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href="#Secondary_contact"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1.2</span> <span>Secondary contact</span> </div> </a> <ul id="toc-Secondary_contact-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-No_selection_(drift/migration_balance)" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#No_selection_(drift/migration_balance)"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.1.3</span> <span>No selection (drift/migration balance)</span> </div> </a> <ul id="toc-No_selection_(drift/migration_balance)-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> </ul> </li> <li id="toc-Clinal_structure_and_terminology" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Clinal_structure_and_terminology"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Clinal structure and terminology</span> </div> </a> <ul id="toc-Clinal_structure_and_terminology-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Types_of_clines" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Types_of_clines"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Types of clines</span> </div> </a> <button aria-controls="toc-Types_of_clines-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 clines subsection</span> </button> <ul id="toc-Types_of_clines-sublist" class="vector-toc-list"> <li id="toc-Continuous_clines" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Continuous_clines"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.1</span> <span>Continuous clines</span> </div> </a> <ul id="toc-Continuous_clines-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Discontinuous_stepped_clines" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Discontinuous_stepped_clines"> <div class="vector-toc-text"> <span class="vector-toc-numb">3.2</span> <span>Discontinuous stepped clines</span> </div> </a> <ul id="toc-Discontinuous_stepped_clines-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Clines_and_speciation" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Clines_and_speciation"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Clines and speciation</span> </div> </a> <ul id="toc-Clines_and_speciation-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Clinal_maps" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Clinal_maps"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Clinal maps</span> </div> </a> <ul id="toc-Clinal_maps-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Examples_of_clines" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Examples_of_clines"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>Examples of clines</span> </div> </a> <ul id="toc-Examples_of_clines-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> </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 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class="mw-page-title-main">Cline (biology)</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 23 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-23" 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">23 languages</span> </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%AA%D9%86%D9%88%D8%B9_%D8%A7%D9%84%D8%B3%D9%85%D8%A7%D8%AA_(%D8%B9%D9%84%D9%85_%D8%A7%D9%84%D8%A3%D8%AD%D9%8A%D8%A7%D8%A1)" title="تنوع السمات (علم الأحياء) – Arabic" lang="ar" hreflang="ar" data-title="تنوع السمات (علم الأحياء)" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target"><span>العربية</span></a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Klina" title="Klina – Bosnian" lang="bs" hreflang="bs" data-title="Klina" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target"><span>Bosanski</span></a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Clina" title="Clina – Catalan" lang="ca" hreflang="ca" data-title="Clina" 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/Klin%C3%A1ln%C3%AD_variabilita" title="Klinální variabilita – Czech" lang="cs" hreflang="cs" data-title="Klinální variabilita" 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/Klinal_variation" title="Klinal variation – Danish" lang="da" hreflang="da" data-title="Klinal variation" 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/Kline_(Biologie)" title="Kline (Biologie) – German" lang="de" hreflang="de" data-title="Kline (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/%C3%96kokliin" title="Ökokliin – Estonian" lang="et" hreflang="et" data-title="Ökokliin" 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/Clina" title="Clina – Spanish" lang="es" hreflang="es" data-title="Clina" 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/Klino_(biologio)" title="Klino (biologio) – Esperanto" lang="eo" hreflang="eo" data-title="Klino (biologio)" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target"><span>Esperanto</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%DA%A9%D9%84%D8%A7%DB%8C%D9%86_(%D8%B2%DB%8C%D8%B3%D8%AA%E2%80%8C%D8%B4%D9%86%D8%A7%D8%B3%DB%8C)" title="کلاین (زیست‌شناسی) – Persian" lang="fa" hreflang="fa" data-title="کلاین (زیست‌شناسی)" 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/Cline_syst%C3%A9matique" title="Cline systématique – French" lang="fr" hreflang="fr" data-title="Cline systématique" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Variazione_clinale" title="Variazione clinale – Italian" lang="it" hreflang="it" data-title="Variazione clinale" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target"><span>Italiano</span></a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/%C3%98koklin" title="Økoklin – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Økoklin" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target"><span>Norsk bokmål</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Ekoklina" title="Ekoklina – Polish" lang="pl" hreflang="pl" data-title="Ekoklina" 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/Varia%C3%A7%C3%A3o_clinal" title="Variação clinal – Portuguese" lang="pt" hreflang="pt" data-title="Variação clinal" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/Varia%C8%9Bie_clinal%C4%83" title="Variație clinală – Romanian" lang="ro" hreflang="ro" data-title="Variație clinală" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target"><span>Română</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%9A%D0%BB%D0%B8%D0%BD%D0%B0%D0%BB%D1%8C%D0%BD%D0%B0%D1%8F_%D0%B8%D0%B7%D0%BC%D0%B5%D0%BD%D1%87%D0%B8%D0%B2%D0%BE%D1%81%D1%82%D1%8C" title="Клинальная изменчивость – Russian" lang="ru" hreflang="ru" data-title="Клинальная изменчивость" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Cline" title="Cline – Simple English" lang="en-simple" hreflang="en-simple" data-title="Cline" 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-sr mw-list-item"><a href="https://sr.wikipedia.org/wiki/%D0%9A%D0%BB%D0%B8%D0%BD%D0%B0_(%D0%B1%D0%B8%D0%BE%D0%BB%D0%BE%D0%B3%D0%B8%D1%98%D0%B0)" title="Клина (биологија) – Serbian" lang="sr" hreflang="sr" data-title="Клина (биологија)" 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/Kliini" title="Kliini – Finnish" lang="fi" hreflang="fi" data-title="Kliini" 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/Klinal_variation" title="Klinal variation – Swedish" lang="sv" hreflang="sv" data-title="Klinal variation" 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/Klin_(biyoloji)" title="Klin (biyoloji) – Turkish" lang="tr" hreflang="tr" data-title="Klin (biyoloji)" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target"><span>Türkçe</span></a></li><li class="interlanguage-link interwiki-uk mw-list-item"><a href="https://uk.wikipedia.org/wiki/%D0%9A%D0%BB%D1%96%D0%BD%D0%B0" title="Кліна – Ukrainian" lang="uk" hreflang="uk" data-title="Кліна" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target"><span>Українська</span></a></li> </ul> <div class="after-portlet after-portlet-lang"><span class="wb-langlinks-edit wb-langlinks-link"><a href="https://www.wikidata.org/wiki/Special:EntityPage/Q966735#sitelinks-wikipedia" title="Edit interlanguage links" class="wbc-editpage">Edit links</a></span></div> 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.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">For other uses, see <a href="/wiki/Cline_(disambiguation)" class="mw-redirect mw-disambig" title="Cline (disambiguation)">Cline</a>.</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">"Gene cline" redirects here. For the baseball player, see <a href="/wiki/Gene_Clines" title="Gene Clines">Gene Clines</a>. For people named Gene Klein, see <a href="/wiki/Eugene_Klein_(disambiguation)" class="mw-redirect mw-disambig" title="Eugene Klein (disambiguation)">Eugene Klein (disambiguation)</a>.</div> <p>In <a href="/wiki/Biology" title="Biology">biology</a>, a <b>cline</b> is a measurable <a href="/wiki/Gradient" title="Gradient">gradient</a> in a single characteristic (or biological trait) of a species across its <a href="/wiki/Range_(biology)" class="mw-redirect" title="Range (biology)">geographical range</a>.<sup id="cite_ref-Huxley_1-0" class="reference"><a href="#cite_note-Huxley-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Clines usually have a genetic (e.g. <a href="/wiki/Allele_frequency" title="Allele frequency">allele frequency</a>, <a href="/wiki/Blood_type" title="Blood type">blood type</a>), or phenotypic (e.g. body size, skin pigmentation) character. They can show either smooth, continuous gradation in a character, or more abrupt changes in the trait from one geographic region to the next.<sup id="cite_ref-Endler_2-0" class="reference"><a href="#cite_note-Endler-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p><p>A cline is a spatial gradient in a single specific trait, rather than in a collection of traits;<sup id="cite_ref-Huxley_biography_3-0" class="reference"><a href="#cite_note-Huxley_biography-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> a single population can therefore have as many clines as it has traits, at least in principle.<sup id="cite_ref-Forest_genetics_4-0" class="reference"><a href="#cite_note-Forest_genetics-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Additionally, as <a href="/wiki/Julian_Huxley" title="Julian Huxley">Julian Huxley</a> recognised, these multiple independent clines may not act in concordance with each other. For example, it has been observed that in Australia, birds generally become smaller the further towards the north of the country they are found. In contrast, the intensity of their plumage colouration follows a different geographical trajectory, being most vibrant where humidity is highest and becoming less vibrant further into the arid centre of the country.<sup id="cite_ref-Mayr1963_5-0" class="reference"><a href="#cite_note-Mayr1963-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> Because of this, Huxley described the notion of clines as an "auxiliary taxonomic principle,” meaning that clinal variation in a species is not awarded taxonomic recognition in the way <a href="/wiki/Subspecies" title="Subspecies">subspecies</a> or <a href="/wiki/Species" title="Species">species</a> are.<sup id="cite_ref-Huxley_1-1" class="reference"><a href="#cite_note-Huxley-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>The term <i>cline</i> was coined by Huxley in 1938 from the <a href="/wiki/Ancient_Greek" title="Ancient Greek">Greek</a> κλίνειν <i>klinein</i>, meaning "to lean.” While it and the term <i><a href="/wiki/Ecotype" title="Ecotype">ecotype</a></i> are sometimes used interchangeably, they do in fact differ in that <i>ecotype</i> refers to a population which differs from other populations in a number of characters, rather than the single character that varies amongst populations in a cline.<sup id="cite_ref-Forest_genetics_4-1" class="reference"><a href="#cite_note-Forest_genetics-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Drivers_and_the_evolution_of_clines">Drivers and the evolution of clines</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cline_(biology)&action=edit&section=1" title="Edit section: Drivers and the evolution of clines"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Gene_flow_(butterflies).svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/92/Gene_flow_%28butterflies%29.svg/220px-Gene_flow_%28butterflies%29.svg.png" decoding="async" width="220" height="68" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/92/Gene_flow_%28butterflies%29.svg/330px-Gene_flow_%28butterflies%29.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/92/Gene_flow_%28butterflies%29.svg/440px-Gene_flow_%28butterflies%29.svg.png 2x" data-file-width="1888" data-file-height="584" /></a><figcaption>Two populations with individuals moving between the populations to demonstrate gene flow</figcaption></figure> <p>Clines are often cited to be the result of two opposing drivers: <a href="/wiki/Natural_selection" title="Natural selection">selection</a> and <a href="/wiki/Gene_flow" title="Gene flow">gene flow</a> (also known as migration).<sup id="cite_ref-Mayr1963_5-1" class="reference"><a href="#cite_note-Mayr1963-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> Selection causes <a href="/wiki/Adaptation" title="Adaptation">adaptation</a> to the local environment, resulting in different genotypes or phenotypes being favoured in different environments. This diversifying force is countered by gene flow, which has a homogenising effect on populations and prevents <a href="/wiki/Speciation" title="Speciation">speciation</a> through causing <a href="/wiki/Genetic_admixture" title="Genetic admixture">genetic admixture</a> and blurring any distinct genetic boundaries.<sup id="cite_ref-Mayr1963_5-2" class="reference"><a href="#cite_note-Mayr1963-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Development_of_clines">Development of clines</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cline_(biology)&action=edit&section=2" title="Edit section: Development of clines"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Clines are generally thought to arise under one of two conditions: "primary differentiation" (also known as "primary contact" or "primary intergradation"), or "secondary contact" (also known as "secondary introgression", or "secondary intergradation").<sup id="cite_ref-Endler_2-1" class="reference"><a href="#cite_note-Endler-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Mayr1942_6-0" class="reference"><a href="#cite_note-Mayr1942-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Joseph_7-0" class="reference"><a href="#cite_note-Joseph-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Primary_differentiation">Primary differentiation</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cline_(biology)&action=edit&section=3" title="Edit section: Primary differentiation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Final_Primary_Differentiation.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/37/Final_Primary_Differentiation.svg/220px-Final_Primary_Differentiation.svg.png" decoding="async" width="220" height="73" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/37/Final_Primary_Differentiation.svg/330px-Final_Primary_Differentiation.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/37/Final_Primary_Differentiation.svg/440px-Final_Primary_Differentiation.svg.png 2x" data-file-width="2000" data-file-height="663" /></a><figcaption>Primary differentiation is demonstrated using the peppered moth as an example, with a change in an environmental variable such as sooty coverage of trees imposing a selective pressure on a previously uniformly coloured moth population. This causes the frequency of melanic morphs to increase the more soot there is on vegetation.</figcaption></figure> <p>Clines produced through this way are generated by spatial heterogeneity in environmental conditions. The mechanism of selection acting upon organisms is therefore external. Species ranges frequently span environmental gradients (e.g. humidity, rainfall, temperature, or day length) and, according to natural selection, different environments will favour different <a href="/wiki/Genotype" title="Genotype">genotypes</a> or <a href="/wiki/Phenotype" title="Phenotype">phenotypes</a>.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> In this way, when previously genetically or phenotypically uniform populations spread into novel environments, they will evolve to be uniquely adapted to the local environment, in the process potentially creating a gradient in a genotypic or phenotypic trait. </p><p>Such clines in characters can not be maintained through selection alone if much gene flow occurred between populations, as this would tend to swamp out the effects of local adaptation. However, because species usually tend to have a limited dispersal range (e.g. in an <a href="/wiki/Isolation_by_distance" title="Isolation by distance">isolation by distance</a> model), restricted gene flow can serve as a type of barrier which encourages geographic differentiation.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> However, some degree of migration is often required to maintain a cline; without it, <a href="/wiki/Speciation" title="Speciation">speciation</a> is likely to eventually occur, as local adaptation can cause reproductive isolation between populations.<sup id="cite_ref-Endler_2-2" class="reference"><a href="#cite_note-Endler-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p><p>A classic example of the role of environmental gradients in creating clines is that of the peppered moth, <a href="/wiki/Peppered_moth" title="Peppered moth"><i>Biston betularia</i></a>, in the UK. During the 19th century, when the industrial sector gained traction, coal emissions blackened vegetation across northwest England and parts of northern Wales. As a result of this, lighter <a href="/wiki/Morph_(zoology)" class="mw-redirect" title="Morph (zoology)">morphs</a> of the moth were more visible to predators against the blackened tree trunks and were therefore more heavily predated relative to the darker morphs. Consequently, the frequency of the more <a href="/wiki/Crypsis" title="Crypsis">cryptic</a> melanic morph of the peppered moth increased drastically in northern England. This cline in morph colour, from a dominance of lighter morphs in the west of England (which did not suffer as heavily from pollution), to the higher frequency of melanic forms in the north, has slowly been degrading since limitations to sooty emissions were introduced in the 1960s.<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> </p> <div class="mw-heading mw-heading4"><h4 id="Secondary_contact">Secondary contact</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cline_(biology)&action=edit&section=4" title="Edit section: Secondary contact"><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_contact" title="Secondary contact">Secondary contact</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Secondary_Introgression.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/01/Secondary_Introgression.svg/220px-Secondary_Introgression.svg.png" decoding="async" width="220" height="73" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/01/Secondary_Introgression.svg/330px-Secondary_Introgression.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/01/Secondary_Introgression.svg/440px-Secondary_Introgression.svg.png 2x" data-file-width="2000" data-file-height="663" /></a><figcaption>Secondary contact between two previously isolated populations. Two previously isolated populations establish contact and therefore gene flow, creating an intermediate zone in the phenotypic or genotypic character between the two populations.</figcaption></figure> <p>Clines generated through this mechanism have arisen through the joining of two formerly isolated populations which differentiated in <a href="/wiki/Allopatric_speciation" title="Allopatric speciation">allopatry</a>, creating an intermediate zone. This secondary contact scenario may occur, for example, when climatic conditions change, allowing the ranges of populations to expand and meet.<sup id="cite_ref-Mayr1942_6-1" class="reference"><a href="#cite_note-Mayr1942-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> Because over time the effect of gene flow will tend to eventually swamp out any regional differences and cause one large homogenous population, for a stable cline to be maintained when two populations join there must usually be a selective pressure maintaining a degree of differentiation between the two populations.<sup id="cite_ref-Endler_2-3" class="reference"><a href="#cite_note-Endler-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p><p>The mechanism of selection maintaining the clines in this scenario is often intrinsic. This means that the fitness of individuals is independent of the external environment, and selection is instead dependent on the genome of the individual. Intrinsic, or endogenous, selection can give rise to clines in characters through a variety of mechanisms. One way it may act is through <a href="/wiki/Underdominance" title="Underdominance">heterozygote disadvantage</a>, in which intermediate genotypes have a lower relative fitness than either homozygote genotypes. Because of this disadvantage, one allele will tend to become fixed in a given population, such that populations will consist largely of either <i>AA</i> (<a href="/wiki/Homozygote" class="mw-redirect" title="Homozygote">homozygous</a> dominant) or <i>aa</i> (homozygous recessive) individuals.<sup id="cite_ref-Bazykin1969_11-0" class="reference"><a href="#cite_note-Bazykin1969-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> The cline of hetero<a href="/wiki/Zygote" title="Zygote">zygotes</a> that is created when these respective populations come into contact is then shaped by the opposing forces of selection and gene flow; even if selection against heterozygotes is great, if there is some degree of gene flow between the two populations, then a steep cline may be able to be maintained.<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Richard1993_13-0" class="reference"><a href="#cite_note-Richard1993-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> Because instrinsic selection is independent of the external environment, clines generated by selection against hybrids are not fixed to any given geographical area and can move around the geographic landscape.<sup id="cite_ref-EoE_14-0" class="reference"><a href="#cite_note-EoE-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> Such <a href="/wiki/Hybrid_zone" title="Hybrid zone">hybrid zones</a> where hybrids are a disadvantage relative to their parental lines (but which are nonetheless maintained through selection being counteracted by gene flow) are known as "tension zones".<sup id="cite_ref-Bazykin1969_11-1" class="reference"><a href="#cite_note-Bazykin1969-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p><p>Another way in which selection can generate clines is through <a href="/wiki/Frequency-dependent_selection" title="Frequency-dependent selection">frequency-dependent selection</a>. Characters that could be maintained by such frequency-dependent selective pressures include warning signals (<a href="/wiki/Aposematism" title="Aposematism">aposematism</a>). For example, aposematic signals in <i><a href="/wiki/Heliconius" title="Heliconius">Heliconius</a></i> butterflies sometimes display steep clines between populations, which are maintained through <a href="/wiki/Frequency-dependent_selection#Positive" title="Frequency-dependent selection">positive frequency dependence</a>.<sup id="cite_ref-Richard1993_13-1" class="reference"><a href="#cite_note-Richard1993-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> This is because heterozygosity, <a href="/wiki/Mutation" title="Mutation">mutations</a> and <a href="/wiki/Genetic_recombination" title="Genetic recombination">recombination</a> can all produce patterns that deviate from those well-established signals which mark prey as being unpalatable. These individuals are then predated more heavily relative to their counterparts with "normal" markings (i.e. selected against), creating populations dominated by a particular pattern of warning signal. As with heterozygote disadvantage, when these populations join, a narrow cline of intermediate individuals could be produced, maintained by gene flow counteracting selection.<sup id="cite_ref-Richard1993_13-2" class="reference"><a href="#cite_note-Richard1993-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><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> </p><p>Secondary contact could lead to a cline with a steep gradient if heterozygote disadvantage or frequency-dependent selection exists, as intermediates are heavily selected against. Alternatively, steep clines could exist because the populations have only recently established secondary contact, and the character in the original allopatric populations had a large degree of differentiation. As genetic admixture between the population increases with time however, the steepness of the cline is likely to decrease as the difference in character is eroded. However, if the character in the original allopatric populations was not very differentiated to begin with, the cline between the populations need not display a very steep gradient.<sup id="cite_ref-Endler_2-4" class="reference"><a href="#cite_note-Endler-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Because both primary differentiation and secondary contact can therefore give rise to similar or identical clinal patterns (e.g. gently sloping clines), distinguishing which of these two processes is responsible for generating a cline is difficult and often impossible.<sup id="cite_ref-Endler_2-5" class="reference"><a href="#cite_note-Endler-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> However, in some circumstances a cline and a geographic variable (such as humidity) may be very tightly linked, with a change in one corresponding closely to a change in the other. In such cases it may be tentatively concluded that the cline is generated by primary differentiation and therefore moulded by environmental selective pressures.<sup id="cite_ref-Joseph_7-1" class="reference"><a href="#cite_note-Joseph-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="No_selection_(drift/migration_balance)"><span id="No_selection_.28drift.2Fmigration_balance.29"></span>No selection (drift/migration balance)</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cline_(biology)&action=edit&section=5" title="Edit section: No selection (drift/migration balance)"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>While selection can therefore clearly play a key role in creating clines, it is theoretically feasible that they might be generated by <a href="/wiki/Genetic_drift" title="Genetic drift">genetic drift</a> alone. It is unlikely that large-scale clines in genotype or phenotype frequency will be produced solely by drift. However, across smaller geographical scales and in smaller populations, drift could produce temporary clines.<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> The fact that drift is a weak force upholding the cline however means that clines produced this way are often random (i.e. uncorrelated with environmental variables) and subject to breakdown or reversal over time.<sup id="cite_ref-Endler_2-6" class="reference"><a href="#cite_note-Endler-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Such clines are therefore unstable and sometimes called "transient clines".<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Clinal_structure_and_terminology">Clinal structure and terminology</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cline_(biology)&action=edit&section=6" title="Edit section: Clinal structure and terminology"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Character-Cline_Diagram.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b6/Character-Cline_Diagram.svg/220px-Character-Cline_Diagram.svg.png" decoding="async" width="220" height="90" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b6/Character-Cline_Diagram.svg/330px-Character-Cline_Diagram.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b6/Character-Cline_Diagram.svg/440px-Character-Cline_Diagram.svg.png 2x" data-file-width="1652" data-file-height="678" /></a><figcaption>Clinal characters change from one end of the geographic range to another. The extent of this change is reflected in the slope of the cline.</figcaption></figure> <p>The steepness, or gradient, of a cline reflects the extent of the differentiation in the character across a geographic range.<sup id="cite_ref-Endler_2-7" class="reference"><a href="#cite_note-Endler-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> For example, a steep cline could indicate large variation in the colour of plumage between adjacent bird populations. It has been previously outlined that such steep clines may be the result of two previously allopatric populations with a large degree of difference in the trait having only recently established gene flow, or where there is strong selection against hybrids. However, it may also reflect a sudden environmental change or boundary. Examples of rapidly changing environmental boundaries like this include abrupt changes in the heavy metal content of soils, and the consequent narrow clines produced between populations of <i><a href="/wiki/Agrostis" title="Agrostis">Agrostis</a></i> that are either adapted to these soils with high metal content, or adapted to "normal" soil.<sup id="cite_ref-Endler_2-8" class="reference"><a href="#cite_note-Endler-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> Conversely, a shallow cline indicates little geographical variation in the character or trait across a given geographical distance. This may have arisen through weak differential environmental selective pressure, or where two populations established secondary contact a long time ago and gene flow has eroded the large character differentiation between the populations. </p><p>The gradient of a cline is related to another commonly referred to property, clinal width. A cline with a steep slope is said to have a small, or narrow, width, while shallower clines have larger widths.<sup id="cite_ref-Joseph_7-2" class="reference"><a href="#cite_note-Joseph-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Types_of_clines">Types of clines</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cline_(biology)&action=edit&section=7" title="Edit section: Types of clines"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Clinal_Variation_by_Huxley.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b6/Clinal_Variation_by_Huxley.svg/220px-Clinal_Variation_by_Huxley.svg.png" decoding="async" width="220" height="140" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b6/Clinal_Variation_by_Huxley.svg/330px-Clinal_Variation_by_Huxley.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b6/Clinal_Variation_by_Huxley.svg/440px-Clinal_Variation_by_Huxley.svg.png 2x" data-file-width="2644" data-file-height="1684" /></a><figcaption>Categories and subcategories of clines, as defined by Huxley</figcaption></figure> <p>According to Huxley, clines can be classified into two categories; continuous clines and discontinuous stepped clines.<sup id="cite_ref-Huxley_1-2" class="reference"><a href="#cite_note-Huxley-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> These types of clines characterise the way that a genetic or phenotypic trait transforms from one end of its geographical range of the species to the other. </p> <div class="mw-heading mw-heading3"><h3 id="Continuous_clines">Continuous clines</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cline_(biology)&action=edit&section=8" title="Edit section: Continuous clines"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In continuous clines, all populations of the species are able to interbreed and there is gene flow throughout the entire range of the species. In this way, these clines are both biologically (no clear subgroups) and geographically (contiguous distribution) continuous. Continuous clines can be further sub-divided into smooth and stepped clines. </p> <ul><li>Continuous smooth clines are characterised by the lack of any abrupt changes or delineation in the genetic or phenotypic trait across the cline, instead displaying a smooth gradation throughout. Huxley recognised that this type of cline, with its uniform slope throughout, was unlikely to be common.</li> <li>Continuous stepped clines consist of an overall shallow cline, interspersed by sections of much steeper slope. The shallow slope represents the populations, and the shorter, steeper sections the larger change in character between populations.<sup id="cite_ref-hedrick_19-0" class="reference"><a href="#cite_note-hedrick-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> Stepped clines can be further subdivided into horizontally stepped clines, and obliquely stepped clines. <ul><li>Horizontally stepped clines show no intra-population variation or gradation in the character, therefore displaying a horizontal gradient. These uniform populations are connected by steeper sections of the cline, characterised by larger changes in the form of the character. However, because in continuous clines all populations exchange genetic material, the intergradation zone between the groups can never have a vertical slope.<sup id="cite_ref-hedrick_19-1" class="reference"><a href="#cite_note-hedrick-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup></li> <li>In obliquely stepped clines, conversely, each population also demonstrates a cline in the character, albeit of a shallower slope than the clines connecting the populations together. Huxley compared obliquely stepped clines to looking like a "stepped ramp", rather than taking on the formation of a staircase as in the case of horizontally stepped clines.<sup id="cite_ref-Huxley_biography_3-1" class="reference"><a href="#cite_note-Huxley_biography-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup></li></ul></li></ul> <div class="mw-heading mw-heading3"><h3 id="Discontinuous_stepped_clines">Discontinuous stepped clines</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cline_(biology)&action=edit&section=9" title="Edit section: Discontinuous stepped clines"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Unlike in continuous clines, in discontinuous clines the populations of species are allopatric, meaning there is very little or no gene flow amongst populations. The genetic or phenotypic trait in question always shows a steeper gradient between groups than within groups, as in continuous clines. Discontinuous clines follow the same principles as continuous clines by displaying either </p> <ul><li>Horizontally stepped clines, where intra-group variation is very small or non-existent and the geographic space separating groups shows a sharp change in character</li> <li>Obliquely stepped clines, where there is some intra-group gradation, but this is less than the gradation in the character between populations<sup id="cite_ref-Huxley_1-3" class="reference"><a href="#cite_note-Huxley-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup></li></ul> <div class="mw-heading mw-heading2"><h2 id="Clines_and_speciation">Clines and speciation</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cline_(biology)&action=edit&section=10" title="Edit section: Clines and speciation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Parapatric_Speciation_Schematic.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/5f/Parapatric_Speciation_Schematic.svg/220px-Parapatric_Speciation_Schematic.svg.png" decoding="async" width="220" height="89" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/5f/Parapatric_Speciation_Schematic.svg/330px-Parapatric_Speciation_Schematic.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/5f/Parapatric_Speciation_Schematic.svg/440px-Parapatric_Speciation_Schematic.svg.png 2x" data-file-width="1683" data-file-height="683" /></a><figcaption><a href="/wiki/Parapatric_speciation" title="Parapatric speciation">Parapatric speciation</a></figcaption></figure> <p>It was originally assumed that geographic isolation was a necessary precursor to speciation (<a href="/wiki/Allopatric_speciation" title="Allopatric speciation">allopatric speciation</a>).<sup id="cite_ref-Mayr1942_6-2" class="reference"><a href="#cite_note-Mayr1942-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> The possibility that clines may be a precursor to speciation was therefore ignored, as they were assumed to be evidence of the fact that in contiguous populations gene flow was too strong a force of homogenisation, and selection too weak a force of differentiation, for speciation to take place.<sup id="cite_ref-Endler_2-9" class="reference"><a href="#cite_note-Endler-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> However, the existence of particular types of clines, such as <a href="/wiki/Ring_species" title="Ring species">ring species</a>, in which populations did not differentiate in allopatry but the terminal ends of the cline nonetheless do not interbreed, cast into doubt whether complete geographical isolation of populations is an absolute requirement for speciation. </p><p>Because clines can exist in populations connected by some degree of gene flow, the generation of new species from a previously clinal population is termed <a href="/wiki/Parapatric_speciation" title="Parapatric speciation">parapatric speciation</a>. Both extrinsic and intrinsic selection can serve to generate varying degrees of <a href="/wiki/Reproductive_isolation" title="Reproductive isolation">reproductive isolation</a> and thereby instigate the process of <a href="/wiki/Speciation" title="Speciation">speciation</a>. For example, through environmental selection acting on populations and favouring particular allele frequencies, large genetic differences between populations may accumulate (this would be reflected in clinal structure by the presence of numerous very steep clines). If the local genetic differences are great enough, it may lead unfavourable combinations of genotypes and therefore to hybrids being at a decreased fitness relative to the parental lines. When this hybrid disadvantage is great enough, natural selection will select for <a href="/wiki/Reproductive_isolation#Pre-zygotic_isolation" title="Reproductive isolation">pre-zygotic</a> traits in the homozygous parental lines that reduce the likelihood of disadvantageous hybridisation - in other words, natural selection will favour traits that promote assortative mating in the parental lines.<sup id="cite_ref-EoE_14-1" class="reference"><a href="#cite_note-EoE-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> This is known as <a href="/wiki/Reinforcement_(speciation)" title="Reinforcement (speciation)">reinforcement</a> and plays an important role in parapatric and <a href="/wiki/Sympatric_speciation" title="Sympatric speciation">sympatric speciation</a>.<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Endler_2-10" class="reference"><a href="#cite_note-Endler-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Clinal_maps">Clinal maps</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cline_(biology)&action=edit&section=11" title="Edit section: Clinal maps"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Clines can be portrayed graphically on maps using lines that show the transition in character state from one end of the geographic range to the other. Character states can however additionally be represented using isophenes, defined by Ernst Mayr as "lines of equal expression of a clinally varying character".<sup id="cite_ref-Mayr1963_5-3" class="reference"><a href="#cite_note-Mayr1963-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> In other words, areas on maps that demonstrate the same biological phenomenon or character will be connected by something that resembles a contour line. When mapping clines therefore, which follow a character gradation from one extreme to the other, isophenes will transect clinal lines at a right angle. </p> <div class="mw-heading mw-heading2"><h2 id="Examples_of_clines">Examples of clines</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cline_(biology)&action=edit&section=12" title="Edit section: Examples of clines"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Northern_red_fox_%26_southern_desert_red_fox.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f9/Northern_red_fox_%26_southern_desert_red_fox.jpg/220px-Northern_red_fox_%26_southern_desert_red_fox.jpg" decoding="async" width="220" height="71" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f9/Northern_red_fox_%26_southern_desert_red_fox.jpg/330px-Northern_red_fox_%26_southern_desert_red_fox.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f9/Northern_red_fox_%26_southern_desert_red_fox.jpg/440px-Northern_red_fox_%26_southern_desert_red_fox.jpg 2x" data-file-width="1040" data-file-height="336" /></a><figcaption>Bergmann's Rule demonstrated showing the difference in size between a larger northern fox, whose range spans colder regions, and a smaller desert fox, whose range is primarily in hot regions</figcaption></figure> <p>Although the term "cline" was first officially coined by Huxley in 1938, gradients and geographic variations in the character states of species have been observed for centuries. Indeed, some gradations have been considered so ubiquitous that they have been labelled <a href="/wiki/Biological_rules" title="Biological rules">ecological "rules"</a>. One commonly cited example of a gradient in morphology is <a href="/wiki/Gloger%27s_rule" title="Gloger's rule">Gloger's Rule</a>, named after <a href="/wiki/C._L._Gloger" title="C. L. Gloger">Constantin Gloger</a>, who observed in 1833 that environmental factors and the pigmentation of avian <a href="/wiki/Plumage" title="Plumage">plumage</a> tend to covary with each other, such that birds found in arid areas near the Equator tend to be much darker than those in less arid areas closer to the Poles. Since then, this rule has been extended to include many other animals, including flies, butterflies, and wolves.<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><p>Other ecogeographical rules include <a href="/wiki/Bergmann%E2%80%99s_rule" class="mw-redirect" title="Bergmann’s rule">Bergmann's Rule</a>, coined by <a href="/wiki/Carl_Bergmann_(anatomist)" title="Carl Bergmann (anatomist)">Carl Bergmann</a> in 1857, which states that <a href="/wiki/Homeothermy" title="Homeothermy">homeotherms</a> closer to the Equator tend to be smaller than their more northerly or southerly conspecifics.<sup id="cite_ref-Wool_22-0" class="reference"><a href="#cite_note-Wool-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> One of the proposed reasons for this cline is that larger animals have a relatively smaller surface area to volume ratio and therefore improved heat conservancy – an important advantage in cold climates.<sup id="cite_ref-Wool_22-1" class="reference"><a href="#cite_note-Wool-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> The role of the environment in imposing a selective pressure and producing this cline has been heavily implicated due to the fact that Bergmann's Rule has been observed across many independent lineages of species and continents. For example, the <a href="/wiki/House_sparrow" title="House sparrow">house sparrow</a>, which was introduced in the early 1850s to the eastern United States, evolved a north-south gradient in size soon after its introduction. This gradient reflects the gradient that already existed in the house sparrow's native range in Europe.<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Ring_Species_(gene_flow_around_a_barrier)_BLANK.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/12/Ring_Species_%28gene_flow_around_a_barrier%29_BLANK.png/220px-Ring_Species_%28gene_flow_around_a_barrier%29_BLANK.png" decoding="async" width="220" height="147" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/12/Ring_Species_%28gene_flow_around_a_barrier%29_BLANK.png/330px-Ring_Species_%28gene_flow_around_a_barrier%29_BLANK.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/12/Ring_Species_%28gene_flow_around_a_barrier%29_BLANK.png/440px-Ring_Species_%28gene_flow_around_a_barrier%29_BLANK.png 2x" data-file-width="3493" data-file-height="2340" /></a><figcaption>Interbreeding populations represented by a gradient of coloured circles around a geographic barrier</figcaption></figure> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Ring_species_seagull.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8d/Ring_species_seagull.svg/220px-Ring_species_seagull.svg.png" decoding="async" width="220" height="248" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8d/Ring_species_seagull.svg/330px-Ring_species_seagull.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8d/Ring_species_seagull.svg/440px-Ring_species_seagull.svg.png 2x" data-file-width="552" data-file-height="622" /></a><figcaption>The <i>Larus</i> gulls interbreed in a ring around the arctic.<br />1: <i>Larus argentatus argentatus</i>, 2: <i>Larus fuscus</i> (<i>sensu stricto</i>), 3: <i>Larus fuscus heuglini</i>, 4: <i>Larus argentatus birulai</i>, 5: <i>Larus argentatus vegae</i>, 6: <i>Larus argentatus smithsonianus</i>, 7: <i>Larus argentatus argenteus</i></figcaption></figure> <p><a href="/wiki/Ring_species" title="Ring species">Ring species</a><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> are a distinct type of cline where the geographical distribution in question is circular in shape, so that the two ends of the cline overlap with one another, giving two adjacent populations that rarely <a href="/wiki/Hybrid_(biology)" title="Hybrid (biology)">interbreed</a> due to the cumulative effect of the many changes in phenotype along the cline. The populations elsewhere along the cline interbreed with their geographically adjacent populations as in a standard cline. In the case of <i><a href="/wiki/Larus" title="Larus">Larus</a></i> gulls, the habitats of the end populations even overlap, which introduces questions as to what constitutes a species: nowhere along the cline can a line be drawn between the populations, but they are unable to interbreed. </p><p>In humans, clines in the frequency of blood types has allowed scientists to infer past population migrations. For example, the Type B blood group reaches its highest frequency in Asia, but become less frequent further west. From this, it has been possible to infer that some Asian populations migrated towards Europe around 2,000 years ago, causing genetic admixture in an <a href="/wiki/Isolation_by_distance" title="Isolation by distance">isolation by distance</a> model. In contrast to this cline, blood Type A shows the reverse pattern, reaching its highest frequency in Europe and declining in frequency towards Asia.<sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cline_(biology)&action=edit&section=13" 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-Huxley-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-Huxley_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Huxley_1-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Huxley_1-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Huxley_1-3"><sup><i><b>d</b></i></sup></a></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFHuxley1938" class="citation journal cs1">Huxley, Julian (30 July 1938). "Clines: an Auxiliary Taxonomic Principle". <i>Nature</i>. <b>142</b> (3587): 219–220. <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/1938Natur.142..219H">1938Natur.142..219H</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2F142219a0">10.1038/142219a0</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:4124055">4124055</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature&rft.atitle=Clines%3A+an+Auxiliary+Taxonomic+Principle&rft.volume=142&rft.issue=3587&rft.pages=219-220&rft.date=1938-07-30&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A4124055%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1038%2F142219a0&rft_id=info%3Abibcode%2F1938Natur.142..219H&rft.aulast=Huxley&rft.aufirst=Julian&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACline+%28biology%29" class="Z3988"></span></span> </li> <li id="cite_note-Endler-2"><span class="mw-cite-backlink">^ <a href="#cite_ref-Endler_2-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Endler_2-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Endler_2-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Endler_2-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-Endler_2-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-Endler_2-5"><sup><i><b>f</b></i></sup></a> <a href="#cite_ref-Endler_2-6"><sup><i><b>g</b></i></sup></a> <a href="#cite_ref-Endler_2-7"><sup><i><b>h</b></i></sup></a> <a href="#cite_ref-Endler_2-8"><sup><i><b>i</b></i></sup></a> <a href="#cite_ref-Endler_2-9"><sup><i><b>j</b></i></sup></a> <a href="#cite_ref-Endler_2-10"><sup><i><b>k</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFEndler1977" class="citation book cs1">Endler, John A. (1977). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=BL12fYUMobwC&q=rassenkreis+cline"><i>Geographic Variation, Speciation, and Clines</i></a>. Vol. 10. Princeton University Press. pp. 1–246. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0691081922" title="Special:BookSources/978-0691081922"><bdi>978-0691081922</bdi></a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/409931">409931</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Geographic+Variation%2C+Speciation%2C+and+Clines&rft.pages=1-246&rft.pub=Princeton+University+Press&rft.date=1977&rft_id=info%3Apmid%2F409931&rft.isbn=978-0691081922&rft.aulast=Endler&rft.aufirst=John+A.&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DBL12fYUMobwC%26q%3Drassenkreis%2Bcline&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACline+%28biology%29" class="Z3988"></span> <span class="cs1-visible-error citation-comment"><code class="cs1-code">{{<a href="/wiki/Template:Cite_book" title="Template:Cite book">cite book</a>}}</code>: </span><span class="cs1-visible-error citation-comment"><code class="cs1-code">|journal=</code> ignored (<a href="/wiki/Help:CS1_errors#periodical_ignored" title="Help:CS1 errors">help</a>)</span></span> </li> <li id="cite_note-Huxley_biography-3"><span class="mw-cite-backlink">^ <a href="#cite_ref-Huxley_biography_3-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Huxley_biography_3-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFDronamrajuNeedham1993" class="citation book cs1">Dronamraju, Krishna R.; Needham, Joseph (1993). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=xrLsCgAAQBAJ&q=continuous+discontinuous+clines&pg=PT195"><i>If I Am To Be Remembered: Correspondence Of Julian Huxley</i></a>. World Scientific. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/9789814505192" title="Special:BookSources/9789814505192"><bdi>9789814505192</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=If+I+Am+To+Be+Remembered%3A+Correspondence+Of+Julian+Huxley&rft.pub=World+Scientific&rft.date=1993&rft.isbn=9789814505192&rft.aulast=Dronamraju&rft.aufirst=Krishna+R.&rft.au=Needham%2C+Joseph&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DxrLsCgAAQBAJ%26q%3Dcontinuous%2Bdiscontinuous%2Bclines%26pg%3DPT195&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACline+%28biology%29" class="Z3988"></span></span> </li> <li id="cite_note-Forest_genetics-4"><span class="mw-cite-backlink">^ <a href="#cite_ref-Forest_genetics_4-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Forest_genetics_4-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWhiteAdamsNeale2007" class="citation book cs1">White, Timothy L.; Adams, W. T.; Neale, David B. (2007). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=UHZCeg4BqtkC&q=ecotypes+and+clines&pg=PA190"><i>Forest Genetics</i></a>. CABI Publishing. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/9781845932862" title="Special:BookSources/9781845932862"><bdi>9781845932862</bdi></a><span class="reference-accessdate">. Retrieved <span class="nowrap">28 March</span> 2018</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Forest+Genetics&rft.pub=CABI+Publishing&rft.date=2007&rft.isbn=9781845932862&rft.aulast=White&rft.aufirst=Timothy+L.&rft.au=Adams%2C+W.+T.&rft.au=Neale%2C+David+B.&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DUHZCeg4BqtkC%26q%3Decotypes%2Band%2Bclines%26pg%3DPA190&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACline+%28biology%29" class="Z3988"></span></span> </li> <li id="cite_note-Mayr1963-5"><span class="mw-cite-backlink">^ <a href="#cite_ref-Mayr1963_5-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Mayr1963_5-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Mayr1963_5-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Mayr1963_5-3"><sup><i><b>d</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMayr1963" class="citation book cs1">Mayr, Ernst (1963). <span class="id-lock-registration" title="Free registration required"><a rel="nofollow" class="external text" href="https://archive.org/details/animalspeciesevo00mayr"><i>Populations, species, and evolution: an abridgment of Animal species and evolution</i></a></span>. Belknap Press of Harvard University Press<span class="reference-accessdate">. Retrieved <span class="nowrap">1 April</span> 2018</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Populations%2C+species%2C+and+evolution%3A+an+abridgment+of+Animal+species+and+evolution&rft.pub=Belknap+Press+of+Harvard+University+Press&rft.date=1963&rft.aulast=Mayr&rft.aufirst=Ernst&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fanimalspeciesevo00mayr&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACline+%28biology%29" class="Z3988"></span></span> </li> <li id="cite_note-Mayr1942-6"><span class="mw-cite-backlink">^ <a href="#cite_ref-Mayr1942_6-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Mayr1942_6-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Mayr1942_6-2"><sup><i><b>c</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMayr1942" class="citation book cs1">Mayr, Ernst (1942). <span class="id-lock-registration" title="Free registration required"><a rel="nofollow" class="external text" href="https://archive.org/details/systematicsorigi0000mayr_p8z7"><i>Systematics and the Origin of Species from the viewpoint of a zoologist</i></a></span>. Columbia University Press.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Systematics+and+the+Origin+of+Species+from+the+viewpoint+of+a+zoologist&rft.pub=Columbia+University+Press&rft.date=1942&rft.aulast=Mayr&rft.aufirst=Ernst&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fsystematicsorigi0000mayr_p8z7&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACline+%28biology%29" class="Z3988"></span></span> </li> <li id="cite_note-Joseph-7"><span class="mw-cite-backlink">^ <a href="#cite_ref-Joseph_7-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Joseph_7-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Joseph_7-2"><sup><i><b>c</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFFelsenstein2013" class="citation book cs1">Felsenstein, Joseph (2013). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=BW7tCAAAQBAJ&q=cline+slope+steepness"><i>Numerical Taxonomy</i></a>. Springer Science & Business Media. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/9783642690242" title="Special:BookSources/9783642690242"><bdi>9783642690242</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Numerical+Taxonomy&rft.pub=Springer+Science+%26+Business+Media&rft.date=2013&rft.isbn=9783642690242&rft.aulast=Felsenstein&rft.aufirst=Joseph&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DBW7tCAAAQBAJ%26q%3Dcline%2Bslope%2Bsteepness&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACline+%28biology%29" class="Z3988"></span></span> </li> <li id="cite_note-8"><span class="mw-cite-backlink"><b><a href="#cite_ref-8">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFDarwin1859" class="citation book cs1">Darwin, Charles R. 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Jones & Bartlett Learning.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Genetics+of+Populations&rft.pub=Jones+%26+Bartlett+Learning&rft.date=2011&rft.aulast=Hedrick&rft.aufirst=Philip&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACline+%28biology%29" class="Z3988"></span></span> </li> <li id="cite_note-20"><span class="mw-cite-backlink"><b><a href="#cite_ref-20">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCoyneOrr2004" class="citation book cs1">Coyne, Jerry A.; Orr, H. Allen (2004). <i>Speciation</i>. Sinauer. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/9780878930890" title="Special:BookSources/9780878930890"><bdi>9780878930890</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Speciation&rft.pub=Sinauer&rft.date=2004&rft.isbn=9780878930890&rft.aulast=Coyne&rft.aufirst=Jerry+A.&rft.au=Orr%2C+H.+Allen&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACline+%28biology%29" class="Z3988"></span></span> </li> <li id="cite_note-21"><span class="mw-cite-backlink"><b><a href="#cite_ref-21">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHuggett2004" class="citation book cs1">Huggett, Richard J. (2004). <i>Fundamentals of Biogeography</i>. Psychology Press.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Fundamentals+of+Biogeography&rft.pub=Psychology+Press&rft.date=2004&rft.aulast=Huggett&rft.aufirst=Richard+J.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACline+%28biology%29" class="Z3988"></span></span> </li> <li id="cite_note-Wool-22"><span class="mw-cite-backlink">^ <a href="#cite_ref-Wool_22-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Wool_22-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWool2006" class="citation book cs1">Wool, David (2006). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=ydWuuBw6xeYC&q=huxley+1942+cline"><i>The Driving Forces of Evolution: Genetic Processes in Populations</i></a>. CRC Press. p. 163. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/9781578084456" title="Special:BookSources/9781578084456"><bdi>9781578084456</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Driving+Forces+of+Evolution%3A+Genetic+Processes+in+Populations&rft.pages=163&rft.pub=CRC+Press&rft.date=2006&rft.isbn=9781578084456&rft.aulast=Wool&rft.aufirst=David&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DydWuuBw6xeYC%26q%3Dhuxley%2B1942%2Bcline&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACline+%28biology%29" class="Z3988"></span></span> </li> <li id="cite_note-23"><span class="mw-cite-backlink"><b><a href="#cite_ref-23">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHuggett1995" class="citation book cs1">Huggett, Richard J. (1995). <span class="id-lock-registration" title="Free registration required"><a rel="nofollow" class="external text" href="https://archive.org/details/geoecologyevolut0000hugg"><i>Geoecology: An evolutionary approach</i></a></span>. Routledge. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/9780415086899" title="Special:BookSources/9780415086899"><bdi>9780415086899</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Geoecology%3A+An+evolutionary+approach&rft.pub=Routledge&rft.date=1995&rft.isbn=9780415086899&rft.aulast=Huggett&rft.aufirst=Richard+J.&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fgeoecologyevolut0000hugg&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACline+%28biology%29" class="Z3988"></span></span> </li> <li id="cite_note-24"><span class="mw-cite-backlink"><b><a href="#cite_ref-24">^</a></b></span> <span class="reference-text"><i>Fundamentals of biogeography</i>, Richard J. Huggett, 2nd ed. (2004), p. 20</span> </li> <li id="cite_note-25"><span class="mw-cite-backlink"><b><a href="#cite_ref-25">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBirx2006" class="citation book cs1">Birx, H. James (2006). <i>Encyclopaedia of Anthropology</i>. SAGE.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Encyclopaedia+of+Anthropology&rft.pub=SAGE&rft.date=2006&rft.aulast=Birx&rft.aufirst=H.+James&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACline+%28biology%29" class="Z3988"></span></span> </li> </ol></div></div> <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 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.navbox-abovebelow,.mw-parser-output .navbox-group,.mw-parser-output .navbox-subgroup .navbox-title{background-color:#ddf}.mw-parser-output .navbox-subgroup .navbox-group,.mw-parser-output .navbox-subgroup .navbox-abovebelow{background-color:#e6e6ff}.mw-parser-output .navbox-even{background-color:#f7f7f7}.mw-parser-output .navbox-odd{background-color:transparent}.mw-parser-output .navbox .hlist td dl,.mw-parser-output .navbox .hlist td ol,.mw-parser-output .navbox .hlist td ul,.mw-parser-output .navbox td.hlist dl,.mw-parser-output .navbox td.hlist ol,.mw-parser-output .navbox td.hlist ul{padding:0.125em 0}.mw-parser-output .navbox .navbar{display:block;font-size:100%}.mw-parser-output .navbox-title .navbar{float:left;text-align:left;margin-right:0.5em}body.skin--responsive .mw-parser-output .navbox-image img{max-width:none!important}@media print{body.ns-0 .mw-parser-output .navbox{display:none!important}}</style></div><div role="navigation" class="navbox" aria-labelledby="Speciation" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><style data-mw-deduplicate="TemplateStyles:r1239400231">.mw-parser-output .navbar{display:inline;font-size:88%;font-weight:normal}.mw-parser-output .navbar-collapse{float:left;text-align:left}.mw-parser-output .navbar-boxtext{word-spacing:0}.mw-parser-output .navbar ul{display:inline-block;white-space:nowrap;line-height:inherit}.mw-parser-output .navbar-brackets::before{margin-right:-0.125em;content:"[ "}.mw-parser-output .navbar-brackets::after{margin-left:-0.125em;content:" ]"}.mw-parser-output .navbar li{word-spacing:-0.125em}.mw-parser-output .navbar a>span,.mw-parser-output .navbar a>abbr{text-decoration:inherit}.mw-parser-output .navbar-mini abbr{font-variant:small-caps;border-bottom:none;text-decoration:none;cursor:inherit}.mw-parser-output .navbar-ct-full{font-size:114%;margin:0 7em}.mw-parser-output .navbar-ct-mini{font-size:114%;margin:0 4em}html.skin-theme-clientpref-night .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Speciation" title="Template:Speciation"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Speciation" title="Template talk:Speciation"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Speciation" title="Special:EditPage/Template:Speciation"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Speciation" style="font-size:114%;margin:0 4em"><a href="/wiki/Speciation" title="Speciation">Speciation</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><a href="/w/index.php?title=Introduction_to_speciation&action=edit&redlink=1" class="new" title="Introduction to speciation (page does not exist)">Introduction</a></li> <li><a href="/wiki/History_of_speciation" title="History of speciation">History</a></li> <li><a href="/wiki/Laboratory_experiments_of_speciation" title="Laboratory experiments of speciation">Laboratory experiments</a></li> <li><a href="/wiki/Glossary_of_speciation" class="mw-redirect" title="Glossary of speciation">Glossary</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Basic concepts</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Species" title="Species">Species</a> (<a href="/wiki/Species_complex" title="Species complex">Species complex</a>)</li> <li><a href="/wiki/Reproductive_isolation" title="Reproductive isolation">Reproductive isolation</a></li> <li><a href="/wiki/Anagenesis" title="Anagenesis">Anagenesis</a></li> <li><a href="/wiki/Cladogenesis" title="Cladogenesis">Cladogenesis</a></li> <li><a href="/wiki/Cospeciation" title="Cospeciation">Cospeciation</a></li> <li><a href="/wiki/Evidence_of_evolution" class="mw-redirect" title="Evidence of evolution">Evidence of evolution</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Geographic modes</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Allopatric_speciation" title="Allopatric speciation">Allopatric</a> (<a href="/wiki/Peripatric_speciation" title="Peripatric speciation">Peripatric</a> · <a href="/wiki/Peripatric_speciation#Quantum_and_budding_speciation" title="Peripatric speciation">Quantum</a> · <a href="/wiki/Peripatric_speciation#Centrifugal_speciation" title="Peripatric speciation">Centrifugal</a> · <a href="/wiki/Founder_effect" title="Founder effect">Founder effect</a>)</li> <li><a href="/wiki/Parapatric_speciation" title="Parapatric speciation">Parapatric</a> (<a class="mw-selflink selflink">Clines</a> · <a href="/wiki/Ring_species" title="Ring species">Ring species</a>)</li> <li><a href="/wiki/Sympatric_speciation" title="Sympatric speciation">Sympatric</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Reproductive_isolation" title="Reproductive isolation">Isolating factors</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Adaptation" title="Adaptation">Adaptation</a></li> <li><a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a></li> <li><a href="/wiki/Sexual_selection" title="Sexual selection">Sexual selection</a></li> <li><a href="/wiki/Ecological_speciation" title="Ecological speciation">Ecological speciation</a> (<a href="/wiki/Parallel_speciation" title="Parallel speciation">Parallel speciation</a> · <a href="/wiki/Allochronic_speciation" title="Allochronic speciation">Allochrony</a>)</li> <li><a href="/wiki/Nonecological_speciation" title="Nonecological speciation">Nonecological speciation</a></li> <li><a href="/wiki/Assortative_mating" title="Assortative mating">Assortative mating</a></li> <li><a href="/wiki/Haldane%27s_rule" title="Haldane's rule">Haldane's rule</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Hybrid_(biology)" title="Hybrid (biology)">Hybrid concepts</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Hybrid_speciation" title="Hybrid speciation">Hybrid speciation</a> (<a href="/wiki/Polyploidy" title="Polyploidy">Polyploidy</a> · <a href="/wiki/Klepton" title="Klepton">Klepton</a> · <a href="/w/index.php?title=Recombinational_speciation&action=edit&redlink=1" class="new" title="Recombinational speciation (page does not exist)">Recombination</a>)</li> <li><a href="/wiki/Reinforcement_(speciation)" title="Reinforcement (speciation)">Reinforcement</a> (<a href="/wiki/Evidence_for_speciation_by_reinforcement" title="Evidence for speciation by reinforcement">evidence</a>)</li> <li><a href="/wiki/Secondary_contact" title="Secondary contact">Secondary contact</a></li> <li><a href="/wiki/Character_displacement" title="Character displacement">Character displacement</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Speciation in taxa</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/w/index.php?title=Speciation_in_birds&action=edit&redlink=1" class="new" title="Speciation in birds (page does not exist)">Birds</a></li> <li><a href="/w/index.php?title=Speciation_in_fish&action=edit&redlink=1" class="new" title="Speciation in fish (page does not exist)">Fish</a></li> <li><a href="/w/index.php?title=Speciation_in_insects&action=edit&redlink=1" class="new" title="Speciation in insects (page does not exist)">Insects</a></li> <li><a href="/w/index.php?title=Speciation_in_plants&action=edit&redlink=1" class="new" title="Speciation in plants (page does not exist)">Plants</a></li> <li><a href="/w/index.php?title=Speciation_in_the_fossil_record&action=edit&redlink=1" class="new" title="Speciation in the fossil record (page does not exist)">Fossils</a> (<a href="/wiki/Paleopolyploidy" title="Paleopolyploidy">Paleopolyploidy</a> · <a href="/wiki/Punctuated_equilibrium" title="Punctuated equilibrium">Punctuated equilibrium</a> · <a href="/wiki/Macroevolution" title="Macroevolution">Macroevolution</a> · <a href="/wiki/Chronospecies" title="Chronospecies">Chronospecies</a>)</li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><span class="noviewer" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <b><a href="/wiki/Category:Speciation" title="Category:Speciation">Category</a></b></li> <li><span class="noviewer" typeof="mw:File"><span title="Commons page"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/12px-Commons-logo.svg.png" decoding="async" width="12" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/18px-Commons-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/24px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></span></span> <b><a href="https://commons.wikimedia.org/wiki/Category:Speciation" class="extiw" title="commons:Category:Speciation">Commons</a></b></li> <li><span class="noviewer" typeof="mw:File"><a href="/wiki/File:Symbol_portal_class.svg" class="mw-file-description" title="Portal"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/16px-Symbol_portal_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/23px-Symbol_portal_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/e/e2/Symbol_portal_class.svg/31px-Symbol_portal_class.svg.png 2x" data-file-width="180" data-file-height="185" /></a></span> <b><a href="/wiki/Portal:Evolutionary_biology" title="Portal:Evolutionary biology">Evolutionary biology Portal</a></b></li> <li><span class="noviewer" typeof="mw:File"><span title="WikiProject"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/3/37/People_icon.svg/16px-People_icon.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/37/People_icon.svg/24px-People_icon.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/37/People_icon.svg/32px-People_icon.svg.png 2x" data-file-width="100" data-file-height="100" /></span></span> <b><a href="/wiki/Wikipedia:WikiProject_Evolutionary_biology" title="Wikipedia:WikiProject Evolutionary biology">WikiProject</a></b></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Evolutionary_biology" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Evolution" title="Template:Evolution"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Evolution" title="Template talk:Evolution"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Evolution" title="Special:EditPage/Template:Evolution"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Evolutionary_biology" style="font-size:114%;margin:0 4em"><a href="/wiki/Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><a href="/wiki/Introduction_to_evolution" title="Introduction to evolution">Introduction</a></li> <li><a href="/wiki/Outline_of_evolution" title="Outline of evolution">Outline</a></li> <li><a href="/wiki/Timeline_of_the_evolutionary_history_of_life" title="Timeline of the evolutionary history of life">Timeline of evolution</a></li> <li><a href="/wiki/History_of_life" title="History of life">History of life</a></li> <li><a href="/wiki/Index_of_evolutionary_biology_articles" title="Index of evolutionary biology articles">Index</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Evolution" title="Evolution">Evolution</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Abiogenesis" title="Abiogenesis">Abiogenesis</a></li> <li><a href="/wiki/Adaptation" title="Adaptation">Adaptation</a></li> <li><a href="/wiki/Adaptive_radiation" title="Adaptive radiation">Adaptive radiation</a></li> <li><a href="/wiki/Altruism_(biology)" title="Altruism (biology)">Altruism</a> <ul><li><a href="/wiki/Cheating_(biology)" title="Cheating (biology)">Cheating</a></li> <li><a href="/wiki/Reciprocal_altruism" title="Reciprocal altruism">Reciprocal</a></li></ul></li> <li><a href="/wiki/Baldwin_effect" title="Baldwin effect">Baldwin effect</a></li> <li><a href="/wiki/Cladistics" title="Cladistics">Cladistics</a></li> <li><a href="/wiki/Coevolution" title="Coevolution">Coevolution</a> <ul><li><a href="/wiki/Mutualism_(biology)" title="Mutualism (biology)">Mutualism</a></li></ul></li> <li><a href="/wiki/Common_descent" title="Common descent">Common descent</a></li> <li><a href="/wiki/Convergent_evolution" title="Convergent evolution">Convergence</a></li> <li><a href="/wiki/Divergent_evolution" title="Divergent evolution">Divergence</a></li> <li><a href="/wiki/Earliest_known_life_forms" title="Earliest known life forms">Earliest known life forms</a></li> <li><a href="/wiki/Evidence_of_common_descent" title="Evidence of common descent">Evidence of evolution</a></li> <li><a href="/wiki/Evolutionary_arms_race" title="Evolutionary arms race">Evolutionary arms race</a></li> <li><a href="/wiki/Evolutionary_pressure" title="Evolutionary pressure">Evolutionary pressure</a></li> <li><a href="/wiki/Exaptation" title="Exaptation">Exaptation</a></li> <li><a href="/wiki/Extinction" title="Extinction">Extinction</a> <ul><li><a href="/wiki/Extinction_event" title="Extinction event">Event</a></li></ul></li> <li><a href="/wiki/Homology_(biology)" title="Homology (biology)">Homology</a></li> <li><a href="/wiki/Last_universal_common_ancestor" title="Last universal common ancestor">Last universal common ancestor</a></li> <li><a href="/wiki/Macroevolution" title="Macroevolution">Macroevolution</a></li> <li><a href="/wiki/Microevolution" title="Microevolution">Microevolution</a></li> <li><a href="/wiki/Evolutionary_mismatch" title="Evolutionary mismatch">Mismatch</a></li> <li><a href="/wiki/Nonadaptive_radiation" title="Nonadaptive radiation">Non-adaptive radiation</a></li> <li><a href="/wiki/Abiogenesis" title="Abiogenesis">Origin of life</a></li> <li><a href="/wiki/Panspermia" title="Panspermia">Panspermia</a></li> <li><a href="/wiki/Parallel_evolution" title="Parallel evolution">Parallel evolution</a></li> <li><a href="/wiki/Signalling_theory" title="Signalling theory">Signalling theory</a> <ul><li><a href="/wiki/Handicap_principle" title="Handicap principle">Handicap principle</a></li></ul></li> <li><a href="/wiki/Speciation" title="Speciation">Speciation</a> <ul><li><a href="/wiki/Species" title="Species">Species</a></li> <li><a href="/wiki/Species_complex" title="Species complex">Species complex</a></li></ul></li> <li><a href="/wiki/Evolutionary_taxonomy" title="Evolutionary taxonomy">Taxonomy</a></li> <li><a href="/wiki/Unit_of_selection" title="Unit of selection">Unit of selection</a> <ul><li><a href="/wiki/Gene-centered_view_of_evolution" title="Gene-centered view of evolution">Gene-centered view of evolution</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Population_genetics" title="Population genetics">Population<br />genetics</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Selective_breeding" title="Selective breeding">Artificial selection</a></li> <li><a href="/wiki/Biodiversity" title="Biodiversity">Biodiversity</a></li> <li><a href="/wiki/Evolutionarily_stable_strategy" title="Evolutionarily stable strategy">Evolutionarily stable strategy</a></li> <li><a href="/wiki/Fisher%27s_principle" title="Fisher's principle">Fisher's principle</a></li> <li><a href="/wiki/Fitness_(biology)" title="Fitness (biology)">Fitness</a> <ul><li><a href="/wiki/Inclusive_fitness" title="Inclusive fitness">Inclusive</a></li></ul></li> <li><a href="/wiki/Gene_flow" title="Gene flow">Gene flow</a></li> <li><a href="/wiki/Genetic_drift" title="Genetic drift">Genetic drift</a></li> <li><a href="/wiki/Kin_selection" title="Kin selection">Kin selection</a> <ul><li><a href="/wiki/Parental_investment" title="Parental investment">Parental investment</a></li> <li><a href="/wiki/Parent%E2%80%93offspring_conflict" title="Parent–offspring conflict">Parent–offspring conflict</a></li></ul></li> <li><a href="/wiki/Mutation" title="Mutation">Mutation</a></li> <li><a href="/wiki/Population" title="Population">Population</a></li> <li><a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a></li> <li><a href="/wiki/Sexual_dimorphism" title="Sexual dimorphism">Sexual dimorphism</a></li> <li><a href="/wiki/Sexual_selection" title="Sexual selection">Sexual selection</a> <ul><li><a href="/wiki/Sexual_selection_in_flowering_plants" title="Sexual selection in flowering plants">Flowering plants</a></li> <li><a href="/wiki/Sexual_selection_in_fungi" title="Sexual selection in fungi">Fungi</a></li> <li><a href="/wiki/Mate_choice" title="Mate choice">Mate choice</a></li></ul></li> <li><a href="/wiki/Social_selection" title="Social selection">Social selection</a></li> <li><a href="/wiki/Trivers%E2%80%93Willard_hypothesis" title="Trivers–Willard hypothesis">Trivers–Willard hypothesis</a></li> <li><a href="/wiki/Genetic_variation" title="Genetic variation">Variation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Development</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Canalisation_(genetics)" title="Canalisation (genetics)">Canalisation</a></li> <li><a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evolutionary developmental biology</a></li> <li><a href="/wiki/Genetic_assimilation" title="Genetic assimilation">Genetic assimilation</a></li> <li><a href="/wiki/Inversion_(evolutionary_biology)" title="Inversion (evolutionary biology)">Inversion</a></li> <li><a href="/wiki/Modularity_(biology)" title="Modularity (biology)">Modularity</a></li> <li><a href="/wiki/Phenotypic_plasticity" title="Phenotypic plasticity">Phenotypic plasticity</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Taxon" title="Taxon">taxa</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_bacteria" title="Evolution of bacteria">Bacteria</a></li> <li><a href="/wiki/Evolution_of_birds" title="Evolution of birds">Birds</a> <ul><li><a href="/wiki/Origin_of_birds" title="Origin of birds">origin</a></li></ul></li> <li><a href="/wiki/Evolution_of_brachiopods" title="Evolution of brachiopods">Brachiopods</a></li> <li><a href="/wiki/Evolution_of_molluscs" title="Evolution of molluscs">Molluscs</a> <ul><li><a href="/wiki/Evolution_of_cephalopods" title="Evolution of cephalopods">Cephalopods</a></li></ul></li> <li><a href="/wiki/Dinosaur#Evolutionary_history" title="Dinosaur">Dinosaurs</a></li> <li><a href="/wiki/Evolution_of_fish" title="Evolution of fish">Fish</a></li> <li><a href="/wiki/Evolution_of_fungi" title="Evolution of fungi">Fungi</a></li> <li><a href="/wiki/Evolution_of_insects" title="Evolution of insects">Insects</a> <ul><li><a href="/wiki/Evolution_of_butterflies" title="Evolution of butterflies">butterflies</a></li></ul></li> <li><a href="/wiki/History_of_life" title="History of life">Life</a></li> <li><a href="/wiki/Evolution_of_mammals" title="Evolution of mammals">Mammals</a> <ul><li><a href="/wiki/Cat_gap" title="Cat gap">cats</a></li> <li><a href="/wiki/Canidae#Evolution" title="Canidae">canids</a> <ul><li><a href="/wiki/Evolution_of_the_wolf" title="Evolution of the wolf">wolves</a></li> <li><a href="/wiki/Domestication_of_the_dog" title="Domestication of the dog">dogs</a></li></ul></li> <li><a href="/wiki/Hyena#Evolution" title="Hyena">hyenas</a></li> <li><a href="/wiki/Evolution_of_cetaceans" title="Evolution of cetaceans">dolphins and whales</a></li> <li><a href="/wiki/Evolution_of_the_horse" title="Evolution of the horse">horses</a></li> <li><a href="/wiki/Evolution_of_Macropodidae" title="Evolution of Macropodidae">Kangaroos</a></li> <li><a href="/wiki/Evolution_of_primates" title="Evolution of primates">primates</a> <ul><li><a href="/wiki/Human_evolution" title="Human evolution">humans</a></li> <li><a href="/wiki/Evolution_of_lemurs" title="Evolution of lemurs">lemurs</a></li></ul></li> <li><a href="/wiki/Evolution_of_sirenians" title="Evolution of sirenians">sea cows</a></li></ul></li> <li><a href="/wiki/Evolutionary_history_of_plants" title="Evolutionary history of plants">Plants</a> <ul><li><a href="/wiki/Pollinator-mediated_selection" title="Pollinator-mediated selection">pollinator-mediated</a></li></ul></li> <li><a href="/wiki/Evolution_of_reptiles" title="Evolution of reptiles">Reptiles</a></li> <li><a href="/wiki/Evolution_of_spiders" title="Evolution of spiders">Spiders</a></li> <li><a href="/wiki/Evolution_of_tetrapods" title="Evolution of tetrapods">Tetrapods</a></li> <li><a href="/wiki/Viral_evolution" title="Viral evolution">Viruses</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Organ_(anatomy)" class="mw-redirect" title="Organ (anatomy)">organs</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_cells" title="Evolution of cells">Cell</a></li> <li><a href="/wiki/Models_of_DNA_evolution" title="Models of DNA evolution">DNA</a></li> <li><a href="/wiki/Evolution_of_flagella" title="Evolution of flagella">Flagella</a></li> <li><a href="/wiki/Eukaryote#Origin_of_eukaryotes" title="Eukaryote">Eukaryotes</a> <ul><li><a href="/wiki/Symbiogenesis" title="Symbiogenesis">symbiogenesis</a></li> <li><a href="/wiki/Chromosome#Eukaryotes" title="Chromosome">chromosome</a></li> <li><a href="/wiki/Endomembrane_system#Evolution" title="Endomembrane system">endomembrane system</a></li> <li><a href="/wiki/Mitochondrion#Origin_and_evolution" title="Mitochondrion">mitochondria</a></li> <li><a href="/wiki/Cell_nucleus#Evolution" title="Cell nucleus">nucleus</a></li> <li><a href="/wiki/Plastid#Origin" title="Plastid">plastids</a></li></ul></li> <li>In animals <ul><li><a href="/wiki/Evolution_of_the_eye" title="Evolution of the eye">eye</a></li> <li><a href="/wiki/Hair#Evolution" title="Hair">hair</a></li> <li><a href="/wiki/Evolution_of_mammalian_auditory_ossicles" title="Evolution of mammalian auditory ossicles">auditory ossicle</a></li> <li><a href="/wiki/Evolution_of_nervous_systems" title="Evolution of nervous systems">nervous system</a></li> <li><a href="/wiki/Evolution_of_the_brain" title="Evolution of the brain">brain</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Biological_process" title="Biological process">processes</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_ageing" title="Evolution of ageing">Aging</a> <ul><li><a href="/wiki/Death#Evolution_of_aging_and_mortality" title="Death">Death</a></li> <li><a href="/wiki/Programmed_cell_death#Evolutionary_origin_of_mitochondrial_apoptosis" title="Programmed cell death">Programmed cell death</a></li></ul></li> <li><a href="/wiki/Origin_of_avian_flight" title="Origin of avian flight">Avian flight</a></li> <li><a href="/wiki/Evolution_of_biological_complexity" title="Evolution of biological complexity">Biological complexity</a></li> <li><a href="/wiki/Cooperation_(evolution)" title="Cooperation (evolution)">Cooperation</a></li> <li><a href="/wiki/Evolution_of_color_vision" title="Evolution of color vision">Color vision</a> <ul><li><a href="/wiki/Evolution_of_color_vision_in_primates" title="Evolution of color vision in primates">in primates</a></li></ul></li> <li><a href="/wiki/Evolution_of_emotion" title="Evolution of emotion">Emotion</a></li> <li><a href="/wiki/Empathy#Evolution_across_species" title="Empathy">Empathy</a></li> <li><a href="/wiki/Evolutionary_ethics" title="Evolutionary ethics">Ethics</a></li> <li><a href="/wiki/Evolution_of_eusociality" title="Evolution of eusociality">Eusociality</a></li> <li><a href="/wiki/Immune_system#Evolution_and_other_mechanisms" title="Immune system">Immune system</a></li> <li><a href="/wiki/Metabolism#Evolution" title="Metabolism">Metabolism</a></li> <li><a href="/wiki/Monogamy_in_animals" title="Monogamy in animals">Monogamy</a></li> <li><a href="/wiki/Evolution_of_morality" title="Evolution of morality">Morality</a></li> <li><a href="/wiki/Mosaic_evolution" title="Mosaic evolution">Mosaic evolution</a></li> <li><a href="/wiki/Multicellular_organism#Evolutionary_history" title="Multicellular organism">Multicellularity</a></li> <li><a href="/wiki/Evolution_of_sexual_reproduction" title="Evolution of sexual reproduction">Sexual reproduction</a> <ul><li><a href="/wiki/Anisogamy#Evolution" title="Anisogamy">Gamete differentiation/sexes</a></li> <li><a href="/wiki/Biological_life_cycle" title="Biological life cycle">Life cycles/nuclear phases</a></li> <li><a href="/wiki/Mating_type" title="Mating type">Mating types</a></li> <li><a href="/wiki/Origin_and_function_of_meiosis" title="Origin and function of meiosis">Meiosis</a></li> <li><a href="/wiki/Sex-determination_system#Evolution" title="Sex-determination system">Sex-determination</a></li></ul></li> <li><a href="/wiki/Evolution_of_snake_venom" title="Evolution of snake venom">Snake venom</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Tempo_and_Mode_in_Evolution" title="Tempo and Mode in Evolution">Tempo and modes</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Phyletic_gradualism" title="Phyletic gradualism">Gradualism</a>/<a href="/wiki/Punctuated_equilibrium" title="Punctuated equilibrium">Punctuated equilibrium</a>/<a href="/wiki/Saltation_(biology)" title="Saltation (biology)">Saltationism</a></li> <li><a href="/wiki/Point_mutation" title="Point mutation">Micromutation</a>/<a href="/wiki/Mutationism#Later_mutationist_theories" title="Mutationism">Macromutation</a></li> <li><a href="/wiki/Uniformitarianism" title="Uniformitarianism">Uniformitarianism</a>/<a href="/wiki/Catastrophism" title="Catastrophism">Catastrophism</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Speciation" title="Speciation">Speciation</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Allopatric_speciation" title="Allopatric speciation">Allopatric</a></li> <li><a href="/wiki/Anagenesis" title="Anagenesis">Anagenesis</a></li> <li><a href="/wiki/Catagenesis_(biology)" title="Catagenesis (biology)">Catagenesis</a></li> <li><a href="/wiki/Cladogenesis" title="Cladogenesis">Cladogenesis</a></li> <li><a href="/wiki/Cospeciation" title="Cospeciation">Cospeciation</a></li> <li><a href="/wiki/Ecological_speciation" title="Ecological speciation">Ecological</a></li> <li><a href="/wiki/Hybrid_speciation" title="Hybrid speciation">Hybrid</a></li> <li><a href="/wiki/Nonecological_speciation" title="Nonecological speciation">Non-ecological</a></li> <li><a href="/wiki/Parapatric_speciation" title="Parapatric speciation">Parapatric</a></li> <li><a href="/wiki/Peripatric_speciation" title="Peripatric speciation">Peripatric</a></li> <li><a href="/wiki/Reinforcement_(speciation)" title="Reinforcement (speciation)">Reinforcement</a></li> <li><a href="/wiki/Sympatric_speciation" title="Sympatric speciation">Sympatric</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">History</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolutionary_ideas_of_the_Renaissance_and_Enlightenment" title="Evolutionary ideas of the Renaissance and Enlightenment">Renaissance and Enlightenment</a></li> <li><a href="/wiki/Transmutation_of_species" title="Transmutation of species">Transmutation of species</a></li> <li><a href="/wiki/David_Hume" title="David Hume">David Hume</a> <ul><li><i><a href="/wiki/Dialogues_Concerning_Natural_Religion" title="Dialogues Concerning Natural Religion">Dialogues Concerning Natural Religion</a></i></li></ul></li> <li><a href="/wiki/Charles_Darwin" title="Charles Darwin">Charles Darwin</a> <ul><li><i><a href="/wiki/On_the_Origin_of_Species" title="On the Origin of Species">On the Origin of Species</a></i></li></ul></li> <li><a href="/wiki/History_of_paleontology" title="History of paleontology">History of paleontology</a></li> <li><a href="/wiki/Transitional_fossil" title="Transitional fossil">Transitional fossil</a></li> <li><a href="/wiki/Blending_inheritance" title="Blending inheritance">Blending inheritance</a></li> <li><a href="/wiki/Mendelian_inheritance" title="Mendelian inheritance">Mendelian inheritance</a></li> <li><a href="/wiki/The_eclipse_of_Darwinism" title="The eclipse of Darwinism">The eclipse of Darwinism</a></li> <li><a href="/wiki/Neo-Darwinism" title="Neo-Darwinism">Neo-Darwinism</a></li> <li><a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis</a></li> <li><a href="/wiki/History_of_molecular_evolution" title="History of molecular evolution">History of molecular evolution</a></li> <li><a href="/wiki/Extended_evolutionary_synthesis" title="Extended evolutionary synthesis">Extended evolutionary synthesis</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Philosophy</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Darwinism" title="Darwinism">Darwinism</a></li> <li><a href="/wiki/Alternatives_to_Darwinian_evolution" title="Alternatives to Darwinian evolution">Alternatives</a> <ul><li><a href="/wiki/Catastrophism" title="Catastrophism">Catastrophism</a></li> <li><a href="/wiki/Lamarckism" title="Lamarckism">Lamarckism</a></li> <li><a href="/wiki/Orthogenesis" title="Orthogenesis">Orthogenesis</a></li> <li><a href="/wiki/Mutationism" title="Mutationism">Mutationism</a></li> <li><a href="/wiki/Saltation_(biology)" title="Saltation (biology)">Saltationism</a></li> <li><a href="/wiki/Structuralism_(biology)" title="Structuralism (biology)">Structuralism</a> <ul><li><a href="/wiki/Spandrel_(biology)" title="Spandrel (biology)">Spandrel</a></li></ul></li> <li><a href="/wiki/Theistic_evolution" title="Theistic evolution">Theistic</a></li> <li><a href="/wiki/Vitalism" title="Vitalism">Vitalism</a></li></ul></li> <li><a href="/wiki/Teleology_in_biology" title="Teleology in biology">Teleology in biology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Biogeography" title="Biogeography">Biogeography</a></li> <li><a href="/wiki/Ecological_genetics" title="Ecological genetics">Ecological genetics</a></li> <li><a href="/wiki/Evolutionary_medicine" title="Evolutionary medicine">Evolutionary medicine</a></li> <li><a href="/wiki/Group_selection" title="Group selection">Group selection</a> <ul><li><a href="/wiki/Cultural_evolution" title="Cultural evolution">Cultural evolution</a></li> <li><a href="/wiki/Cultural_group_selection" title="Cultural group selection">Cultural group selection</a></li> <li><a href="/wiki/Dual_inheritance_theory" title="Dual inheritance theory">Dual inheritance theory</a></li></ul></li> <li><a href="/wiki/Hologenome_theory_of_evolution" title="Hologenome theory of evolution">Hologenome theory of evolution</a></li> <li><a href="/wiki/Missing_heritability_problem" title="Missing heritability problem">Missing heritability problem</a></li> <li><a href="/wiki/Molecular_evolution" title="Molecular evolution">Molecular evolution</a></li> <li><a href="/wiki/Astrobiology" title="Astrobiology">Astrobiology</a></li> <li><a href="/wiki/Phylogenetics" title="Phylogenetics">Phylogenetics</a> <ul><li><a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">Tree</a></li></ul></li> <li><a href="/wiki/Polymorphism_(biology)" title="Polymorphism (biology)">Polymorphism</a></li> <li><a href="/wiki/Protocell" title="Protocell">Protocell</a></li> <li><a href="/wiki/Systematics" title="Systematics">Systematics</a></li> <li><a href="/wiki/Transgenerational_epigenetic_inheritance" title="Transgenerational epigenetic inheritance">Transgenerational epigenetic inheritance</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><span class="noviewer" 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