after the initial <a href="/wiki/Colonisation_(biology)" title="Colonisation (biology)">colonization</a> of a newly created habitat, or after a <a href="/wiki/Disturbance_(ecology)" title="Disturbance (ecology)">disturbance</a> substantially alters a pre-existing habitat.<sup id="cite_ref-Fisher-2018_1-0" class="reference"><a href="#cite_note-Fisher-2018-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Succession that begins in new habitats, uninfluenced by pre-existing communities, is called <a href="/wiki/Primary_succession" title="Primary succession">primary succession</a>, whereas succession that follows disruption of a pre-existing community is called <a href="/wiki/Secondary_succession" title="Secondary succession">secondary succession</a>.<sup id="cite_ref-Fisher-2018_1-1" class="reference"><a href="#cite_note-Fisher-2018-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Primary succession may happen after a <a href="/wiki/Lava_flow" class="mw-redirect" title="Lava flow">lava flow</a> or the emergence of a new <a href="/wiki/Island" title="Island">island</a> from the ocean. <a href="/wiki/Surtsey" title="Surtsey">Surtsey</a>, a volcanic island off the southern coast of <a href="/wiki/Iceland" title="Iceland">Iceland</a>, is an important example of a place where primary succession has been observed.<sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><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> On the other hand, secondary succession happens after <a href="/wiki/Disturbance_(ecology)" title="Disturbance (ecology)">disturbance</a> of a community, such as from a <a href="/wiki/Fire" title="Fire">fire</a>, severe <a href="/wiki/Windthrow" title="Windthrow">windthrow</a>, or <a href="/wiki/Logging" title="Logging">logging</a>. </p><p>Succession was among the first theories advanced in <a href="/wiki/Ecology" title="Ecology">ecology</a>. Ecological succession was first documented in the <a href="/wiki/Indiana_Dunes" class="mw-redirect" title="Indiana Dunes">Indiana Dunes</a> of Northwest Indiana and remains an important <a href="/wiki/Ecology" title="Ecology">ecological</a> topic of study.<sup id="cite_ref-southshorejournal.org_4-0" class="reference"><a href="#cite_note-southshorejournal.org-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Over time, the understanding of succession has changed from a linear progression to a stable climax state, to a more complex, cyclical model that de-emphasizes the idea of organisms having fixed roles or relationships.<sup id="cite_ref-middleton_5-0" class="reference"><a href="#cite_note-middleton-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="History">History</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=1" title="Edit section: History"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Precursors of the idea of ecological succession go back to the beginning of the 19th century. As early as 1742 French naturalist <a href="/wiki/Georges-Louis_Leclerc,_Comte_de_Buffon" title="Georges-Louis Leclerc, Comte de Buffon">Buffon</a> noted that poplars precede oaks and beeches in the natural evolution of a forest. Buffon was later forced by the theological committee at the <a href="/wiki/University_of_Paris" title="University of Paris">University of Paris</a> to recant many of his ideas because they contradicted the biblical narrative of Creation.<sup id="cite_ref-ecology_6-0" class="reference"><a href="#cite_note-ecology-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p><p>Swiss geologist <a href="/wiki/Jean-Andr%C3%A9_Deluc" title="Jean-André Deluc">Jean-André Deluc</a> and the later French naturalist <a href="/wiki/Adolphe_Dureau_de_la_Malle" title="Adolphe Dureau de la Malle">Adolphe Dureau de la Malle</a> were the first to make use of the word <i>succession</i> concerning the vegetation development after forest clear-cutting.<sup id="cite_ref-Wetherington_7-0" class="reference"><a href="#cite_note-Wetherington-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Deluc-1813_8-0" class="reference"><a href="#cite_note-Deluc-1813-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> In 1859 <a href="/wiki/Henry_David_Thoreau" title="Henry David Thoreau">Henry David Thoreau</a> wrote an address called "The Succession of Forest Trees"<sup id="cite_ref-Thoreau-1887_9-0" class="reference"><a href="#cite_note-Thoreau-1887-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> in which he described succession in an oak-pine forest. "It has long been known to observers that squirrels bury nuts in the ground, but I am not aware that any one has thus accounted for the regular succession of forests."<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 Austrian botanist <a href="/wiki/Anton_Kerner_von_Marilaun" title="Anton Kerner von Marilaun">Anton Kerner</a> published a study about the succession of plants in the <a href="/wiki/River_Danube" class="mw-redirect" title="River Danube">Danube</a> river basin in 1863.<sup id="cite_ref-Bazzaz_11-0" class="reference"><a href="#cite_note-Bazzaz-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p><p><a href="/wiki/Ragnar_Hult" title="Ragnar Hult">Ragnar Hult</a>'s 1885 study on the stages of forest development in <a href="/wiki/Blekinge" title="Blekinge">Blekinge</a> noted that <a href="/wiki/Grassland" title="Grassland">grassland</a> becomes <a href="/wiki/Heath" title="Heath">heath</a> before the heath develops into forest. <a href="/wiki/Birch" title="Birch">Birch</a> dominated the early stages of forest development, then <a href="/wiki/Pine" title="Pine">pine</a> (on dry soil) and <a href="/wiki/Spruce" title="Spruce">spruce</a> (on wet soil). If the birch is replaced by <a href="/wiki/Oak" title="Oak">oak</a> it eventually develops to <a href="/wiki/Beech_tree" class="mw-redirect" title="Beech tree">beechwood</a>. Swamps proceed from <a href="/wiki/Moss" title="Moss">moss</a> to <a href="/wiki/Sedges" class="mw-redirect" title="Sedges">sedges</a> to moor vegetation followed by birch and finally spruce.<sup id="cite_ref-ecology_6-1" class="reference"><a href="#cite_note-ecology-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="H._C._Cowles">H. C. Cowles</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=2" title="Edit section: H. C. Cowles"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Indiana_dunes.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/f/f1/Indiana_dunes.jpg" decoding="async" width="300" height="429" class="mw-file-element" data-file-width="300" data-file-height="429" /></a><figcaption>The Indiana Dunes on Lake Michigan, which stimulated Cowles' development of his theories of ecological succession</figcaption></figure><p> Between 1899 and 1910, <a href="/wiki/Henry_Chandler_Cowles" title="Henry Chandler Cowles">Henry Chandler Cowles</a>, at the <a href="/wiki/University_of_Chicago" title="University of Chicago">University of Chicago</a>, developed a more formal concept of succession. Inspired by studies of <a href="/wiki/Denmark" title="Denmark">Danish</a> dunes by <a href="/wiki/Eugen_Warming" class="mw-redirect" title="Eugen Warming">Eugen Warming</a>, Cowles studied <a href="/wiki/Vegetation" title="Vegetation">vegetation</a> development on sand dunes on the shores of <a href="/wiki/Lake_Michigan" title="Lake Michigan">Lake Michigan</a> (the <a href="/wiki/Indiana_Dunes_National_Lakeshore" class="mw-redirect" title="Indiana Dunes National Lakeshore">Indiana Dunes</a>). He recognized that vegetation on dunes of different ages might be interpreted as different stages of a general trend of vegetation development on dunes (an approach to the study of vegetation change later termed space-for-time substitution, or <a href="/wiki/Chronosequence" title="Chronosequence">chronosequence</a> studies). He first published this work as a paper in the <i>Botanical Gazette</i> in 1899 ("The ecological relations of the vegetation of the sand dunes of Lake Michigan").<sup id="cite_ref-Cowles_12-0" class="reference"><a href="#cite_note-Cowles-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> In this classic publication and subsequent papers, he formulated the idea of primary succession and the notion of a <a href="/wiki/Sere_(ecology)" class="mw-redirect" title="Sere (ecology)">sere</a>—a repeatable sequence of community changes specific to particular environmental circumstances.<sup id="cite_ref-southshorejournal.org_4-1" class="reference"><a href="#cite_note-southshorejournal.org-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p><div class="mw-heading mw-heading3"><h3 id="Gleason_and_Clements">Gleason and Clements</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=3" title="Edit section: Gleason and Clements"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>From about 1900 to 1960, however, understanding of succession was dominated by the theories of <a href="/wiki/Frederic_Clements" title="Frederic Clements">Frederic Clements</a>, a contemporary of Cowles, who held that <a href="/wiki/Seral_community" title="Seral community">seres</a> were highly predictable and deterministic and converged on a climatically determined stable <a href="/wiki/Climax_(biology)" class="mw-redirect" title="Climax (biology)">climax community</a> regardless of starting conditions. Clements explicitly analogized the successional development of ecological communities with <a href="/wiki/Ontogenetic_niche_shift" title="Ontogenetic niche shift">ontogenetic development</a> of individual organisms, and his model is often referred to as the pseudo-organismic theory of community ecology. Clements and his followers developed a complex taxonomy of communities and successional pathways. </p><p><a href="/wiki/Henry_Gleason" class="mw-redirect" title="Henry Gleason">Henry Gleason</a> offered a contrasting framework as early as the 1920s. The Gleasonian model was more complex and much less deterministic than the Clementsian. It differs most fundamentally from the Clementsian view in suggesting a much greater role of <a href="/wiki/Stochastic_process" title="Stochastic process">chance factors</a> and in denying the existence of coherent, sharply bounded community types. Gleason argued that species distributions responded individualistically to environmental factors, and communities were best regarded as artifacts of the juxtaposition of species distributions. Gleason's ideas, first published in 1926, were largely ignored until the late 1950s. </p><p>Two quotes illustrate the contrasting views of Clements and Gleason. Clements wrote in 1916: </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>The developmental study of vegetation necessarily rests upon the assumption that the unit or climax formation is an organic entity. As an organism the formation arises, grows, matures, and dies. Furthermore, each climax formation is able to reproduce itself, repeating with essential fidelity the stages of its development.</p><div class="templatequotecite">— <cite>Frederic Clements<sup id="cite_ref-Clements_14-0" class="reference"><a href="#cite_note-Clements-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup></cite></div></blockquote> <p>while Gleason, in his 1926 paper, said: </p> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1244412712"><blockquote class="templatequote"><p>An association is not an organism, scarcely even a vegetational unit, but merely a coincidence.</p><div class="templatequotecite">— <cite>Henry Gleason<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup></cite></div></blockquote> <p>Gleason's ideas were, in fact, more consistent with Cowles' original thinking about succession. About Clements' distinction between <a href="/wiki/Primary_succession" title="Primary succession">primary succession</a> and <a href="/wiki/Secondary_succession" title="Secondary succession">secondary succession</a>, <a href="/wiki/Henry_Chandler_Cowles" title="Henry Chandler Cowles">Cowles</a> wrote (1911): </p> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1244412712"><blockquote class="templatequote"><p>This classification seems not to be of fundamental value, since it separates such closely related phenomena as those of erosion and deposition, and it places together such unlike things as human agencies and the subsidence of land.</p><div class="templatequotecite">— <cite>Henry Cowles<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup></cite></div></blockquote> <div class="mw-heading mw-heading3"><h3 id="Eugene_Odum">Eugene Odum</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=4" title="Edit section: Eugene Odum"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In 1969, Eugene Odum published <i>The Strategy of Ecosystem Development</i>, a paper that was highly influential to conservation and environmental restoration. Odum argued that ecological succession was an orderly progression toward a climax state where “maximum biomass and symbiotic function between organisms are maintained per unit energy flow."<sup id="cite_ref-Christensen_17-0" class="reference"><a href="#cite_note-Christensen-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> Odum highlighted how succession was not merely a change in the species composition of an ecosystem, but also created change in more complex attributes of the ecosystem, such as structure and <a href="/wiki/Nutrient_cycling" class="mw-redirect" title="Nutrient cycling">nutrient cycling</a>.<sup id="cite_ref-Anyomi_18-0" class="reference"><a href="#cite_note-Anyomi-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Modern_era">Modern era</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=5" title="Edit section: Modern era"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A more rigorous, data-driven testing of successional models and community theory generally began with the work of <a href="/wiki/Robert_Whittaker_(ecologist)" title="Robert Whittaker (ecologist)">Robert Whittaker</a> and <a href="/wiki/John_Thomas_Curtis" title="John Thomas Curtis">John Curtis</a> in the 1950s and 1960s. Succession theory has since become less monolithic and more complex. <a href="/wiki/Connell%E2%80%93Slatyer_model_of_ecological_succession" title="Connell–Slatyer model of ecological succession">J. Connell and R. Slatyer</a> attempted a codification of successional processes by mechanism. Among British and North American ecologists, the notion of a stable <a href="/wiki/Climax_vegetation" class="mw-redirect" title="Climax vegetation">climax vegetation</a> has been largely abandoned, and successional processes have come to be seen as much less deterministic, with important roles for historical <a href="/wiki/Contingency_(philosophy)" title="Contingency (philosophy)">contingency</a> and for alternate pathways in the actual development of communities. Debates continue as to the general predictability of successional dynamics and the relative importance of equilibrial vs. non-equilibrial processes. Former Harvard professor <a href="/wiki/Fakhri_A._Bazzaz" title="Fakhri A. Bazzaz">Fakhri A. Bazzaz</a> introduced the notion of <i>scale</i> into the discussion, as he considered that at local or small area scale the processes are stochastic and patchy, but taking bigger regional areas into consideration, certain tendencies can not be denied.<sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> </p><p>More recent definitions of succession highlight change as the central characteristic.<sup id="cite_ref-Christensen_17-1" class="reference"><a href="#cite_note-Christensen-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> New research techniques are greatly enhancing contemporary scientists' ability to study succession, which is now seen as neither entirely random nor entirely predictable.<sup id="cite_ref-Anyomi_18-1" class="reference"><a href="#cite_note-Anyomi-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Factors">Factors</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=6" title="Edit section: Factors"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Both consistent patterns and variability are observed in ecological succession. Theories of ecological succession identify different factors that help explain why plant communities change the way they do.<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Diversity_of_possible_trajectories">Diversity of possible trajectories</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=7" title="Edit section: Diversity of possible trajectories"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Ecological succession was formerly seen as an orderly progression through distinct stages, where several plant communities would replace each other in a fixed order and eventually reach a stable end point known as the <a href="/wiki/Climax_community" title="Climax community">climax</a>. The climax community was sometimes referred to as the 'potential vegetation' of a site, and thought to be primarily determined by the local climate. This idea has been largely abandoned by modern ecologists in favor of nonequilibrium ideas of ecosystems dynamics. Most natural ecosystems experience disturbance at a rate that makes a "climax" community unattainable. <a href="/wiki/Climate_change_(general_concept)" class="mw-redirect" title="Climate change (general concept)">Climate change</a> often occurs at a rate and frequency sufficient to prevent arrival at a climax state. </p><p>The trajectory of successional change can be influenced by initial site conditions, by the type of disturbance that triggers succession, by the interactions of the species present, and by more <a href="/wiki/Stochastic" title="Stochastic">random</a> factors such as availability of colonists or seeds or <a href="/wiki/Weather" title="Weather">weather</a> conditions at the time of disturbance. Some aspects of succession are broadly predictable; others may proceed more unpredictably than in the classical view of ecological succession. Coupled with the <a href="/wiki/Stochastic" title="Stochastic">stochastic</a> nature of disturbance events and other long-term (e.g., climatic) changes, such dynamics make it doubtful whether the 'climax' concept ever applies or is particularly useful in considering actual vegetation.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Stochastic_events">Stochastic events</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=8" title="Edit section: Stochastic events"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Succession is influenced partially by random chance, but it is debated how much random chance directs the trajectory of succession, as opposed to more deterministic factors. The timing of a disturbance such as a weather event may be random and unpredictable. Dispersal of propagules to a new site may also be random. However, community assembly is also determined by processes that select species non-randomly from the local species pool.<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Dispersal_limitation_vs._environmental_filtering">Dispersal limitation vs. environmental filtering</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=9" title="Edit section: Dispersal limitation vs. environmental filtering"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Succession is impacted both by the ability of seeds to disperse to new sites, and the suitability of site conditions for those seeds to grow and survive.<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> Dispersal limitation means that even though favorable sites for a plant to live might exist, the plant's seeds may be unable to reach those sites. Environmental filtering, also called establishment limitation, implies that although seeds may be distributed to a site, those seeds may be unable to survive due to various characteristics of the site. The predicted impact of these two factors varies under different models of ecological succession. </p> <div class="mw-heading mw-heading3"><h3 id="Feedback_loops">Feedback loops</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=10" title="Edit section: Feedback loops"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Ecological succession is driven by feedbacks between plants and their environment.<sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> As plants grow following a disturbance, they change their environment, for example by creating shade, attracting seed dispersers, contributing organic matter to the soil, changing the availability of soil nutrients, creating microhabitats, and buffering temperature and moisture fluctuations. This creates opportunities for different plants to grow, which causes directional change in the ecosystem. The development of some ecosystem attributes, such as <a href="/wiki/Pedogenesis" class="mw-redirect" title="Pedogenesis">soil properties</a> and <a href="/wiki/Nutrient_cycles" class="mw-redirect" title="Nutrient cycles">nutrient cycles</a>, are both influenced by community properties, and, in turn, influence further successional development. This feed-back process may occur over centuries or millennia. Plants may <a href="/wiki/Ecological_facilitation" title="Ecological facilitation">facilitate</a> the establishment of other plants by creating suitable conditions for them to grow, for example by providing shade or allowing for soil formation. Plants may also competitively exclude or otherwise prevent the growth of other plants. </p> <div class="mw-heading mw-heading3"><h3 id="Patterns">Patterns</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=11" title="Edit section: Patterns"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Though the idea of a fixed, predictable process of succession with a single well-defined climax is an overly simplified model, several predictions made by the classical model are accurate. Species diversity, overall plant biomass, plant lifespans, the importance of <a href="/wiki/Decomposer" title="Decomposer">decomposer</a> organisms, and overall stability all increase as a community approaches a climax state, while the rate at which soil nutrients are consumed, rate of biogeochemical cycling, and rate of net primary productivity all decrease as a community approaches a climax state.<sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> </p><p>Communities in early succession will be dominated by fast-growing, well-<a href="/wiki/Biological_dispersal" title="Biological dispersal">dispersed</a> species (<a href="/wiki/Opportunism#Biological" title="Opportunism">opportunist</a>, <a href="/wiki/Glossary_of_evolutionary_biology#fugitive_species" class="mw-redirect" title="Glossary of evolutionary biology">fugitive</a>, or <a href="/wiki/R-selected" class="mw-redirect" title="R-selected">r-selected</a> life-histories). These are also called <a href="/wiki/Pioneer_species" title="Pioneer species">pioneer species</a>. As succession proceeds, these species will tend to be replaced by more competitive (<a href="/wiki/K-selected" class="mw-redirect" title="K-selected">k-selected</a>) species. </p><p>Some of these trends do not apply in all cases. For example, <a href="/wiki/Species_diversity" title="Species diversity">species diversity</a> almost necessarily increases during early succession as new species arrive, but may decline in later succession as competition eliminates opportunistic species and leads to <a href="/wiki/Competitive_exclusion_principle" title="Competitive exclusion principle">dominance by locally superior competitors</a>. <a href="/wiki/Primary_production" title="Primary production">Net Primary Productivity</a>, <a href="/wiki/Biomass_(ecology)" title="Biomass (ecology)">biomass</a>, and <a href="/wiki/Trophic_level" title="Trophic level">trophic</a> properties all show variable patterns over succession, depending on the particular system and site. </p> <div class="mw-heading mw-heading3"><h3 id="Disruptions">Disruptions</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=12" title="Edit section: Disruptions"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Two important perturbation factors today are <a href="/wiki/Human_impact_on_the_environment" title="Human impact on the environment">human actions</a> and <a href="/wiki/Global_warming" class="mw-redirect" title="Global warming">climatic change</a>.<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> Additions to available species pools through range expansions and <a href="/wiki/Introduced_species" title="Introduced species">introductions</a> can also continually reshape communities.<sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Types_of_succession">Types of succession</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=13" title="Edit section: Types of succession"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Primary_succession">Primary succession</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=14" title="Edit section: Primary succession"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Primary_succession" title="Primary succession">Primary succession</a></div> <p>The successional dynamics beginning with colonization of an area with little to no biotic factors is referred to as primary succession.<sup id="cite_ref-Fisher-2018_1-2" class="reference"><a href="#cite_note-Fisher-2018-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> This includes newly exposed rock or sand surfaces, lava flows, and newly exposed glacial tills.<sup id="cite_ref-Fisher-2018_1-3" class="reference"><a href="#cite_note-Fisher-2018-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> The stages of primary succession include pioneer species such as microorganisms,<sup id="cite_ref-Ortiz-Álvarez-2018_29-0" class="reference"><a href="#cite_note-Ortiz-Álvarez-2018-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> plants (lichens and mosses), grasses, smaller shrubs, and trees. Small animals move into the area when enough suitable habitat has been established. When it is a fully functioning ecosystem, it has reached the climax community stage.<sup id="cite_ref-biologydictionary_30-0" class="reference"><a href="#cite_note-biologydictionary-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Secondary_succession">Secondary succession</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=15" title="Edit section: Secondary succession"><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/Secondary_succession" title="Secondary succession">Secondary succession</a></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Secondary_Succession.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/47/Secondary_Succession.png/450px-Secondary_Succession.png" decoding="async" width="450" height="175" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/47/Secondary_Succession.png/675px-Secondary_Succession.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/47/Secondary_Succession.png/900px-Secondary_Succession.png 2x" data-file-width="1459" data-file-height="568" /></a><figcaption>An example of secondary succession by stages: <div><ol><li>A stable deciduous forest community</li><li>A disturbance, such as a wild fire, destroys the forest</li><li>The fire burns the forest to the ground</li><li>The fire leaves behind empty, but not destroyed, soil</li><li>Grasses and other herbaceous plants grow back first</li><li>Small bushes and trees begin to colonize the area</li><li>Fast-growing evergreen trees develop to their fullest, while shade-tolerant trees develop in the understory</li><li>The short-lived and shade-intolerant evergreen trees die as the larger deciduous trees overtop them. The ecosystem is now back to a similar state to where it began.</li></ol></div></figcaption></figure> <p>Secondary succession follows severe disturbance or removal of a preexisting community that has remnants of the previous ecosystem.<sup id="cite_ref-Fisher-2018_1-4" class="reference"><a href="#cite_note-Fisher-2018-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Secondary succession is strongly influenced by pre-disturbance conditions such as soil development, <a href="/wiki/Soil_seed_bank" title="Soil seed bank">seed banks</a>, remaining organic matter, and residual living organisms.<sup id="cite_ref-Fisher-2018_1-5" class="reference"><a href="#cite_note-Fisher-2018-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Because of residual fertility and preexisting organisms, community change in early stages of secondary succession can be relatively rapid.<sup id="cite_ref-Fisher-2018_1-6" class="reference"><a href="#cite_note-Fisher-2018-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>Secondary succession is much more commonly observed and studied than primary succession. Particularly common types of secondary succession include responses to natural disturbances such as fire, flood, and severe winds, and to human-caused disturbances such as logging and agriculture. In secondary succession, the soils and organisms need to be left unharmed so there is a way for the new material to rebuild.<sup id="cite_ref-Thoreau-1887_9-1" class="reference"><a href="#cite_note-Thoreau-1887-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p><p>As an example, in a fragmented old field habitat created in eastern Kansas, woody plants "colonized more rapidly (per unit area) on large and nearby <a href="/wiki/Landscape_ecology#Patch_and_mosaic" title="Landscape ecology">patches</a>".<sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Secondary_succesion_cm01.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a2/Secondary_succesion_cm01.jpg/450px-Secondary_succesion_cm01.jpg" decoding="async" width="450" height="300" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a2/Secondary_succesion_cm01.jpg/675px-Secondary_succesion_cm01.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/a/a2/Secondary_succesion_cm01.jpg 2x" data-file-width="800" data-file-height="533" /></a><figcaption><i>Secondary succession</i>: trees are colonizing uncultivated fields and meadows.</figcaption></figure> <p>Secondary succession can quickly change a landscape. In the 1900s, <a href="/wiki/Acadia_National_Park" title="Acadia National Park">Acadia National Park</a> had a wildfire that destroyed much of the landscape. Originally evergreen trees grew in the landscape. After the fire, the area took at least a year to grow shrubs. Eventually, deciduous trees started to grow instead of evergreens.<sup id="cite_ref-biologydictionary_30-1" class="reference"><a href="#cite_note-biologydictionary-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> </p><p>Secondary succession has been occurring in <a href="/wiki/Shenandoah_National_Park" title="Shenandoah National Park">Shenandoah National Park</a> following the 1995 flood of the <a href="/wiki/Moorman%27s_River" class="mw-redirect" title="Moorman's River">Moorman's</a> and Rapidan rivers, which destroyed plant and animal life.<sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Seasonal_and_cyclic_dynamics">Seasonal and cyclic dynamics</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=16" title="Edit section: Seasonal and cyclic dynamics"><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/Cyclic_succession" title="Cyclic succession">Cyclic succession</a></div> <p>Unlike secondary succession, these types of vegetation change are not dependent on <a href="/wiki/Disturbance_(ecology)" title="Disturbance (ecology)">disturbance</a> but are periodic changes arising from fluctuating species interactions or recurring events. These models modify the <a href="/wiki/Climax_community" title="Climax community">climax concept</a> towards one of dynamic states. </p> <div class="mw-heading mw-heading3"><h3 id="Causes_of_plant_succession">Causes of plant succession</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=17" title="Edit section: Causes of plant succession"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Autogenic_succession" title="Autogenic succession">Autogenic succession</a> can be brought by changes in the soil caused by the organisms there. These changes include accumulation of organic matter in litter or humic layer, alteration of soil nutrients, or change in the pH of soil due to the plants growing there. The structure of the plants themselves can also alter the community.<sup id="cite_ref-33" class="reference"><a href="#cite_note-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup> For example, when larger species like trees mature, they produce shade on to the developing <a href="/wiki/Forest_floor" title="Forest floor">forest floor</a> that tends to exclude light-requiring species. Shade-tolerant species will invade the area. </p><p><a href="/wiki/Allogenic_succession" title="Allogenic succession">Allogenic succession</a> is caused by external environmental influences and not by the vegetation. For example, soil changes due to erosion, leaching or the deposition of silt and clays can alter the nutrient content and water relationships in the ecosystems. Animals also play an important role in allogenic changes as they are pollinators, seed dispersers and herbivores. They can also increase nutrient content of the soil in certain areas, or shift soil about (as termites, ants, and moles do) creating patches in the habitat. This may create regeneration sites that favor certain species. </p><p>Climatic factors may be very important, but on a much longer time-scale than any other. Changes in temperature and rainfall patterns will promote changes in communities. As the climate warmed at the end of each ice age, great successional changes took place. The tundra vegetation and bare glacial till deposits underwent succession to mixed deciduous forest. The <a href="/wiki/Greenhouse_effect" title="Greenhouse effect">greenhouse effect</a> resulting in increase in temperature is likely to bring profound Allogenic changes in the next century. Geological and climatic catastrophes such as volcanic eruptions, earthquakes, avalanches, meteors, floods, fires, and high wind also bring allogenic changes. </p> <div class="mw-heading mw-heading3"><h3 id="Mechanisms">Mechanisms</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=18" title="Edit section: Mechanisms"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In 1916, <a href="/wiki/Frederic_Clements" title="Frederic Clements">Frederic Clements</a> published a descriptive theory of succession and advanced it as a general ecological concept.<sup id="cite_ref-Clements_14-1" class="reference"><a href="#cite_note-Clements-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> His theory of succession had a powerful influence on ecological thought. Clements' concept is usually termed classical <a href="/wiki/Ecological_theory" class="mw-redirect" title="Ecological theory">ecological theory</a>.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (January 2021)">citation needed</span></a></i>]</sup> According to Clements, succession is a process involving several phases:<sup id="cite_ref-Clements_14-2" class="reference"><a href="#cite_note-Clements-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup><sup class="noprint Inline-Template" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citing_sources" title="Wikipedia:Citing sources"><span title="This citation requires a reference to the specific page or range of pages in which the material appears. (June 2016)">page needed</span></a></i>]</sup> </p> <ol><li>Nudation: Succession begins with the development of a bare site, called Nudation (disturbance).<sup id="cite_ref-Clements_14-3" class="reference"><a href="#cite_note-Clements-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup></li> <li>Migration: refers to arrival of <a href="/wiki/Propagule" title="Propagule">propagules</a>.<sup id="cite_ref-Clements_14-4" class="reference"><a href="#cite_note-Clements-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup></li> <li>Ecesis: involves establishment and initial growth of vegetation.<sup id="cite_ref-Clements_14-5" class="reference"><a href="#cite_note-Clements-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup></li> <li>Competition: as vegetation becomes well established, grows, and spreads, various species begin to compete for space, light and nutrients.<sup id="cite_ref-Clements_14-6" class="reference"><a href="#cite_note-Clements-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup></li> <li>Reaction: during this phase autogenic changes such as the buildup of humus affect the habitat, and one plant community replaces another.<sup id="cite_ref-Clements_14-7" class="reference"><a href="#cite_note-Clements-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup></li> <li>Stabilization: a supposedly stable climax community forms.<sup id="cite_ref-Clements_14-8" class="reference"><a href="#cite_note-Clements-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup></li></ol> <div class="mw-heading mw-heading2"><h2 id="Seral_communities">Seral communities</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=19" title="Edit section: Seral communities"><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/Seral_community" title="Seral community">Seral community</a></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Aquatic_Succession_or_Hydrosere.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/2b/Aquatic_Succession_or_Hydrosere.svg/255px-Aquatic_Succession_or_Hydrosere.svg.png" decoding="async" width="255" height="434" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/2b/Aquatic_Succession_or_Hydrosere.svg/383px-Aquatic_Succession_or_Hydrosere.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/2b/Aquatic_Succession_or_Hydrosere.svg/511px-Aquatic_Succession_or_Hydrosere.svg.png 2x" data-file-width="512" data-file-height="869" /></a><figcaption><a href="/wiki/Aquatic_succession" title="Aquatic succession">Aquatic succession</a> or <a href="/wiki/Hydrosere" title="Hydrosere">hydrosere</a>: A) Emergent plant life, or pond <a href="/wiki/Pioneer_species" title="Pioneer species">pioneers</a>, surround an open water lake. Some <a href="/wiki/Terrestrial_plant" title="Terrestrial plant">terrestrial plant</a> life surrounds it as well. B) Emergent plant life begins to move inward and submerge, filling up the lake. Sediment accumulates as the plants grow and die. Terrestrial plant life increases. C) Sediment fills the lake basin, leaving a swampy center at the surface. Terrestrial plants take over and begin to slowly dry out the damp surface.</figcaption></figure> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Schleienloecher2-1-_C.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8f/Schleienloecher2-1-_C.jpg/450px-Schleienloecher2-1-_C.jpg" decoding="async" width="450" height="245" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8f/Schleienloecher2-1-_C.jpg/675px-Schleienloecher2-1-_C.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8f/Schleienloecher2-1-_C.jpg/900px-Schleienloecher2-1-_C.jpg 2x" data-file-width="3008" data-file-height="1635" /></a><figcaption>A <a href="/wiki/Hydrosere" title="Hydrosere">hydrosere</a> community</figcaption></figure> <p>A seral community is an intermediate stage found in an ecosystem advancing towards its <a href="/wiki/Climax_community" title="Climax community">climax community</a>. In many cases more than one seral stage evolves until climax conditions are attained.<sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> A <i><a href="/wiki/Prisere" class="mw-redirect" title="Prisere">prisere</a></i> is a collection of seres making up the development of an area from non-vegetated surfaces to a climax community. Depending on the substratum and climate, different seres are found. </p> <div class="mw-heading mw-heading2"><h2 id="Changes_in_animal_life">Changes in animal life</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=20" title="Edit section: Changes in animal life"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Succession theory was developed primarily by botanists. The study of succession applied to whole <a href="/wiki/Ecosystems" class="mw-redirect" title="Ecosystems">ecosystems</a> initiated in the writings of <a href="/wiki/Ramon_Margalef" title="Ramon Margalef">Ramon Margalef</a>, while <a href="/wiki/Eugene_Odum" title="Eugene Odum">Eugene Odum</a>'s publication of <i>The Strategy of Ecosystem Development</i> is considered its formal starting point.<sup id="cite_ref-35" class="reference"><a href="#cite_note-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> </p><p>Animal life also exhibits changes with changing communities. In the lichen stage, fauna is sparse. It comprises a few mites, ants, and spiders living in cracks and crevices. The fauna undergoes a qualitative increase during the herb grass stage. The animals found during this stage include nematodes, insect larvae, ants, spiders, mites, etc. The animal population increases and diversifies with the development of the forest climax community. The fauna consists of invertebrates like slugs, snails, worms, millipedes, centipedes, ants, bugs; and vertebrates such as squirrels, foxes, mice, moles, snakes, various birds, salamanders and frogs. </p><p>A review of succession research by Hodkinson et al. (2002)<sup id="cite_ref-36" class="reference"><a href="#cite_note-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> documented what was likely first noted by Darwin during his voyage on the H.M.S. Beagle: </p> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1244412712"><blockquote class="templatequote"><p>The often repeated description of the stately palm and other nobel plants, then birds, and lastly man, taking possession of the coral islets as soon as formed in the Pacific, is probably not quite correct; I fear it destroys the poetry of this story, that feather and dirt-feeding and parasitic insects and spiders should be the first inhabitants of newly-formed oceanic land.</p><div class="templatequotecite">— <cite>C. Darwin <sup id="cite_ref-37" class="reference"><a href="#cite_note-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup></cite></div></blockquote> <p>These naturalists note that prior to the establishment of autotrophs, there is a foodweb formed by heterotrophs built on allochthonous inputs of dead organic matter (necromass). Work on volcanic systems such as Kasatochi Volcano in the Aleutians by Sikes and Slowik (2010)<sup id="cite_ref-38" class="reference"><a href="#cite_note-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> supports this idea. </p><p><br /> </p> <div class="mw-heading mw-heading2"><h2 id="Microsuccession">Microsuccession</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=21" title="Edit section: Microsuccession"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Succession of <a href="/wiki/Micro-organisms" class="mw-redirect" title="Micro-organisms">micro-organisms</a> including <a href="/wiki/Fungi" class="mw-redirect" title="Fungi">fungi</a> and <a href="/wiki/Bacteria" title="Bacteria">bacteria</a> occurring within a microhabitat is known as microsuccession or serule. In artificial bacterial meta-communities of motile strains <i>on-chip</i> it has been shown that ecological succession is based on a trade-off between colonization and competition abilities. To exploit locations or explore the landscape? <i>Escherichia coli</i> is a fugitive species, whereas <i>Pseudomonas aeruginosa</i> is a slower colonizer but superior competitor.<sup id="cite_ref-Wetherington_7-1" class="reference"><a href="#cite_note-Wetherington-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> Like in plants, microbial succession can occur in newly available habitats (<a href="/wiki/Primary_succession" title="Primary succession">primary succession</a>) such as surfaces of plant leaves, recently exposed rock surfaces (i.e., glacial till) or animal infant guts,<sup id="cite_ref-Ortiz-Álvarez-2018_29-1" class="reference"><a href="#cite_note-Ortiz-Álvarez-2018-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> and also on disturbed communities (<a href="/wiki/Secondary_succession" title="Secondary succession">secondary succession</a>) like those growing in recently dead trees, decaying fruits,<sup id="cite_ref-39" class="reference"><a href="#cite_note-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup> or animal droppings. Microbial communities may also change due to products secreted by the bacteria present. Changes of pH in a habitat could provide ideal conditions for a new species to inhabit the area. In some cases the new species may outcompete the present ones for nutrients leading to the primary species demise. Changes can also occur by microbial succession with variations in water availability and temperature. Theories of <a href="/wiki/Macroecology" title="Macroecology">macroecology</a> have only recently been applied to <a href="/wiki/Microbiology" title="Microbiology">microbiology</a> and so much remains to be understood about this growing field. A recent study of microbial succession evaluated the balances between <a href="/wiki/Stochastic" title="Stochastic">stochastic</a> and deterministic processes in the bacterial colonization of a salt marsh <a href="/wiki/Chronosequence" title="Chronosequence">chronosequence</a>. The results of this study show that, much like in macro succession, early colonization (<a href="/wiki/Primary_succession" title="Primary succession">primary succession</a>) is mostly influenced by stochasticity while secondary succession of these bacterial communities was more strongly influenced by deterministic factors.<sup id="cite_ref-40" class="reference"><a href="#cite_note-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> </p> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Succesion_in_Bacteria.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/03/Succesion_in_Bacteria.jpg/500px-Succesion_in_Bacteria.jpg" decoding="async" width="500" height="386" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/03/Succesion_in_Bacteria.jpg/750px-Succesion_in_Bacteria.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/03/Succesion_in_Bacteria.jpg/1000px-Succesion_in_Bacteria.jpg 2x" data-file-width="1056" data-file-height="816" /></a><figcaption>Ecological micro-succession in a bacterial meta-community <i>on-chip</i>. (<b>A</b>) sketch of a micron-scale structured bacterial environment based on microfluidics technology; (<b>B</b>) Fluorescent microscopy image of <i>Escherichia coli</i> (magenta) and <i>Pseudomonas aeruginosa</i> (green) inhabiting a device of the type depicted in A and which has been wettened with growth media and inoculated with both species; (<b>C</b>) a sequence of five snapshots of the bacterial community distributed over five patches (of an array with 85) depicting the spatial dynamics of competition between <i>E. coli</i> (magenta) and <i>P. aeruginosa</i> (green); (<b>D</b>) Representation of the succession pattern exhibited by the two bacterial species when competing for space and resources in a patchy environment.<sup id="cite_ref-Wetherington_7-2" class="reference"><a href="#cite_note-Wetherington-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup></figcaption></figure> <div class="mw-heading mw-heading2"><h2 id="Climax_concept">Climax concept</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=22" title="Edit section: Climax concept"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>According to classical <a href="/wiki/Ecological_theory" class="mw-redirect" title="Ecological theory">ecological theory</a>, succession stops when the sere has arrived at an equilibrium or steady state with the physical and biotic environment. Barring major disturbances, it will persist indefinitely.<sup id="cite_ref-Fisher-2018_1-7" class="reference"><a href="#cite_note-Fisher-2018-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> This end point of succession is called climax. </p> <div class="mw-heading mw-heading3"><h3 id="Climax_community">Climax community</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=23" title="Edit section: Climax community"><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/Climax_community" title="Climax community">Climax community</a></div> <p>The final or stable community in a sere is the <i>climax community</i> or <i>climatic vegetation</i>. It is self-perpetuating and in equilibrium with the physical habitat.<sup id="cite_ref-Fisher-2018_1-8" class="reference"><a href="#cite_note-Fisher-2018-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> There is no net annual accumulation of organic matter in a climax community. The annual production and use of energy is balanced in such a community. </p> <div class="mw-heading mw-heading3"><h3 id="Characteristics">Characteristics</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=24" title="Edit section: Characteristics"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li>The vegetation is tolerant of environmental conditions.</li> <li>It has a wide diversity of species, a well-drained spatial structure, and complex food chains.</li> <li>The climax ecosystem is balanced. There is equilibrium between <a href="/wiki/Gross_primary_production" class="mw-redirect" title="Gross primary production">gross primary production</a> and total respiration, between energy used from sunlight and energy released by decomposition, between uptake of nutrients from the soil and the return of nutrient by litter fall to the soil.</li> <li>Individuals in the climax stage are replaced by others of the same kind. Thus the <a href="/wiki/Species_composition" class="mw-redirect" title="Species composition">species composition</a> maintains equilibrium.</li> <li>It is an index of the climate of the area. The life or growth forms indicate the climatic type.</li></ul> <div class="mw-heading mw-heading3"><h3 id="Types_of_climax">Types of climax</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=25" title="Edit section: Types of climax"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <dl><dt>Climatic Climax</dt> <dd>If there is only a single climax and the development of climax community is controlled by the climate of the region, it is termed as climatic climax. For example, development of Maple-beech climax community over moist soil. Climatic climax is theoretical and develops where physical conditions of the substrate are not so extreme as to modify the effects of the prevailing regional climate.</dd> <dt>Edaphic Climax</dt> <dd>When there are more than one climax communities in the region, modified by local conditions of the substrate such as soil moisture, soil nutrients, topography, slope exposure, fire, and animal activity, it is called <i>edaphic climax</i>. Succession ends in an edaphic climax where topography, soil, water, fire, or other disturbances are such that a climatic climax cannot develop.</dd> <dt>Catastrophic Climax</dt> <dd>Climax vegetation vulnerable to a catastrophic event such as a wildfire. For example, in <a href="/wiki/California" title="California">California</a>, <a href="/wiki/Chaparral" title="Chaparral">chaparral</a> vegetation is the final vegetation. The wildfire removes the mature vegetation and decomposers. A rapid development of herbaceous vegetation follows until the shrub dominance is re-established. This is known as catastrophic climax.</dd> <dt>Disclimax</dt> <dd>When a stable community, which is not the climatic or edaphic climax for the given site, is maintained by man or his domestic animals, it is designated as Disclimax (disturbance climax) or anthropogenic subclimax (man-generated). For example, <a href="/wiki/Overgrazing" title="Overgrazing">overgrazing</a> by stock may produce a desert community of bushes and cacti where the local climate actually would allow grassland to maintain itself.</dd> <dt>Subclimax</dt> <dd>The prolonged stage in succession just preceding the climatic climax is <i>subclimax</i>.</dd> <dt>Preclimax and Postclimax</dt> <dd>In certain areas different climax communities develop under similar climatic conditions. If the community has life forms lower than those in the expected climatic climax, it is called <i>preclimax</i>; a community that has life forms higher than those in the expected climatic climax is <i>postclimax</i>. Preclimax strips develop in less moist and hotter areas, whereas Postclimax strands develop in more moist and cooler areas than that of surrounding climate.</dd></dl> <div class="mw-heading mw-heading3"><h3 id="Theories">Theories</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=26" title="Edit section: Theories"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>There are three schools of interpretations explaining the climax concept: </p> <ul><li>Monoclimax or Climatic Climax Theory was advanced by <a href="/wiki/Frederic_Clements" title="Frederic Clements">Clements</a> (1916) and recognizes only one climax whose characteristics are determined solely by climate (climatic climax). The processes of succession and modification of environment overcome the effects of differences in topography, parent material of the soil, and other factors. The whole area would be covered with uniform plant community. Communities other than the climax are related to it, and are recognized as subclimax, postclimax and disclimax.</li> <li>Polyclimax Theory was advanced by Tansley (1935). It proposes that the climax vegetation of a region consists of more than one vegetation climaxes controlled by soil moisture, soil nutrients, topography, slope exposure, fire, and animal activity.</li> <li>Climax Pattern Theory was proposed by Whittaker (1953). The climax pattern theory recognizes a variety of climaxes governed by responses of species populations to biotic and abiotic conditions. According to this theory the total environment of the ecosystem determines the composition, species structure, and balance of a climax community. The environment includes the species' responses to moisture, temperature, and nutrients, their biotic relationships, availability of flora and fauna to colonize the area, chance dispersal of seeds and animals, soils, climate, and disturbance such as fire and wind. The nature of climax vegetation will change as the environment changes. The climax community represents a pattern of populations that corresponds to and changes with the pattern of environment. The central and most widespread community is the climatic climax.</li></ul> <p>The theory of <a href="/wiki/Alternative_stable_states" class="mw-redirect" title="Alternative stable states">alternative stable states</a> suggests there is not one end point but many which transition between each other over ecological time. </p> <div class="mw-heading mw-heading2"><h2 id="Succession_by_habitat_type">Succession by habitat type</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=27" title="Edit section: Succession by habitat type"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Forest_succession">Forest succession</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=28" title="Edit section: Forest succession"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Forest_succession_depicted_over_time.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/41/Forest_succession_depicted_over_time.png/750px-Forest_succession_depicted_over_time.png" decoding="async" width="750" height="536" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/41/Forest_succession_depicted_over_time.png/1125px-Forest_succession_depicted_over_time.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/41/Forest_succession_depicted_over_time.png/1500px-Forest_succession_depicted_over_time.png 2x" data-file-width="4000" data-file-height="2859" /></a><figcaption></figcaption></figure> <p>Forests, being an ecological system, are subject to the species succession process.<sup id="cite_ref-41" class="reference"><a href="#cite_note-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> There are "opportunistic" or "pioneer" species that produce great quantities of seed that are disseminated by the wind, and therefore can colonize big empty extensions.<sup id="cite_ref-Budowski_42-0" class="reference"><a href="#cite_note-Budowski-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> They are capable of germinating and growing in direct sunlight. Once they have produced a <i><a href="/w/index.php?title=Closed_canopy&action=edit&redlink=1" class="new" title="Closed canopy (page does not exist)">closed canopy</a></i>, the lack of direct sun radiation at the soil makes it difficult for their own seedlings to develop. It is then the opportunity for <a href="/wiki/Shade-tolerant" class="mw-redirect" title="Shade-tolerant">shade-tolerant</a> species to become established under the protection of the pioneers. When the pioneers die, the shade-tolerant species replace them. These species are capable of growing beneath the canopy, and therefore, in the absence of disturbances, will stay. For this reason it is then said the <a href="/wiki/Stand_level_modelling" class="mw-redirect" title="Stand level modelling">stand</a> has reached its climax. When a disturbance occurs, the opportunity for the pioneers opens up again, provided they are present or within a reasonable range. </p><p>An example of pioneer species, in forests of northeastern North America are <i>Betula papyrifera</i> (<a href="/wiki/White_birch" class="mw-redirect" title="White birch">White birch</a>) and <i>Prunus serotina</i> (<a href="/wiki/Black_cherry" class="mw-redirect" title="Black cherry">Black cherry</a>), that are particularly well-adapted to exploit large gaps in forest canopies, but are intolerant of shade and are eventually replaced by other <a href="/wiki/Shade-tolerant" class="mw-redirect" title="Shade-tolerant">shade-tolerant</a> species in the absence of disturbances that create such gaps. In the tropics, well known pioneer forest species can be found among the genera <i>Cecropia</i>, <i>Ochroma</i> and <i>Trema</i>.<sup id="cite_ref-Budowski_42-1" class="reference"><a href="#cite_note-Budowski-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> </p><p>Things in nature are not black and white, and there are intermediate stages. It is therefore normal that between the two extremes of light and shade there is a gradient, and there are species that may act as pioneer or tolerant, depending on the circumstances. It is of paramount importance to know the tolerance of species in order to practice an effective <a href="/wiki/Silviculture" title="Silviculture">silviculture</a>. </p> <div class="mw-heading mw-heading3"><h3 id="Wetland_succession">Wetland succession</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=29" title="Edit section: Wetland succession"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Since many types of <a href="/wiki/Wetland" title="Wetland">wetland</a> environments exist, succession may follow a wide array of trajectories and patterns in wetlands. Under the classical model, the process of secondary succession holds that a wetland progresses over time from an initial state of open water with few plants, to a <a href="/wiki/Swamp" title="Swamp">forested</a> climax state where decayed organic matter has built up over time, forming <a href="/wiki/Peat" title="Peat">peat</a>. However, many wetlands are maintained by regular disturbance or natural processes at an equilibrium state that does not resemble the predicted forested "climax."<sup id="cite_ref-Moseley_43-0" class="reference"><a href="#cite_note-Moseley-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> The idea that ponds and wetlands gradually fill in to become dry land has been criticized and called into question due to lack of evidence.<sup id="cite_ref-middleton_5-1" class="reference"><a href="#cite_note-middleton-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>Wetland succession is a uniquely complex, non-linear process shaped by <a href="/wiki/Hydrology" title="Hydrology">hydrology</a>.<sup id="cite_ref-Zweig_44-0" class="reference"><a href="#cite_note-Zweig-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> Hydrological factors often work against linear processes that predict a succession to a "climax" state. The energy carried by moving water may create a continuous source of disturbance. For example, in coastal wetlands, the <a href="/wiki/Tides" class="mw-redirect" title="Tides">tides</a> moving in and out continuously acts upon the ecological community. <a href="/wiki/Fire_ecology" title="Fire ecology">Fire</a> may also maintain an equilibrium state in a wetland by burning off vegetation, thus interrupting the accumulation of peat.<sup id="cite_ref-Moseley_43-1" class="reference"><a href="#cite_note-Moseley-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> Water entering and leaving the wetland follows patterns that are broadly cyclical but erratic. For example, seasonal flooding and drying may occur with yearly changes in precipitation, causing seasonal changes in the wetland community that maintain it at a stable state.<sup id="cite_ref-middleton_5-2" class="reference"><a href="#cite_note-middleton-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> However, unusually heavy rain or unusually severe <a href="/wiki/Drought" title="Drought">drought</a> may cause the wetland to enter a positive feedback loop where it begins to change in a linear direction.<sup id="cite_ref-Zweig_44-1" class="reference"><a href="#cite_note-Zweig-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> Since wetlands are sensitive to changes in the natural processes that maintain them, human activities, <a href="/wiki/Invasive_species" title="Invasive species">invasive species</a>, and <a href="/wiki/Climate_change" title="Climate change">climate change</a> could initiate long-term changes in wetland ecosystems.<sup id="cite_ref-Moseley_43-2" class="reference"><a href="#cite_note-Moseley-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Grassland_succession">Grassland succession</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=30" title="Edit section: Grassland succession"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>For a long time, grasslands were thought to be early stages of succession, dominated by <a href="/wiki/Weed" title="Weed">weedy</a> species and with little conservation value. However, comparing grasslands that form after recovery from long-term disruptions like agricultural <a href="/wiki/Tillage" title="Tillage">tillage</a> with ancient or "old-growth" grasslands has shown that grasslands are not inherently early-successional communities. Rather, grasslands undergo a centuries-long process of succession, and a grassland that is tilled up for agriculture or otherwise destroyed is estimated to take a minimum of 100 years, and potentially on average 1,400 years, to recover to its previous level of biodiversity.<sup id="cite_ref-High_plant_diversity_and_slow_assem_45-0" class="reference"><a href="#cite_note-High_plant_diversity_and_slow_assem-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> However, planting a high diversity of late-successional grassland species in a disturbed environment can accelerate the recovery of the soil's ability to sequester carbon, resulting in twice as much carbon storage as a naturally recovering grassland over the same period of time.<sup id="cite_ref-46" class="reference"><a href="#cite_note-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> </p><p>Many grassland ecosystems are maintained by disturbance, such as fire and grazing by large animals, or else the process of succession will change them to forest or shrubland. In fact, it is debated whether fire should be considered disturbance at all for the North American <a href="/wiki/Prairie" title="Prairie">prairie</a> ecosystems, since it maintains, rather than disrupts, an equilibrium state.<sup id="cite_ref-47" class="reference"><a href="#cite_note-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> Many late-successional grassland species have adaptations that allow them to store nutrients underground and re-sprout rapidly after "aboveground" disturbances like fire or grazing. Disturbance events that severely disrupt or destroy the soil, such as tilling, eliminate these late-successional species, reverting the grassland to an early successional stage dominated by <a href="/wiki/Pioneer_species" title="Pioneer species">pioneers</a>, whereas fire and grazing benefit late-successional species.<sup id="cite_ref-High_plant_diversity_and_slow_assem_45-1" class="reference"><a href="#cite_note-High_plant_diversity_and_slow_assem-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> Both too much and too little disturbance can damage the biodiversity of disturbance-dependent ecosystems like grasslands.<sup id="cite_ref-Schnoor_48-0" class="reference"><a href="#cite_note-Schnoor-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> </p><p>In North American semi-arid grasslands, the introduction of livestock ranching and absence of fire was observed to cause a transition away from grasses to woody vegetation, particularly <a href="/wiki/Prosopis" title="Prosopis">mesquite</a>.<sup id="cite_ref-49" class="reference"><a href="#cite_note-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> However, the means by which ecological succession under frequent disturbance results in ecosystems of the sort seen in remnant prairies is poorly understood.<sup id="cite_ref-50" class="reference"><a href="#cite_note-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Schnoor_48-1" class="reference"><a href="#cite_note-Schnoor-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Ecological_succession&action=edit&section=31" title="Edit section: See also"><span>edit</span></a><span 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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=Ecological_succession&action=edit&section=32" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-Fisher-2018-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-Fisher-2018_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Fisher-2018_1-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Fisher-2018_1-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Fisher-2018_1-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-Fisher-2018_1-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-Fisher-2018_1-5"><sup><i><b>f</b></i></sup></a> <a href="#cite_ref-Fisher-2018_1-6"><sup><i><b>g</b></i></sup></a> <a href="#cite_ref-Fisher-2018_1-7"><sup><i><b>h</b></i></sup></a> <a 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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="CITEREFFisher2018" class="citation book cs1">Fisher MR (2018). <i>Environmental Biology</i>. 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Bruun, Hans Henrik; Olsson, Pal Axel (2015). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4395216">"Soil Disturbance as a Grassland Restoration Measure—Effects on Plant Species Composition and Plant Functional Traits"</a>. <i>PLOS ONE</i>. <b>10</b> (4): e0123698. <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/2015PLoSO..1023698S">2015PLoSO..1023698S</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/10.1371%2Fjournal.pone.0123698">10.1371/journal.pone.0123698</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/PMC4395216">4395216</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/25875745">25875745</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=PLOS+ONE&rft.atitle=Soil+Disturbance+as+a+Grassland+Restoration+Measure%E2%80%94Effects+on+Plant+Species+Composition+and+Plant+Functional+Traits&rft.volume=10&rft.issue=4&rft.pages=e0123698&rft.date=2015&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4395216%23id-name%3DPMC&rft_id=info%3Apmid%2F25875745&rft_id=info%3Adoi%2F10.1371%2Fjournal.pone.0123698&rft_id=info%3Abibcode%2F2015PLoSO..1023698S&rft.aulast=Schnoor&rft.aufirst=Tim&rft.au=Bruun%2C+Hans+Henrik&rft.au=Olsson%2C+Pal+Axel&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4395216&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEcological+succession" class="Z3988"></span></span> </li> <li id="cite_note-49"><span class="mw-cite-backlink"><b><a href="#cite_ref-49">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFVan_Auken2000" class="citation journal cs1">Van Auken, O.W. 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"Mechanisms of succession in natural communities and their role in community stability and organization". <i><a href="/wiki/The_American_Naturalist" title="The American Naturalist">The American Naturalist</a></i>. <b>111</b> (982): <span class="nowrap">1119–</span>44. <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/1977ANat..111.1119C">1977ANat..111.1119C</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1086%2F283241">10.1086/283241</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:3587878">3587878</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+American+Naturalist&rft.atitle=Mechanisms+of+succession+in+natural+communities+and+their+role+in+community+stability+and+organization&rft.volume=111&rft.issue=982&rft.pages=%3Cspan+class%3D%22nowrap%22%3E1119-%3C%2Fspan%3E44&rft.date=1977&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A3587878%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1086%2F283241&rft_id=info%3Abibcode%2F1977ANat..111.1119C&rft.aulast=Connell&rft.aufirst=JH&rft.au=Slatyer%2C+RO&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEcological+succession" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFFrouzPrachPižlHáněl2008" class="citation journal cs1">Frouz J, Prach K, Pižl V, Háněl L, Starý J, Tajovský K, et al. (2008). "Interactions between soil development, vegetation and soil fauna during spontaneous succession in post mining sites". <i>European Journal of Soil Biology</i>. <b>44</b> (1): <span class="nowrap">109–</span>121. <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/2008EJSB...44..109F">2008EJSB...44..109F</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.ejsobi.2007.09.002">10.1016/j.ejsobi.2007.09.002</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=European+Journal+of+Soil+Biology&rft.atitle=Interactions+between+soil+development%2C+vegetation+and+soil+fauna+during+spontaneous+succession+in+post+mining+sites.&rft.volume=44&rft.issue=1&rft.pages=%3Cspan+class%3D%22nowrap%22%3E109-%3C%2Fspan%3E121&rft.date=2008&rft_id=info%3Adoi%2F10.1016%2Fj.ejsobi.2007.09.002&rft_id=info%3Abibcode%2F2008EJSB...44..109F&rft.aulast=Frouz&rft.aufirst=J&rft.au=Prach%2C+K&rft.au=Pi%C5%BEl%2C+V&rft.au=H%C3%A1n%C4%9Bl%2C+L&rft.au=Star%C3%BD%2C+J&rft.au=Tajovsk%C3%BD%2C+K&rft.au=Materna%2C+J&rft.au=Balik%2C+V&rft.au=Kal%C4%8D%C3%ADk%2C+J&rft.au=%C5%98ehounkov%C3%A1%2C+K&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEcological+succession" class="Z3988"></span></li></ul> </div> <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=Ecological_succession&action=edit&section=34" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1235681985">.mw-parser-output .side-box{margin:4px 0;box-sizing:border-box;border:1px solid #aaa;font-size:88%;line-height:1.25em;background-color:var(--background-color-interactive-subtle,#f8f9fa);display:flow-root}.mw-parser-output .side-box-abovebelow,.mw-parser-output .side-box-text{padding:0.25em 0.9em}.mw-parser-output .side-box-image{padding:2px 0 2px 0.9em;text-align:center}.mw-parser-output .side-box-imageright{padding:2px 0.9em 2px 0;text-align:center}@media(min-width:500px){.mw-parser-output .side-box-flex{display:flex;align-items:center}.mw-parser-output .side-box-text{flex:1;min-width:0}}@media(min-width:720px){.mw-parser-output .side-box{width:238px}.mw-parser-output .side-box-right{clear:right;float:right;margin-left:1em}.mw-parser-output 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alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/30px-Commons-logo.svg.png" decoding="async" width="30" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/45px-Commons-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/59px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></a></span></div> <div class="side-box-text plainlist">Wikimedia Commons has media related to <span style="font-weight: bold; font-style: italic;"><a href="https://commons.wikimedia.org/wiki/Category:Ecological_succession" class="extiw" title="commons:Category:Ecological succession">Ecological succession</a></span>.</div></div> </div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1235681985"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1237033735"><div class="side-box side-box-right plainlinks sistersitebox"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1126788409"> <div class="side-box-flex"> <div class="side-box-image"><span class="noviewer" typeof="mw:File"><a href="/wiki/File:Wikibooks-logo-en-noslogan.svg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/df/Wikibooks-logo-en-noslogan.svg/40px-Wikibooks-logo-en-noslogan.svg.png" decoding="async" width="40" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/df/Wikibooks-logo-en-noslogan.svg/60px-Wikibooks-logo-en-noslogan.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/df/Wikibooks-logo-en-noslogan.svg/80px-Wikibooks-logo-en-noslogan.svg.png 2x" data-file-width="400" data-file-height="400" /></a></span></div> <div class="side-box-text plainlist">Wikibooks has a book on the topic of: <i><b><a href="https://en.wikibooks.org/wiki/Ecology/Community_succession_and_stability" class="extiw" title="wikibooks:Ecology/Community succession and stability">Ecology/Community succession and stability</a></b></i></div></div> </div> <ul><li><a rel="nofollow" class="external text" href="http://scienceaid.co.uk/biology/ecology/ecosystems.html">Science Aid: Succession</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20210506095947/http://scienceaid.co.uk/biology/ecology/ecosystems.html">Archived</a> 2021-05-06 at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a> Explanation of succession for high school students.</li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20031012214336/http://oz.plymouth.edu/~lts/ecology/ecohistory/cowles.html">Biographical sketch of Henry Chandler Cowles.</a></li> <li><a rel="nofollow" class="external text" href="http://www.edwardgoldsmith.com/page119.html">Robbert Murphy sees a significantly ideological, rather than scientific, basis for the disfavour shown towards succession by the current ecological orthodoxy and seeks to reinstate succession by holistic and teleological argument.</a></li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline ol,.mw-parser-output .hlist.inline ul,.mw-parser-output .hlist dl dl,.mw-parser-output .hlist dl ol,.mw-parser-output .hlist dl ul,.mw-parser-output .hlist ol dl,.mw-parser-output .hlist ol ol,.mw-parser-output .hlist ol ul,.mw-parser-output .hlist ul dl,.mw-parser-output .hlist ul ol,.mw-parser-output .hlist ul ul{display:inline}.mw-parser-output .hlist .mw-empty-li{display:none}.mw-parser-output .hlist dt::after{content:": "}.mw-parser-output .hlist dd::after,.mw-parser-output .hlist li::after{content:" 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href="/wiki/Ecosystem_model" title="Ecosystem model">Modelling ecosystems</a>: <a href="/wiki/Trophic_level" title="Trophic level">Trophic</a> components</div></th></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">General</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Abiotic_component" title="Abiotic component">Abiotic component</a></li> <li><a href="/wiki/Abiotic_stress" title="Abiotic stress">Abiotic stress</a></li> <li><a href="/wiki/Behavioral_ecology" title="Behavioral ecology">Behaviour</a></li> <li><a href="/wiki/Biogeochemical_cycle" title="Biogeochemical cycle">Biogeochemical cycle</a></li> <li><a href="/wiki/Biomass_(ecology)" title="Biomass (ecology)">Biomass</a></li> <li><a href="/wiki/Biotic_component" class="mw-redirect" title="Biotic component">Biotic component</a></li> <li><a href="/wiki/Biotic_stress" title="Biotic stress">Biotic stress</a></li> <li><a href="/wiki/Carrying_capacity" title="Carrying capacity">Carrying capacity</a></li> <li><a href="/wiki/Competition_(biology)" title="Competition (biology)">Competition</a></li> <li><a href="/wiki/Ecosystem" title="Ecosystem">Ecosystem</a></li> <li><a href="/wiki/Ecosystem_ecology" title="Ecosystem ecology">Ecosystem ecology</a></li> <li><a href="/wiki/Ecosystem_model" title="Ecosystem model">Ecosystem model</a></li> <li><a href="/wiki/Green_world_hypothesis" title="Green world hypothesis">Green world hypothesis</a></li> <li><a href="/wiki/Keystone_species" title="Keystone species">Keystone species</a></li> <li><a href="/wiki/List_of_feeding_behaviours" title="List of feeding behaviours">List of feeding behaviours</a></li> <li><a href="/wiki/Metabolic_theory_of_ecology" title="Metabolic theory of ecology">Metabolic theory of ecology</a></li> <li><a href="/wiki/Productivity_(ecology)" title="Productivity (ecology)">Productivity</a></li> <li><a href="/wiki/Resource_(biology)" title="Resource (biology)">Resource</a></li> <li><a href="/wiki/Restoration_ecology" class="mw-redirect" title="Restoration ecology">Restoration</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Autotroph" title="Autotroph">Producers</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Autotroph" title="Autotroph">Autotrophs</a></li> <li><a href="/wiki/Chemosynthesis" title="Chemosynthesis">Chemosynthesis</a></li> <li><a href="/wiki/Chemotroph" title="Chemotroph">Chemotrophs</a></li> <li><a href="/wiki/Foundation_species" title="Foundation species">Foundation species</a></li> <li><a href="/wiki/Kinetotroph" title="Kinetotroph">Kinetotrophs</a></li> <li><a href="/wiki/Mixotroph" title="Mixotroph">Mixotrophs</a></li> <li><a href="/wiki/Myco-heterotrophy" title="Myco-heterotrophy">Myco-heterotrophy</a></li> <li><a href="/wiki/Mycotroph" title="Mycotroph">Mycotroph</a></li> <li><a href="/wiki/Organotroph" title="Organotroph">Organotrophs</a></li> <li><a href="/wiki/Photoheterotroph" title="Photoheterotroph">Photoheterotrophs</a></li> <li><a href="/wiki/Photosynthesis" title="Photosynthesis">Photosynthesis</a></li> <li><a href="/wiki/Photosynthetic_efficiency" title="Photosynthetic efficiency">Photosynthetic efficiency</a></li> <li><a href="/wiki/Phototroph" title="Phototroph">Phototrophs</a></li> <li><a href="/wiki/Primary_nutritional_groups" title="Primary nutritional groups">Primary nutritional groups</a></li> <li><a href="/wiki/Primary_production" title="Primary production">Primary production</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Consumer_(food_chain)" title="Consumer (food chain)">Consumers</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Apex_predator" title="Apex predator">Apex predator</a></li> <li><a href="/wiki/Bacterivore" title="Bacterivore">Bacterivore</a></li> <li><a href="/wiki/Carnivore" title="Carnivore">Carnivores</a></li> <li><a href="/wiki/Chemoorganotroph" class="mw-redirect" title="Chemoorganotroph">Chemoorganotroph</a></li> <li><a href="/wiki/Foraging" title="Foraging">Foraging</a></li> <li><a href="/wiki/Generalist_and_specialist_species" title="Generalist and specialist species">Generalist and specialist species</a></li> <li><a href="/wiki/Intraguild_predation" title="Intraguild predation">Intraguild predation</a></li> <li><a href="/wiki/Herbivore" title="Herbivore">Herbivores</a></li> <li><a href="/wiki/Heterotroph" title="Heterotroph">Heterotroph</a></li> <li><a href="/wiki/Heterotrophic_nutrition" title="Heterotrophic nutrition">Heterotrophic nutrition</a></li> <li><a href="/wiki/Insectivore" title="Insectivore">Insectivore</a></li> <li><a href="/wiki/Mesopredator" title="Mesopredator">Mesopredators</a></li> <li><a href="/wiki/Mesopredator_release_hypothesis" title="Mesopredator release hypothesis">Mesopredator release hypothesis</a></li> <li><a href="/wiki/Omnivore" title="Omnivore">Omnivores</a></li> <li><a href="/wiki/Optimal_foraging_theory" title="Optimal foraging theory">Optimal foraging theory</a></li> <li><a href="/wiki/Planktivore" title="Planktivore">Planktivore</a></li> <li><a href="/wiki/Predation" title="Predation">Predation</a></li> <li><a href="/wiki/Prey_switching" title="Prey switching">Prey switching</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Decomposer" title="Decomposer">Decomposers</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Chemoorganoheterotrophy" class="mw-redirect" title="Chemoorganoheterotrophy">Chemoorganoheterotrophy</a></li> <li><a href="/wiki/Decomposition" title="Decomposition">Decomposition</a></li> <li><a href="/wiki/Detritivore" title="Detritivore">Detritivores</a></li> <li><a href="/wiki/Detritus" title="Detritus">Detritus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Microorganism#Habitats_and_ecology" title="Microorganism">Microorganisms</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Archaea" title="Archaea">Archaea</a></li> <li><a href="/wiki/Bacteriophage" title="Bacteriophage">Bacteriophage</a></li> <li><a href="/wiki/Lithoautotroph" title="Lithoautotroph">Lithoautotroph</a></li> <li><a href="/wiki/Lithotroph" title="Lithotroph">Lithotrophy</a></li> <li><a href="/wiki/Marine_microorganisms" title="Marine microorganisms">Marine</a></li> <li><a href="/wiki/Microbial_cooperation" title="Microbial cooperation">Microbial cooperation</a></li> <li><a href="/wiki/Microbial_ecology" title="Microbial ecology">Microbial ecology</a></li> <li><a href="/wiki/Microbial_food_web" title="Microbial food web">Microbial food web</a></li> <li><a href="/wiki/Microbial_intelligence" title="Microbial intelligence">Microbial intelligence</a></li> <li><a href="/wiki/Microbial_loop" title="Microbial loop">Microbial loop</a></li> <li><a href="/wiki/Microbial_mat" title="Microbial mat">Microbial mat</a></li> <li><a href="/wiki/Microbial_metabolism" title="Microbial metabolism">Microbial metabolism</a></li> <li><a href="/wiki/Phage_ecology" title="Phage ecology">Phage ecology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Food_web" title="Food web">Food webs</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Biomagnification" title="Biomagnification">Biomagnification</a></li> <li><a href="/wiki/Ecological_efficiency" title="Ecological efficiency">Ecological efficiency</a></li> <li><a href="/wiki/Ecological_pyramid" title="Ecological pyramid">Ecological pyramid</a></li> <li><a href="/wiki/Energy_flow_(ecology)" title="Energy flow (ecology)">Energy flow</a></li> <li><a href="/wiki/Food_chain" title="Food chain">Food chain</a></li> <li><a href="/wiki/Trophic_level" title="Trophic level">Trophic level</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Example webs</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Lake_ecosystem#Trophic_relationships" title="Lake ecosystem">Lakes</a></li> <li><a href="/wiki/River_ecosystem#Trophic_relationships" title="River ecosystem">Rivers</a></li> <li><a href="/wiki/Soil_food_web" title="Soil food web">Soil</a></li> <li><a href="/wiki/Tritrophic_interactions_in_plant_defense" title="Tritrophic interactions in plant defense">Tritrophic interactions in plant defense</a></li> <li><a href="/wiki/Marine_food_web" title="Marine food web">Marine food webs</a> <ul><li><a href="/wiki/Cold_seep" title="Cold seep">cold seeps</a></li> <li><a href="/wiki/Hydrothermal_vent#Biological_communities" title="Hydrothermal vent">hydrothermal vents</a></li> <li><a href="/wiki/Intertidal_ecology" title="Intertidal ecology">intertidal</a></li> <li><a href="/wiki/Kelp_forest#Trophic_ecology" title="Kelp forest">kelp forests</a></li> <li><a href="/wiki/North_Pacific_Gyre" title="North Pacific Gyre">North Pacific Gyre</a></li> <li><a href="/wiki/Ecology_of_the_San_Francisco_Estuary#Food_web" title="Ecology of the San Francisco Estuary">San Francisco Estuary</a></li> <li><a href="/wiki/Tide_pool" title="Tide pool">tide pool</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Processes</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Ascendency" title="Ascendency">Ascendency</a></li> <li><a href="/wiki/Bioaccumulation" title="Bioaccumulation">Bioaccumulation</a></li> <li><a href="/wiki/Cascade_effect_(ecology)" title="Cascade effect (ecology)">Cascade effect</a></li> <li><a href="/wiki/Climax_community" title="Climax community">Climax community</a></li> <li><a href="/wiki/Competitive_exclusion_principle" title="Competitive exclusion principle">Competitive exclusion principle</a></li> <li><a href="/wiki/Consumer%E2%80%93resource_interactions" title="Consumer–resource interactions">Consumer–resource interactions</a></li> <li><a href="/wiki/Copiotroph" title="Copiotroph">Copiotrophs</a></li> <li><a href="/wiki/Dominance_(ecology)" title="Dominance (ecology)">Dominance</a></li> <li><a href="/wiki/Ecological_network" title="Ecological network">Ecological network</a></li> <li><a class="mw-selflink selflink">Ecological succession</a></li> <li><a href="/wiki/Energy_quality" title="Energy quality">Energy quality</a></li> <li><a href="/wiki/Energy_systems_language" title="Energy systems language">Energy systems language</a></li> <li><a href="/wiki/F-ratio_(oceanography)" title="F-ratio (oceanography)">f-ratio</a></li> <li><a href="/wiki/Feed_conversion_ratio" title="Feed conversion ratio">Feed conversion ratio</a></li> <li><a href="/wiki/Feeding_frenzy" title="Feeding frenzy">Feeding frenzy</a></li> <li><a href="/wiki/Mesotrophic_soil" title="Mesotrophic soil">Mesotrophic soil</a></li> <li><a href="/wiki/Nutrient_cycle" title="Nutrient cycle">Nutrient cycle</a></li> <li><a href="/wiki/Oligotroph" title="Oligotroph">Oligotroph</a></li> <li><a href="/wiki/Paradox_of_the_plankton" title="Paradox of the plankton">Paradox of the plankton</a></li> <li><a href="/wiki/Trophic_cascade" title="Trophic cascade">Trophic cascade</a></li> <li><a href="/wiki/Trophic_mutualism" title="Trophic mutualism">Trophic mutualism</a></li> <li><a href="/wiki/Trophic_state_index" title="Trophic state index">Trophic state index</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Defense,<br />counter</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Animal_coloration" title="Animal coloration">Animal coloration</a></li> <li><a href="/wiki/Anti-predator_adaptation" title="Anti-predator adaptation">Anti-predator adaptations</a></li> <li><a href="/wiki/Camouflage" title="Camouflage">Camouflage</a></li> <li><a href="/wiki/Deimatic_behaviour" title="Deimatic behaviour">Deimatic behaviour</a></li> <li><a href="/wiki/Herbivore_adaptations_to_plant_defense" title="Herbivore adaptations to plant defense">Herbivore adaptations to plant defense</a></li> <li><a href="/wiki/Mimicry" title="Mimicry">Mimicry</a></li> <li><a href="/wiki/Plant_defense_against_herbivory" title="Plant defense against herbivory">Plant defense against herbivory</a></li> <li><a href="/wiki/Shoaling_and_schooling#Predator_avoidance" title="Shoaling and schooling">Predator avoidance in schooling fish</a></li></ul> </div></td></tr></tbody></table></div><div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Ecology:_Modelling_ecosystems:_Other_components285" style="padding:3px"><table class="nowraplinks hlist mw-collapsible mw-collapsed navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Modelling_ecosystems" title="Template:Modelling ecosystems"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Modelling_ecosystems" title="Template talk:Modelling ecosystems"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Modelling_ecosystems" title="Special:EditPage/Template:Modelling ecosystems"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Ecology:_Modelling_ecosystems:_Other_components285" style="font-size:114%;margin:0 4em"><a href="/wiki/Ecology" title="Ecology">Ecology</a>: <a href="/wiki/Ecosystem_model" title="Ecosystem model">Modelling ecosystems</a>: Other components</div></th></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Population_ecology" title="Population ecology">Population<br />ecology</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Abundance_(ecology)" title="Abundance (ecology)">Abundance</a></li> <li><a href="/wiki/Allee_effect" title="Allee effect">Allee effect</a></li> <li><a href="/wiki/Consumer-resource_model" title="Consumer-resource model">Consumer-resource model</a></li> <li><a href="/wiki/Depensation" title="Depensation">Depensation</a></li> <li><a href="/wiki/Ecological_yield" title="Ecological yield">Ecological yield</a></li> <li><a href="/wiki/Effective_population_size" title="Effective population size">Effective population size</a></li> <li><a href="/wiki/Intraspecific_competition" title="Intraspecific competition">Intraspecific competition</a></li> <li><a href="/wiki/Logistic_function" title="Logistic function">Logistic function</a></li> <li><a href="/wiki/Malthusian_growth_model" title="Malthusian growth model">Malthusian growth model</a></li> <li><a href="/wiki/Maximum_sustainable_yield" title="Maximum sustainable yield">Maximum sustainable yield</a></li> <li><a href="/wiki/Overpopulation" title="Overpopulation">Overpopulation</a></li> <li><a href="/wiki/Overexploitation" title="Overexploitation">Overexploitation</a></li> <li><a href="/wiki/Population_cycle" title="Population cycle">Population cycle</a></li> <li><a href="/wiki/Population_dynamics" title="Population dynamics">Population dynamics</a></li> <li><a href="/wiki/Population_model" title="Population model">Population modeling</a></li> <li><a href="/wiki/Population_size" title="Population size">Population size</a></li> <li><a href="/wiki/Lotka%E2%80%93Volterra_equations" title="Lotka–Volterra equations">Predator–prey (Lotka–Volterra) equations</a></li> <li><a href="/wiki/Recruitment_(biology)" title="Recruitment (biology)">Recruitment</a></li> <li><a href="/wiki/Small_population_size" title="Small population size">Small population size</a></li> <li><a href="/wiki/Ecological_stability" title="Ecological stability">Stability</a> <ul><li><a href="/wiki/Ecological_resilience" title="Ecological resilience">Resilience</a></li> <li><a href="/wiki/Resistance_(ecology)" title="Resistance (ecology)">Resistance</a></li></ul></li> <li><a href="/wiki/Random_generalized_Lotka%E2%80%93Volterra_model" title="Random generalized Lotka–Volterra model">Random generalized Lotka–Volterra model</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Species</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Biodiversity" title="Biodiversity">Biodiversity</a></li> <li><a href="/wiki/Density_dependence" title="Density dependence">Density-dependent inhibition</a></li> <li><a href="/wiki/Ecological_effects_of_biodiversity" title="Ecological effects of biodiversity">Ecological effects of biodiversity</a></li> <li><a href="/wiki/Ecological_extinction" title="Ecological extinction">Ecological extinction</a></li> <li><a href="/wiki/Endemism" title="Endemism">Endemic species</a></li> <li><a href="/wiki/Flagship_species" title="Flagship species">Flagship species</a></li> <li><a href="/wiki/Ordination_(statistics)" title="Ordination (statistics)">Gradient analysis</a></li> <li><a href="/wiki/Bioindicator" title="Bioindicator">Indicator species</a></li> <li><a href="/wiki/Introduced_species" title="Introduced species">Introduced species</a></li> <li><a href="/wiki/Invasive_species" title="Invasive species">Invasive species</a> / <a href="/wiki/Native_species" title="Native species">Native species</a></li> <li><a href="/wiki/Latitudinal_gradients_in_species_diversity" title="Latitudinal gradients in species diversity">Latitudinal gradients in species diversity</a></li> <li><a href="/wiki/Minimum_viable_population" title="Minimum viable population">Minimum viable population</a></li> <li><a href="/wiki/Unified_neutral_theory_of_biodiversity" title="Unified neutral theory of biodiversity">Neutral theory</a></li> <li><a href="/wiki/Occupancy%E2%80%93abundance_relationship" title="Occupancy–abundance relationship">Occupancy–abundance relationship</a></li> <li><a href="/wiki/Population_viability_analysis" title="Population viability analysis">Population viability analysis</a></li> <li><a href="/wiki/Priority_effect" title="Priority effect">Priority effect</a></li> <li><a href="/wiki/Rapoport%27s_rule" title="Rapoport's rule">Rapoport's rule</a></li> <li><a href="/wiki/Relative_abundance_distribution" title="Relative abundance distribution">Relative abundance distribution</a></li> <li><a href="/wiki/Relative_species_abundance" title="Relative species abundance">Relative species abundance</a></li> <li><a href="/wiki/Species_diversity" title="Species diversity">Species diversity</a></li> <li><a href="/wiki/Species_homogeneity" title="Species homogeneity">Species homogeneity</a></li> <li><a href="/wiki/Species_richness" title="Species richness">Species richness</a></li> <li><a href="/wiki/Species_distribution" title="Species distribution">Species distribution</a></li> <li><a href="/wiki/Species%E2%80%93area_relationship" title="Species–area relationship">Species–area curve</a></li> <li><a href="/wiki/Umbrella_species" title="Umbrella species">Umbrella species</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Species<br />interaction</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Antibiosis" title="Antibiosis">Antibiosis</a></li> <li><a href="/wiki/Biological_interaction" title="Biological interaction">Biological interaction</a></li> <li><a href="/wiki/Commensalism" title="Commensalism">Commensalism</a></li> <li><a href="/wiki/Community_(ecology)" title="Community (ecology)">Community ecology</a></li> <li><a href="/wiki/Ecological_facilitation" title="Ecological facilitation">Ecological facilitation</a></li> <li><a href="/wiki/Interspecific_competition" title="Interspecific competition">Interspecific competition</a></li> <li><a href="/wiki/Mutualism_(biology)" title="Mutualism (biology)">Mutualism</a></li> <li><a href="/wiki/Parasitism" title="Parasitism">Parasitism</a></li> <li><a href="/wiki/Storage_effect" title="Storage effect">Storage effect</a></li> <li><a href="/wiki/Symbiosis" title="Symbiosis">Symbiosis</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Spatial_ecology" title="Spatial ecology">Spatial<br />ecology</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Biogeography" title="Biogeography">Biogeography</a></li> <li><a href="/wiki/Cross-boundary_subsidy" title="Cross-boundary subsidy">Cross-boundary subsidy</a></li> <li><a href="/wiki/Cline_(biology)" title="Cline (biology)">Ecocline</a></li> <li><a href="/wiki/Ecotone" title="Ecotone">Ecotone</a></li> <li><a href="/wiki/Ecotype" title="Ecotype">Ecotype</a></li> <li><a href="/wiki/Disturbance_(ecology)" title="Disturbance (ecology)">Disturbance</a></li> <li><a href="/wiki/Edge_effects" title="Edge effects">Edge effects</a></li> <li><a href="/wiki/Foster%27s_rule" title="Foster's rule">Foster's rule</a></li> <li><a href="/wiki/Habitat_fragmentation" title="Habitat fragmentation">Habitat fragmentation</a></li> <li><a href="/wiki/Ideal_free_distribution" title="Ideal free distribution">Ideal free distribution</a></li> <li><a href="/wiki/Intermediate_disturbance_hypothesis" title="Intermediate disturbance hypothesis">Intermediate disturbance hypothesis</a></li> <li><a href="/wiki/Insular_biogeography" title="Insular biogeography">Insular biogeography</a></li> <li><a href="/wiki/Land_change_modeling" title="Land change modeling">Land change modeling</a></li> <li><a href="/wiki/Landscape_ecology" title="Landscape ecology">Landscape ecology</a></li> <li><a href="/wiki/Landscape_epidemiology" title="Landscape epidemiology">Landscape epidemiology</a></li> <li><a href="/wiki/Landscape_limnology" title="Landscape limnology">Landscape limnology</a></li> <li><a href="/wiki/Metapopulation" title="Metapopulation">Metapopulation</a></li> <li><a href="/wiki/Patch_dynamics" title="Patch dynamics">Patch dynamics</a></li> <li><a href="/wiki/R/K_selection_theory" title="R/K selection theory"><i>r</i>/<i>K</i> selection theory</a></li> <li><a href="/wiki/Resource_selection_function" title="Resource selection function">Resource selection function</a></li> <li><a href="/wiki/Source%E2%80%93sink_dynamics" title="Source–sink dynamics">Source–sink dynamics</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Ecological_niche" title="Ecological niche">Niche</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Ecological_trap" title="Ecological trap">Ecological trap</a></li> <li><a href="/wiki/Ecosystem_engineer" title="Ecosystem engineer">Ecosystem engineer</a></li> <li><a href="/wiki/Species_distribution_modelling" title="Species distribution modelling">Environmental niche modelling</a></li> <li><a href="/wiki/Guild_(ecology)" title="Guild (ecology)">Guild</a></li> <li><a href="/wiki/Habitat" title="Habitat">Habitat</a> <ul><li><a href="/wiki/Marine_habitat" title="Marine habitat">Marine</a></li> <li><a href="/wiki/Semiaquatic" title="Semiaquatic">Semiaquatic</a></li> <li><a href="/wiki/Terrestrial_animal" title="Terrestrial animal">Terrestrial</a></li></ul></li> <li><a href="/wiki/Limiting_similarity" title="Limiting similarity">Limiting similarity</a></li> <li><a href="/wiki/Niche_apportionment_models" title="Niche apportionment models">Niche apportionment models</a></li> <li><a href="/wiki/Niche_construction" title="Niche construction">Niche construction</a></li> <li><a href="/wiki/Niche_differentiation" class="mw-redirect" title="Niche differentiation">Niche differentiation</a></li> <li><a href="/wiki/Ontogenetic_niche_shift" title="Ontogenetic niche shift">Ontogenetic niche shift</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Non-trophic_networks" title="Non-trophic networks">Other<br />networks</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Assembly_rules" title="Assembly rules">Assembly rules</a></li> <li><a href="/wiki/Bateman%27s_principle" title="Bateman's principle">Bateman's principle</a></li> <li><a href="/wiki/Bioluminescence" title="Bioluminescence">Bioluminescence</a></li> <li><a href="/wiki/Ecological_collapse" class="mw-redirect" title="Ecological collapse">Ecological collapse</a></li> <li><a href="/wiki/Ecological_debt" title="Ecological debt">Ecological debt</a></li> <li><a href="/wiki/Ecological_debt" title="Ecological debt">Ecological deficit</a></li> <li><a href="/wiki/Energy_flow_(ecology)" title="Energy flow (ecology)">Ecological energetics</a></li> <li><a href="/wiki/Ecological_indicator" title="Ecological indicator">Ecological indicator</a></li> <li><a href="/wiki/Ecological_threshold" title="Ecological threshold">Ecological threshold</a></li> <li><a href="/wiki/Ecosystem_diversity" title="Ecosystem diversity">Ecosystem diversity</a></li> <li><a href="/wiki/Emergence" title="Emergence">Emergence</a></li> <li><a href="/wiki/Extinction_debt" title="Extinction debt">Extinction debt</a></li> <li><a href="/wiki/Kleiber%27s_law" title="Kleiber's law">Kleiber's law</a></li> <li><a href="/wiki/Liebig%27s_law_of_the_minimum" title="Liebig's law of the minimum">Liebig's law of the minimum</a></li> <li><a href="/wiki/Marginal_value_theorem" title="Marginal value theorem">Marginal value theorem</a></li> <li><a href="/wiki/Thorson%27s_rule" title="Thorson's rule">Thorson's rule</a></li> <li><a href="/wiki/Xerosere" title="Xerosere">Xerosere</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Other</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Allometry" title="Allometry">Allometry</a></li> <li><a href="/wiki/Alternative_stable_state" title="Alternative stable state">Alternative stable state</a></li> <li><a href="/wiki/Balance_of_nature" title="Balance of nature">Balance of nature</a></li> <li><a href="/wiki/Biological_data_visualization" title="Biological data visualization">Biological data visualization</a></li> <li><a href="/wiki/Ecological_economics" title="Ecological economics">Ecological economics</a></li> <li><a href="/wiki/Ecological_footprint" title="Ecological footprint">Ecological footprint</a></li> <li><a href="/wiki/Ecological_forecasting" title="Ecological forecasting">Ecological forecasting</a></li> <li><a href="/wiki/Environmental_humanities" title="Environmental humanities">Ecological humanities</a></li> <li><a href="/wiki/Ecological_stoichiometry" title="Ecological stoichiometry">Ecological stoichiometry</a></li> <li><a href="/wiki/Ecopath" title="Ecopath">Ecopath</a></li> <li><a href="/wiki/Ecosystem_based_fisheries" class="mw-redirect" title="Ecosystem based fisheries">Ecosystem based fisheries</a></li> <li><a href="/wiki/Endolith" title="Endolith">Endolith</a></li> <li><a href="/wiki/Evolutionary_ecology" title="Evolutionary ecology">Evolutionary ecology</a></li> <li><a href="/wiki/Functional_ecology" title="Functional ecology">Functional ecology</a></li> <li><a href="/wiki/Industrial_ecology" title="Industrial ecology">Industrial ecology</a></li> <li><a href="/wiki/Macroecology" title="Macroecology">Macroecology</a></li> <li><a href="/wiki/Microecosystem" title="Microecosystem">Microecosystem</a></li> <li><a href="/wiki/Natural_environment" title="Natural environment">Natural environment</a></li> <li><a href="/wiki/Regime_shift" title="Regime shift">Regime shift</a></li> <li><a href="/wiki/Sexecology" title="Sexecology">Sexecology</a></li> <li><a href="/wiki/Systems_ecology" title="Systems ecology">Systems ecology</a></li> <li><a href="/wiki/Urban_ecology" title="Urban ecology">Urban ecology</a></li> <li><a href="/wiki/Theoretical_ecology" title="Theoretical ecology">Theoretical ecology</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div><a href="/wiki/Outline_of_ecology" title="Outline of ecology">Outline of ecology</a></div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"><style data-mw-deduplicate="TemplateStyles:r1038841319">.mw-parser-output .tooltip-dotted{border-bottom:1px dotted;cursor:help}</style></div><div role="navigation" 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