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Evolution of the wolf - Wikipedia

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id="toc-Canis_mosbachensis" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Canis_mosbachensis"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.6</span> <span><i>Canis mosbachensis</i></span> </div> </a> <ul id="toc-Canis_mosbachensis-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Canis_lupus" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Canis_lupus"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.7</span> <span><i>Canis lupus</i></span> </div> </a> <ul id="toc-Canis_lupus-sublist" class="vector-toc-list"> <li id="toc-Canis_lupus_bohemica" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Canis_lupus_bohemica"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.7.1</span> <span><i>Canis lupus bohemica</i></span> </div> </a> <ul id="toc-Canis_lupus_bohemica-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Canis_c.f._familiaris_-_Paleolithic_dog" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Canis_c.f._familiaris_-_Paleolithic_dog"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.8</span> <span><i>Canis c.f. familiaris</i> - Paleolithic dog</span> </div> </a> <ul id="toc-Canis_c.f._familiaris_-_Paleolithic_dog-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Canis_familiaris_(domestic_dog)" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Canis_familiaris_(domestic_dog)"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.9</span> <span><i>Canis familiaris</i> (domestic dog)</span> </div> </a> <ul id="toc-Canis_familiaris_(domestic_dog)-sublist" class="vector-toc-list"> <li id="toc-Comparison_with_modern_wolves" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Comparison_with_modern_wolves"> <div class="vector-toc-text"> <span class="vector-toc-numb">1.9.1</span> <span>Comparison with modern wolves</span> </div> </a> <ul id="toc-Comparison_with_modern_wolves-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> </ul> </li> <li id="toc-Genetic_record" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Genetic_record"> <div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Genetic record</span> </div> </a> <button aria-controls="toc-Genetic_record-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 Genetic record subsection</span> </button> <ul id="toc-Genetic_record-sublist" class="vector-toc-list"> <li id="toc-DNA_sequences" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#DNA_sequences"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1</span> <span>DNA sequences</span> </div> </a> <ul id="toc-DNA_sequences-sublist" class="vector-toc-list"> <li id="toc-Marker_issue" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Marker_issue"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1.1</span> <span>Marker issue</span> </div> </a> <ul id="toc-Marker_issue-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Timing_issue" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Timing_issue"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.1.2</span> <span>Timing issue</span> </div> </a> <ul id="toc-Timing_issue-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Wolf-like_canids" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Wolf-like_canids"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2</span> <span>Wolf-like canids</span> </div> </a> <ul id="toc-Wolf-like_canids-sublist" class="vector-toc-list"> <li id="toc-Admixture_with_an_extinct_unknown_canid" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Admixture_with_an_extinct_unknown_canid"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.2.1</span> <span>Admixture with an extinct unknown canid</span> </div> </a> <ul id="toc-Admixture_with_an_extinct_unknown_canid-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Out_of_Beringia" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Out_of_Beringia"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.3</span> <span>Out of Beringia</span> </div> </a> <ul id="toc-Out_of_Beringia-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Into_America_and_Japan" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Into_America_and_Japan"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.4</span> <span>Into America and Japan</span> </div> </a> <ul id="toc-Into_America_and_Japan-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Divergence_with_the_coyote" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Divergence_with_the_coyote"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.5</span> <span>Divergence with the coyote</span> </div> </a> <ul id="toc-Divergence_with_the_coyote-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Domestic_dog" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Domestic_dog"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.6</span> <span>Domestic dog</span> </div> </a> <ul id="toc-Domestic_dog-sublist" class="vector-toc-list"> <li id="toc-Dingo" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Dingo"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.6.1</span> <span>Dingo</span> </div> </a> <ul id="toc-Dingo-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Canis_variabilis" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Canis_variabilis"> <div class="vector-toc-text"> <span class="vector-toc-numb">2.7</span> <span><i>Canis variabilis</i></span> </div> </a> <ul id="toc-Canis_variabilis-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Rise_to_dominant_predator" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Rise_to_dominant_predator"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Rise to dominant predator</span> </div> </a> <ul id="toc-Rise_to_dominant_predator-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Wolf_population_differences" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Wolf_population_differences"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Wolf population differences</span> </div> </a> <button 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class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Notes"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Notes</span> </div> </a> <ul id="toc-Notes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>References</span> </div> </a> <ul id="toc-References-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Works" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Works"> <div class="vector-toc-text"> <span class="vector-toc-numb">7</span> <span>Works</span> </div> </a> <ul id="toc-Works-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 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class="firstHeading mw-first-heading"><span class="mw-page-title-main">Evolution of the wolf</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. 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dir="ltr"><p class="mw-empty-elt"> </p> <figure typeof="mw:File/Thumb"><a href="/wiki/File:British_Pleistocene_Mammalia_(1866)_Wolf_Cranium.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f9/British_Pleistocene_Mammalia_%281866%29_Wolf_Cranium.png/250px-British_Pleistocene_Mammalia_%281866%29_Wolf_Cranium.png" decoding="async" width="250" height="341" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/f/f9/British_Pleistocene_Mammalia_%281866%29_Wolf_Cranium.png 1.5x" data-file-width="350" data-file-height="478" /></a><figcaption>Illustration of a Pleistocene wolf cranium that was found in <a href="/wiki/Kents_Cavern" title="Kents Cavern">Kents Cavern</a>, Torquay, England<sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">&#91;</span>1<span class="cite-bracket">&#93;</span></a></sup></figcaption></figure> <p>It is widely agreed that the evolutionary lineage of the <a href="/wiki/Grey_wolf" class="mw-redirect" title="Grey wolf">grey wolf</a> can be traced back 2 million years to the <a href="/wiki/Early_Pleistocene" title="Early Pleistocene">Early Pleistocene</a> species <i><a href="/wiki/Canis_etruscus" title="Canis etruscus">Canis etruscus</a>,</i> and its successor the <a href="/wiki/Middle_Pleistocene" class="mw-redirect" title="Middle Pleistocene">Middle Pleistocene</a> <i><a href="/wiki/Canis_mosbachensis" title="Canis mosbachensis">Canis mosbachensis</a></i>.<sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">&#91;</span>2<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">&#91;</span>3<span class="cite-bracket">&#93;</span></a></sup> The grey wolf <i>Canis lupus</i> is a highly adaptable species that is able to exist in a range of environments and which possesses a wide distribution across the <a href="/wiki/Holarctic" class="mw-redirect" title="Holarctic">Holarctic</a>. Studies of modern grey wolves have identified distinct sub-populations that live in close proximity to each other.<sup id="cite_ref-musiani2007_4-0" class="reference"><a href="#cite_note-musiani2007-4"><span class="cite-bracket">&#91;</span>4<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-schweizer2016a_5-0" class="reference"><a href="#cite_note-schweizer2016a-5"><span class="cite-bracket">&#91;</span>5<span class="cite-bracket">&#93;</span></a></sup> This variation in sub-populations is closely linked to differences in habitat – precipitation, temperature, vegetation, and prey specialization – which affect cranio-dental plasticity.<sup id="cite_ref-geffen2004_6-0" class="reference"><a href="#cite_note-geffen2004-6"><span class="cite-bracket">&#91;</span>6<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-pilot2006_7-0" class="reference"><a href="#cite_note-pilot2006-7"><span class="cite-bracket">&#91;</span>7<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-flower2014_8-0" class="reference"><a href="#cite_note-flower2014-8"><span class="cite-bracket">&#91;</span>8<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-leonard2014_9-0" class="reference"><a href="#cite_note-leonard2014-9"><span class="cite-bracket">&#91;</span>9<span class="cite-bracket">&#93;</span></a></sup> </p><p>The earliest specimens of the modern grey wolf date to around 400,000 years ago,<sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">&#91;</span>10<span class="cite-bracket">&#93;</span></a></sup> or possibly earlier to 1 million years ago.<sup id="cite_ref-tedford2009_11-0" class="reference"><a href="#cite_note-tedford2009-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup> Most modern wolves share most of their common ancestry within the last 25-23,000 years from earlier Siberian wolf populations.<sup id="cite_ref-freedman2014_12-0" class="reference"><a href="#cite_note-freedman2014-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-fan2016_13-0" class="reference"><a href="#cite_note-fan2016-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-skoglund2015_14-0" class="reference"><a href="#cite_note-skoglund2015-14"><span class="cite-bracket">&#91;</span>14<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Loog2018_15-0" class="reference"><a href="#cite_note-Loog2018-15"><span class="cite-bracket">&#91;</span>15<span class="cite-bracket">&#93;</span></a></sup> While some sources have suggested that this is the result of a <a href="/wiki/Population_bottleneck" title="Population bottleneck">population bottleneck</a>, others suggest that this is a normal consequence of <a href="/wiki/Gene_flow" title="Gene flow">gene flow</a> homogenising wolf genomes across their range.<sup id="cite_ref-:2_16-0" class="reference"><a href="#cite_note-:2-16"><span class="cite-bracket">&#91;</span>16<span class="cite-bracket">&#93;</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Fossil_record">Fossil record</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=1" title="Edit section: Fossil record"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Canis#Taxonomy" title="Canis">Evolution of Canis</a> and <a href="/wiki/Canidae#Phylogenetic_relationships" title="Canidae">Canid relationships</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Canisetr.JPG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/39/Canisetr.JPG/220px-Canisetr.JPG" decoding="async" width="220" height="300" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/39/Canisetr.JPG/330px-Canisetr.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/39/Canisetr.JPG/440px-Canisetr.JPG 2x" data-file-width="1628" data-file-height="2221" /></a><figcaption><i>Canis etruscus</i> skull in the <a href="/wiki/Montevarchi" title="Montevarchi">Montevarchi</a> Paleontological Museum</figcaption></figure> <p>The fossil record for ancient vertebrates is composed of rarely occurring fragments from which it is often impossible to obtain genetic material. Researchers are limited to <a href="/wiki/Morphology_(biology)" title="Morphology (biology)">morphologic analysis</a> but it is difficult to estimate the intra-species and inter-species variations and relationships that existed between specimens across time and place. Some observations are debated by researchers who do not always agree, and hypotheses that are supported by some authors are challenged by others.<sup id="cite_ref-sardella2014_17-0" class="reference"><a href="#cite_note-sardella2014-17"><span class="cite-bracket">&#91;</span>17<span class="cite-bracket">&#93;</span></a></sup> </p><p>The <a href="/wiki/Cretaceous%E2%80%93Paleogene_extinction_event" title="Cretaceous–Paleogene extinction event">Cretaceous–Paleogene extinction event</a> occurred 66 million years ago and brought an end to the non-avian dinosaurs and the appearance of the first carnivorans.<sup id="cite_ref-FOOTNOTEWangTedford20088_18-0" class="reference"><a href="#cite_note-FOOTNOTEWangTedford20088-18"><span class="cite-bracket">&#91;</span>18<span class="cite-bracket">&#93;</span></a></sup> The name carnivoran is given to a member of the order <a href="/wiki/Carnivora" title="Carnivora">Carnivora</a>. Carnivorans possess a common arrangement of teeth called <a href="/wiki/Carnassials" class="mw-redirect" title="Carnassials">carnassials</a>, in which the first lower <a href="/wiki/Molar_(tooth)" title="Molar (tooth)">molar</a> and the last upper <a href="/wiki/Premolar" title="Premolar">premolar</a> possess blade-like enamel <a href="/wiki/Crown_(tooth)" title="Crown (tooth)">crowns</a> that act similar to a pair of shears for cutting meat. This dental arrangement has been modified by adaptation over the past 60 million years for diets composed of meat, for crushing vegetation, or for the loss of the carnassial function altogether as in seals, sea lions, and walruses. Today, not all carnivorans are <a href="/wiki/Carnivores" class="mw-redirect" title="Carnivores">carnivores</a>, such as the insect-eating <a href="/wiki/Aardwolf" title="Aardwolf">aardwolf</a>.<sup id="cite_ref-FOOTNOTEWangTedford20081_19-0" class="reference"><a href="#cite_note-FOOTNOTEWangTedford20081-19"><span class="cite-bracket">&#91;</span>19<span class="cite-bracket">&#93;</span></a></sup> </p><p>The carnivoran ancestors of the dog-like <a href="/wiki/Caniforms" class="mw-redirect" title="Caniforms">caniforms</a> and the cat-like <a href="/wiki/Feliforms" class="mw-redirect" title="Feliforms">feliforms</a> began their separate evolutionary paths just after the end of the dinosaurs. The first members of the dog family <a href="/wiki/Canidae" title="Canidae">Canidae</a> appeared 40 million years ago,<sup id="cite_ref-FOOTNOTEWangTedford2008116_20-0" class="reference"><a href="#cite_note-FOOTNOTEWangTedford2008116-20"><span class="cite-bracket">&#91;</span>20<span class="cite-bracket">&#93;</span></a></sup> of which only its subfamily the <a href="/wiki/Caninae" title="Caninae">Caninae</a> survives today in the form of the wolf-like and fox-like canines. The caniforms included the fox-like genus <i><a href="/wiki/Leptocyon" title="Leptocyon">Leptocyon</a></i> whose various species existed from 34 million YBP before branching 11.9 million YBP into <i><a href="/wiki/Vulpes" title="Vulpes">Vulpes</a></i> (foxes) and <a href="/wiki/Canis" title="Canis">Canini</a> (canines). The jackal-sized <i><a href="/wiki/Eucyon" title="Eucyon">Eucyon</a></i> existed in North America from 10 million YBP and by the <a href="/wiki/Early_Pliocene" class="mw-redirect" title="Early Pliocene">Early Pliocene</a> about 6–5 million YBP the coyote-like <i><a href="/wiki/Eucyon#Eucyon_davisi" title="Eucyon">Eucyon davisi</a></i><sup id="cite_ref-fossilworks5_21-0" class="reference"><a href="#cite_note-fossilworks5-21"><span class="cite-bracket">&#91;</span>21<span class="cite-bracket">&#93;</span></a></sup> invaded Eurasia. In North America it gave rise to early <i>Canis</i> which first appeared in the <a href="/wiki/Miocene" title="Miocene">Miocene</a> (6 million YBP) in south-western US and Mexico. By 5 million YBP the larger <i><a href="/wiki/Canis_lepophagus" title="Canis lepophagus">Canis lepophagus</a></i> appeared in the same region.<sup id="cite_ref-FOOTNOTEWangTedford200858_22-0" class="reference"><a href="#cite_note-FOOTNOTEWangTedford200858-22"><span class="cite-bracket">&#91;</span>22<span class="cite-bracket">&#93;</span></a></sup> </p><p>The canids that had immigrated from North America to Eurasia – <i><a href="/wiki/Eucyon" title="Eucyon">Eucyon</a></i>, <i><a href="/wiki/Vulpes" title="Vulpes">Vulpes</a></i>, and <i><a href="/wiki/Nyctereutes" title="Nyctereutes">Nyctereutes</a></i> – were small to medium-sized predators during the Late Miocene and Early Pliocene but they were not the top predators. The position of the canids would change with the arrival of <i>Canis</i> to become a dominant predator across the <a href="/wiki/Holarctic" class="mw-redirect" title="Holarctic">Holarctic</a>. The wolf-sized <i><a href="#Canis_chihliensis">C. chihliensis</a></i> appeared in northern China in the Mid-Pliocene around 4–3 million YBP.<sup id="cite_ref-FOOTNOTEWangTedford2008148_23-0" class="reference"><a href="#cite_note-FOOTNOTEWangTedford2008148-23"><span class="cite-bracket">&#91;</span>23<span class="cite-bracket">&#93;</span></a></sup> The large wolf-sized <i>Canis</i> appeared in the Middle <a href="/wiki/Pliocene" title="Pliocene">Pliocene</a> about 3 million years ago in the Yushe Basin, <a href="/wiki/Shanxi" title="Shanxi">Shanxi</a> Province, China. By 2.5 million years ago its range included the Nihewan Basin in <a href="/wiki/Yangyuan_County" title="Yangyuan County">Yangyuan County</a>, <a href="/wiki/Hebei" title="Hebei">Hebei</a>, China and Kuruksay, Tadzhikistan.<sup id="cite_ref-vislobokova2003_24-0" class="reference"><a href="#cite_note-vislobokova2003-24"><span class="cite-bracket">&#91;</span>24<span class="cite-bracket">&#93;</span></a></sup> This was followed by an explosion of <i>Canis</i> evolution across Eurasia in the Early Pleistocene around 1.8 million YBP in what is commonly referred to as the <i>wolf event</i>. It is associated with the formation of the <a href="/wiki/Mammoth_steppe" title="Mammoth steppe">mammoth steppe</a> and continental glaciation. <i>Canis</i> spread to Europe in the forms of <i><a href="/wiki/Canis_arnensis" title="Canis arnensis">C. arnensis</a></i>, <i><a href="/wiki/Canis_etruscus" title="Canis etruscus">C. etruscus</a></i>, and <i><a href="/wiki/Canis_falconeri" class="mw-redirect" title="Canis falconeri">C. falconeri</a></i>.<sup id="cite_ref-FOOTNOTEWangTedford2008148_23-1" class="reference"><a href="#cite_note-FOOTNOTEWangTedford2008148-23"><span class="cite-bracket">&#91;</span>23<span class="cite-bracket">&#93;</span></a></sup> Other studies state that the oldest <i>Canis</i> remains that have been found in Europe were from France and dated to 3.1 million YBP,<sup id="cite_ref-lacombat2008_25-0" class="reference"><a href="#cite_note-lacombat2008-25"><span class="cite-bracket">&#91;</span>25<span class="cite-bracket">&#93;</span></a></sup> followed by <i>Canis cf. etruscus</i> (where <a href="/wiki/Open_nomenclature#Usage_of_open_nomenclature" title="Open nomenclature">cf.</a> in Latin means confer, uncertain) from Italy dated to 2.2 million YBP.<sup id="cite_ref-rook1996_26-0" class="reference"><a href="#cite_note-rook1996-26"><span class="cite-bracket">&#91;</span>26<span class="cite-bracket">&#93;</span></a></sup> </p><p>The fossil record is incomplete but it is likely that wolves arose from a population of small, early canids.<sup id="cite_ref-nowak2003_27-0" class="reference"><a href="#cite_note-nowak2003-27"><span class="cite-bracket">&#91;</span>27<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: p241">&#58;&#8202;p241&#8202;</span></sup> Morphological evidence<sup id="cite_ref-nowak2003_27-1" class="reference"><a href="#cite_note-nowak2003-27"><span class="cite-bracket">&#91;</span>27<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: p239">&#58;&#8202;p239&#8202;</span></sup><sup id="cite_ref-nowak1979_28-0" class="reference"><a href="#cite_note-nowak1979-28"><span class="cite-bracket">&#91;</span>28<span class="cite-bracket">&#93;</span></a></sup> and genetic evidence<sup id="cite_ref-wayne1995_29-0" class="reference"><a href="#cite_note-wayne1995-29"><span class="cite-bracket">&#91;</span>29<span class="cite-bracket">&#93;</span></a></sup> both suggest that wolves evolved during the <a href="/wiki/Pliocene" title="Pliocene">Pliocene</a> and Early <a href="/wiki/Pleistocene" title="Pleistocene">Pleistocene</a> eras from the same lineage that also led to the coyote,<sup id="cite_ref-nowak2003_27-2" class="reference"><a href="#cite_note-nowak2003-27"><span class="cite-bracket">&#91;</span>27<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: p239">&#58;&#8202;p239&#8202;</span></sup> with fossil specimens indicating that the coyote and the wolf diverged from a common ancestor 1.5 million years ago.<sup id="cite_ref-nowak2003_27-3" class="reference"><a href="#cite_note-nowak2003-27"><span class="cite-bracket">&#91;</span>27<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: p240">&#58;&#8202;p240&#8202;</span></sup><sup id="cite_ref-nowak1979_28-1" class="reference"><a href="#cite_note-nowak1979-28"><span class="cite-bracket">&#91;</span>28<span class="cite-bracket">&#93;</span></a></sup> The ancestor of the jackal and the other extant members of the genus <i>Canis</i> had split from the lineage before this time.<sup id="cite_ref-nowak2003_27-4" class="reference"><a href="#cite_note-nowak2003-27"><span class="cite-bracket">&#91;</span>27<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: p240">&#58;&#8202;p240&#8202;</span></sup> </p><p>After this separation from a common ancestor the species that were believed to be involved in the further evolution of the wolf and coyote – and the beliefs of some <a href="/wiki/Paleontology" title="Paleontology">paleontologists</a> – diverged.<sup id="cite_ref-nowak2003_27-5" class="reference"><a href="#cite_note-nowak2003-27"><span class="cite-bracket">&#91;</span>27<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: p240">&#58;&#8202;p240&#8202;</span></sup> A number of researchers believed that the lines of <i><a href="/wiki/Canis_edwardii" title="Canis edwardii">C. priscolatrans</a></i>, <i>C. etruscus</i>, <i><a href="/wiki/Canis_rufus" class="mw-redirect" title="Canis rufus">C. rufus</a></i>, <i><a href="/wiki/Canis_lycaon" class="mw-redirect" title="Canis lycaon">C. lycaon</a></i>, and <i><a href="/wiki/Canis_lupus" class="mw-redirect" title="Canis lupus">C. lupus</a></i> were components involved in some way that lead to the modern wolf and coyote.<sup id="cite_ref-nowak2003_27-6" class="reference"><a href="#cite_note-nowak2003-27"><span class="cite-bracket">&#91;</span>27<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: p240">&#58;&#8202;p240&#8202;</span></sup><sup id="cite_ref-hoffmeister1954_30-0" class="reference"><a href="#cite_note-hoffmeister1954-30"><span class="cite-bracket">&#91;</span>30<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-lawrence1967_31-0" class="reference"><a href="#cite_note-lawrence1967-31"><span class="cite-bracket">&#91;</span>31<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-lawrence1975_32-0" class="reference"><a href="#cite_note-lawrence1975-32"><span class="cite-bracket">&#91;</span>32<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-martin1974_33-0" class="reference"><a href="#cite_note-martin1974-33"><span class="cite-bracket">&#91;</span>33<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-webb1974_34-0" class="reference"><a href="#cite_note-webb1974-34"><span class="cite-bracket">&#91;</span>34<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-wayne1991_35-0" class="reference"><a href="#cite_note-wayne1991-35"><span class="cite-bracket">&#91;</span>35<span class="cite-bracket">&#93;</span></a></sup> </p> <table style="border: 1px solid #ccc; vertical-align: middle; width: 248px; margin:0.2em 0 0.2em 1em; float:right; clear:right;"> <tbody><tr> <td style="text-align: center; font-weight: bold; border: 0; padding: 0;">Wolf evolution </td></tr> <tr> <td style="padding: 5px"> <div class="clade"><style data-mw-deduplicate="TemplateStyles:r1258728058">body.skin-vector-2022 .mw-parser-output div.clade,body.skin-minerva .mw-parser-output div.clade{overflow-x:auto;overflow-y:hidden}body.skin-minerva .mw-parser-output div.clade p{font-size:inherit}.mw-parser-output table.clade{border-spacing:0;margin:0;font-size:100%;line-height:100%;border-collapse:separate;width:auto}.mw-parser-output table.clade table.clade{width:100%;line-height:inherit}.mw-parser-output table.clade td.clade-label{min-width:0.2em;width:0.2em;padding:0.1em 0.25em;vertical-align:bottom;text-align:center;border-left:1px solid;border-bottom:1px solid;white-space:nowrap}.mw-parser-output table.clade td.clade-label::before,.mw-parser-output table.clade td.clade-slabel::before{content:"\2060 "}.mw-parser-output table.clade td.clade-fixed-width{overflow:hidden;text-overflow:ellipsis}.mw-parser-output table.clade td.clade-fixed-width:hover{overflow:visible}.mw-parser-output table.clade td.clade-label.first{border-left:none;border-right:none}.mw-parser-output table.clade td.clade-label.reverse{border-left:none;border-right:1px solid}.mw-parser-output table.clade td.clade-slabel{padding:0.1em 0.25em;vertical-align:top;text-align:center;border-left:1px solid;white-space:nowrap}.mw-parser-output table.clade td.clade-slabel:hover{overflow:visible}.mw-parser-output table.clade td.clade-slabel.last{border-left:none;border-right:none}.mw-parser-output table.clade td.clade-slabel.reverse{border-left:none;border-right:1px solid}.mw-parser-output table.clade td.clade-bar{vertical-align:middle;text-align:left;padding:0 0.5em;position:relative}.mw-parser-output table.clade td.clade-bar.reverse{text-align:right;position:relative}.mw-parser-output table.clade td.clade-leaf{border:0;padding:0;text-align:left}.mw-parser-output table.clade td.clade-leafR{border:0;padding:0;text-align:right}.mw-parser-output table.clade td.clade-leaf.reverse{text-align:right}.mw-parser-output table.clade:hover span.linkA{background-color:yellow}.mw-parser-output table.clade:hover span.linkB{background-color:green}</style> <table class="clade" style="font-size:75%; line-height:65%;width:450px;"> <tbody><tr> <td class="clade-label first">†<i><a href="/wiki/Canis_lepophagus" title="Canis lepophagus">Canis&#160;lepophagus</a></i> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"><i><a href="/wiki/Canis_latrans" class="mw-redirect" title="Canis latrans">Canis&#160;latrans</a></i>&#160;(Coyote) </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p><span typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_IX).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/41/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IX%29.jpg/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IX%29.jpg" decoding="async" width="50" height="42" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/41/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IX%29.jpg/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IX%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/41/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IX%29.jpg/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IX%29.jpg 2x" data-file-width="414" data-file-height="347" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label">†<i><a href="/wiki/Canis_edwardii#Canis_priscolatrans" title="Canis edwardii">Canis&#160;priscolatrans</a></i> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p>extinct taxa </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label">†<i><a href="/wiki/Canis_armbrusteri" class="mw-redirect" title="Canis armbrusteri">Canis&#160;armbrusteri</a></i> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first">†<i><a href="/wiki/Canis_dirus" class="mw-redirect" title="Canis dirus">Canis&#160;dirus</a></i>&#160;(Dire&#160;wolf) </td> <td rowspan="2" class="clade-leaf"> <p><span typeof="mw:File"><a href="/wiki/File:The_American_Museum_journal_(c1900-(1918))_(Aenocyon_dirus).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fd/The_American_Museum_journal_%28c1900-%281918%29%29_%28Aenocyon_dirus%29.jpg/80px-The_American_Museum_journal_%28c1900-%281918%29%29_%28Aenocyon_dirus%29.jpg" decoding="async" width="80" height="44" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fd/The_American_Museum_journal_%28c1900-%281918%29%29_%28Aenocyon_dirus%29.jpg/120px-The_American_Museum_journal_%28c1900-%281918%29%29_%28Aenocyon_dirus%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fd/The_American_Museum_journal_%28c1900-%281918%29%29_%28Aenocyon_dirus%29.jpg/160px-The_American_Museum_journal_%28c1900-%281918%29%29_%28Aenocyon_dirus%29.jpg 2x" data-file-width="1720" data-file-height="944" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label">†<i><a href="/wiki/Canis_nehringi" title="Canis nehringi">Canis&#160;nehringi</a></i> </td> <td rowspan="2" class="clade-leaf"> <p><span typeof="mw:File"><a href="/wiki/File:Large_sized_canid_icon.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/72/Large_sized_canid_icon.jpg/50px-Large_sized_canid_icon.jpg" decoding="async" width="50" height="31" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/72/Large_sized_canid_icon.jpg/75px-Large_sized_canid_icon.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/72/Large_sized_canid_icon.jpg/100px-Large_sized_canid_icon.jpg 2x" data-file-width="500" data-file-height="314" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label">†<i><a href="/wiki/Canis_nehringi#Canis_gezi" title="Canis nehringi">Canis&#160;gezi</a></i> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p><span typeof="mw:File"><a href="/wiki/File:Large_sized_canid_icon.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/72/Large_sized_canid_icon.jpg/50px-Large_sized_canid_icon.jpg" decoding="async" width="50" height="31" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/72/Large_sized_canid_icon.jpg/75px-Large_sized_canid_icon.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/72/Large_sized_canid_icon.jpg/100px-Large_sized_canid_icon.jpg 2x" data-file-width="500" data-file-height="314" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label">†<i>Canis&#160;etruscus</i> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p>†<i><a href="/wiki/Canis_variabilis" class="mw-redirect" title="Canis variabilis">Canis variabilis</a></i><span typeof="mw:File"><a href="/wiki/File:Canis_lupus_variabilis.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/96/Canis_lupus_variabilis.jpg/50px-Canis_lupus_variabilis.jpg" decoding="async" width="50" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/96/Canis_lupus_variabilis.jpg/75px-Canis_lupus_variabilis.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/96/Canis_lupus_variabilis.jpg/100px-Canis_lupus_variabilis.jpg 2x" data-file-width="301" data-file-height="243" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label">†<i><a href="#Canis_mosbachensis">Canis&#160;mosbachensis</a></i> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"><i><a href="/wiki/Canis_rufus" class="mw-redirect" title="Canis rufus">Canis&#160;rufus</a></i>&#160;(red&#160;wolf) </td> <td rowspan="2" class="clade-leaf"> <p><span typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_V)_C._l._rufus_mod.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fa/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_V%29_C._l._rufus_mod.jpg/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_V%29_C._l._rufus_mod.jpg" decoding="async" width="50" height="35" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fa/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_V%29_C._l._rufus_mod.jpg/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_V%29_C._l._rufus_mod.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fa/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_V%29_C._l._rufus_mod.jpg/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_V%29_C._l._rufus_mod.jpg 2x" data-file-width="491" data-file-height="344" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"><i><a href="/wiki/Canis_lupus" class="mw-redirect" title="Canis lupus">Canis&#160;lupus</a></i>&#160;(grey&#160;wolf) </td> <td rowspan="2" class="clade-leaf"> <p><span typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_I).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c4/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.jpg/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.jpg" decoding="async" width="50" height="31" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c4/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.jpg/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c4/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.jpg/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.jpg 2x" data-file-width="500" data-file-height="314" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"><i><a href="/wiki/Canis_familiaris" class="mw-redirect" title="Canis familiaris">Canis&#160;familiaris</a></i>&#160;(dog) </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p><span style="display: inline-block; transform: scaleX(-1);"><span typeof="mw:File"><a href="/wiki/File:202104_Dog.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f3/202104_Dog.svg/50px-202104_Dog.svg.png" decoding="async" width="50" height="38" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f3/202104_Dog.svg/75px-202104_Dog.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f3/202104_Dog.svg/100px-202104_Dog.svg.png 2x" data-file-width="512" data-file-height="384" /></a></span></span> </p> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td style="text-align: left; font-size: 88%; border: 0; padding: 0;">Proposed evolution and branching of genus <i>Canis</i> towards the wolf.<sup id="cite_ref-nowak2003_27-7" class="reference"><a href="#cite_note-nowak2003-27"><span class="cite-bracket">&#91;</span>27<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: p240">&#58;&#8202;p240&#8202;</span></sup> </td></tr></tbody></table> <div class="mw-heading mw-heading3"><h3 id="Canis_lepophagus"><i>Canis lepophagus</i></h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=2" title="Edit section: Canis lepophagus"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><i><a href="/wiki/Canis_lepophagus" title="Canis lepophagus">Canis lepophagus</a></i> lived in the early Pliocene in North America.<sup id="cite_ref-wang2008_36-0" class="reference"><a href="#cite_note-wang2008-36"><span class="cite-bracket">&#91;</span>36<span class="cite-bracket">&#93;</span></a></sup> <a href="/wiki/Bj%C3%B6rn_Kurt%C3%A9n" title="Björn Kurtén">Kurten</a> proposed that the <a href="/wiki/Blancan" title="Blancan">Blancan</a> <i>C. lepophagus</i><sup id="cite_ref-fossilworks1_37-0" class="reference"><a href="#cite_note-fossilworks1-37"><span class="cite-bracket">&#91;</span>37<span class="cite-bracket">&#93;</span></a></sup> derived from smaller <a href="/wiki/Miocene" title="Miocene">Miocene</a> <i>Canis</i> species in North America. It then became widespread across Eurasia where it was either identical to, or closely related with, <i>C. arnensis</i> of Europe.<sup id="cite_ref-nowak2003_27-8" class="reference"><a href="#cite_note-nowak2003-27"><span class="cite-bracket">&#91;</span>27<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: p241">&#58;&#8202;p241&#8202;</span></sup><sup id="cite_ref-kurten1974_38-0" class="reference"><a href="#cite_note-kurten1974-38"><span class="cite-bracket">&#91;</span>38<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-kurten1980_39-0" class="reference"><a href="#cite_note-kurten1980-39"><span class="cite-bracket">&#91;</span>39<span class="cite-bracket">&#93;</span></a></sup> </p><p>Johnston describes <i>C. lepophagus</i> as having a more slender skull and skeleton than in the modern coyote.<sup id="cite_ref-johnston1938_40-0" class="reference"><a href="#cite_note-johnston1938-40"><span class="cite-bracket">&#91;</span>40<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: 385">&#58;&#8202;385&#8202;</span></sup> Robert M. Nowak found that the early populations had small, delicate and narrowly proportioned skulls that resemble small coyotes and appear to be ancestral to <i><a href="/wiki/Coyote" title="Coyote">C. latrans</a></i>.<sup id="cite_ref-nowak2003_27-9" class="reference"><a href="#cite_note-nowak2003-27"><span class="cite-bracket">&#91;</span>27<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: p241">&#58;&#8202;p241&#8202;</span></sup> Johnson noted that some specimens found in Cita Canyon, Texas, had larger, broader skulls,<sup id="cite_ref-johnston1938_40-1" class="reference"><a href="#cite_note-johnston1938-40"><span class="cite-bracket">&#91;</span>40<span class="cite-bracket">&#93;</span></a></sup> and along with other fragments Nowak suggested that these were evolving into wolves.<sup id="cite_ref-nowak2003_27-10" class="reference"><a href="#cite_note-nowak2003-27"><span class="cite-bracket">&#91;</span>27<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: p241">&#58;&#8202;p241&#8202;</span></sup><sup id="cite_ref-nowak1979_28-2" class="reference"><a href="#cite_note-nowak1979-28"><span class="cite-bracket">&#91;</span>28<span class="cite-bracket">&#93;</span></a></sup> </p><p>Tedford disagreed with previous authors and found that its cranio-dental morphology lacked some characteristics that are shared by <i>C. lupus</i> and <i>C. latrans</i>, and therefore there was not a close relationship but it did suggest <i>C. lepophagus</i> was the ancestor of both wolves and coyotes.<sup id="cite_ref-tedford2009_11-1" class="reference"><a href="#cite_note-tedford2009-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: p119">&#58;&#8202;p119&#8202;</span></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Canis_armbrusteri"><i>Canis armbrusteri</i></h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=3" title="Edit section: Canis armbrusteri"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The North American wolves became larger, with tooth specimens indicating that <i>C. priscolatrans</i> diverged into the large wolf <i><a href="/wiki/Canis_ambrusteri" class="mw-redirect" title="Canis ambrusteri">C. armbrusteri</a></i>.<sup id="cite_ref-nowak2003_27-11" class="reference"><a href="#cite_note-nowak2003-27"><span class="cite-bracket">&#91;</span>27<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: p242">&#58;&#8202;p242&#8202;</span></sup><sup id="cite_ref-berta1995_41-0" class="reference"><a href="#cite_note-berta1995-41"><span class="cite-bracket">&#91;</span>41<span class="cite-bracket">&#93;</span></a></sup> during the Middle Pleistocene in North America.<sup id="cite_ref-nowak1979_28-3" class="reference"><a href="#cite_note-nowak1979-28"><span class="cite-bracket">&#91;</span>28<span class="cite-bracket">&#93;</span></a></sup> Robert A. Martin disagreed, and believed that <i>C. armbrusteri</i><sup id="cite_ref-fossilworks3_42-0" class="reference"><a href="#cite_note-fossilworks3-42"><span class="cite-bracket">&#91;</span>42<span class="cite-bracket">&#93;</span></a></sup> was <i>C. lupus</i>.<sup id="cite_ref-martin1974_33-1" class="reference"><a href="#cite_note-martin1974-33"><span class="cite-bracket">&#91;</span>33<span class="cite-bracket">&#93;</span></a></sup> Nowak disagreed with Martin and proposed that <i>C. armbrusteri</i> was not related to <i>C. lupus</i> but <i>C. priscolatrans</i>, which then gave rise to <i><a href="/wiki/Dire_wolf" title="Dire wolf">A. dirus</a></i>. Tedford proposed that the South American <i><a href="/wiki/Canis_nehringi#Canis_gezi" title="Canis nehringi">C. gezi</a></i> and <i><a href="/wiki/Canis_nehringi" title="Canis nehringi">C. nehringi</a></i> share dental and cranial similarities developed for hypercarnivory, suggesting <i>C. armbrusteri</i> was the common ancestor of <i>C. gezi</i>, <i>C. nehringi</i> and <i>A. dirus</i>.<sup id="cite_ref-tedford2009_11-2" class="reference"><a href="#cite_note-tedford2009-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: 148">&#58;&#8202;148&#8202;</span></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Aenocyon_dirus"><i>Aenocyon dirus</i></h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=4" title="Edit section: Aenocyon dirus"><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/Dire_wolf" title="Dire wolf">Dire wolf</a></div> <p>In 1908 the paleontologist <a href="/wiki/John_Campbell_Merriam" title="John Campbell Merriam">John Campbell Merriam</a> began retrieving numerous fossilized bone fragments of a large wolf from the Rancho La<span class="nowrap">&#160;</span>Brea tar pits. By 1912 he had found a skeleton sufficiently complete to be able to formally recognize these and the previously found specimens under the name <i>C.<span class="nowrap">&#160;</span>dirus</i> (Leidy 1858)<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">&#91;<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (May 2022)">citation needed</span></a></i>&#93;</sup>. </p><p><i>Canis dirus</i>,<sup id="cite_ref-fossilworks4_43-0" class="reference"><a href="#cite_note-fossilworks4-43"><span class="cite-bracket">&#91;</span>43<span class="cite-bracket">&#93;</span></a></sup> or as it is now widely recognized, <i>Aenocyon dirus</i>, lived in the late Pleistocene to early Holocene in North and South America.<sup id="cite_ref-kurten1984_44-0" class="reference"><a href="#cite_note-kurten1984-44"><span class="cite-bracket">&#91;</span>44<span class="cite-bracket">&#93;</span></a></sup> In 1987, a new hypothesis proposed that a mammal population could give rise to a larger form called a hypermorph during times when food was abundant, but when food later became scarce the hypermorph would either adapt to a smaller form or go extinct. This hypothesis might explain the large body sizes found in many Late Pleistocene mammals compared to their modern counterparts. Both extinction and <a href="/wiki/Speciation" title="Speciation">speciation</a> – a new species splitting from an older one – could occur together during periods of climatic extremes.<sup id="cite_ref-geist1998_45-0" class="reference"><a href="#cite_note-geist1998-45"><span class="cite-bracket">&#91;</span>45<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-geist1987_46-0" class="reference"><a href="#cite_note-geist1987-46"><span class="cite-bracket">&#91;</span>46<span class="cite-bracket">&#93;</span></a></sup> Gloria<span class="nowrap">&#160;</span>D.<span class="nowrap">&#160;</span>Goulet agreed with Martin and further proposed that this hypothesis might explain the sudden appearance of <i>A.<span class="nowrap">&#160;</span>dirus</i> in North America, and that because of the similarities in their skull shapes that <i>C.<span class="nowrap">&#160;</span>lupus</i> gave rise to the <i>A.<span class="nowrap">&#160;</span>dirus</i> hypermorph due to abundant game, a stable environment, and large competitors.<sup id="cite_ref-goulet1993_47-0" class="reference"><a href="#cite_note-goulet1993-47"><span class="cite-bracket">&#91;</span>47<span class="cite-bracket">&#93;</span></a></sup> Nowak, Kurten and Berta disagreed with Goulet and proposed that <i>A. dirus</i> was not derived from <i>C. lupus</i>.<sup id="cite_ref-nowak1979_28-4" class="reference"><a href="#cite_note-nowak1979-28"><span class="cite-bracket">&#91;</span>28<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-kurten1980_39-1" class="reference"><a href="#cite_note-kurten1980-39"><span class="cite-bracket">&#91;</span>39<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-berta1988_48-0" class="reference"><a href="#cite_note-berta1988-48"><span class="cite-bracket">&#91;</span>48<span class="cite-bracket">&#93;</span></a></sup> The three noted paleontologists Xiaoming Wang, R. H. Tedford and R. M. Nowak have all proposed that <i>A. dirus</i> had evolved from <i>C. armbrusteri</i>,<sup id="cite_ref-FOOTNOTEWangTedford200852_49-0" class="reference"><a href="#cite_note-FOOTNOTEWangTedford200852-49"><span class="cite-bracket">&#91;</span>49<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-tedford2009_11-3" class="reference"><a href="#cite_note-tedford2009-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: 181">&#58;&#8202;181&#8202;</span></sup> with Nowak stating that there were specimens from Cumberland Cave, Maryland, that indicated <i>C. armbrusteri</i> diverging into <i>A. dirus</i>.<sup id="cite_ref-nowak2003_27-12" class="reference"><a href="#cite_note-nowak2003-27"><span class="cite-bracket">&#91;</span>27<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: p243">&#58;&#8202;p243&#8202;</span></sup><sup id="cite_ref-nowak2002_50-0" class="reference"><a href="#cite_note-nowak2002-50"><span class="cite-bracket">&#91;</span>50<span class="cite-bracket">&#93;</span></a></sup> The two taxa share a number of characteristics (<a href="/wiki/Synapomorphy" class="mw-redirect" title="Synapomorphy">synapomorphy</a>), which suggests an origin of <i>A. dirus</i> in the late <a href="/wiki/Irvingtonian" title="Irvingtonian">Irvingtonian</a> in the open terrain in the midcontinent, and then later expanding eastward and displacing its ancestor <i>C. armbrusteri</i>.<sup id="cite_ref-tedford2009_11-4" class="reference"><a href="#cite_note-tedford2009-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: 181">&#58;&#8202;181&#8202;</span></sup> </p><p>However, in 2021, a study indicated the dire wolf to be a highly divergent lineage which last shared a <a href="/wiki/Most_recent_common_ancestor" title="Most recent common ancestor">most recent common ancestor</a> with the wolf-like canines 5.7 million years ago. The morphological similarity between dire wolves and gray wolves was concluded to be due to <a href="/wiki/Convergent_evolution" title="Convergent evolution">convergent evolution</a>. This finding indicates that the wolf and coyote lineages evolved in isolation from the dire wolf lineage. The study proposes an early origin of the dire wolf lineage in the Americas, and that this geographic isolation allowed them to develop a degree of <a href="/wiki/Reproductive_isolation" title="Reproductive isolation">reproductive isolation</a> since their divergence 5.7 million years ago. Coyotes, dholes, gray wolves, and the extinct <i><a href="/wiki/Xenocyon" title="Xenocyon">Xenocyon</a></i> evolved in Eurasia and expanded into North America relatively recently during the Late Pleistocene, therefore there was no admixture with the dire wolf. As a result, the study found that the correct binomial name of the dire wolf is <i>Aenocyon dirus</i>, as proposed by Merriam in 1918. The long-term isolation of the dire wolf lineage implies that other American fossil taxa, including <i>C. armbrusteri</i> and <i>C. edwardii</i>, may also belong to the dire wolf's lineage.<sup id="cite_ref-Perri2021_51-0" class="reference"><a href="#cite_note-Perri2021-51"><span class="cite-bracket">&#91;</span>51<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Canis_chihliensis"><i>Canis chihliensis</i></h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=5" title="Edit section: Canis chihliensis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <table style="border: 1px solid #ccc; vertical-align: middle; width: 248px; margin:0.2em 0 0.2em 1em; float:right; clear:right;"> <tbody><tr> <td style="text-align: center; font-weight: bold; border: 0; padding: 0;">Wolf evolution – alternate proposal </td></tr> <tr> <td style="padding: 5px"> <div class="clade"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade" style="font-size:75%; line-height:65%;width:450px;"> <tbody><tr> <td class="clade-label first">†<i><a href="/wiki/Eucyon" title="Eucyon">Eucyon</a>&#160;davisi</i> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first">†<i><a href="/wiki/Canis_lepophagus" title="Canis lepophagus">Canis&#160;lepophagus</a></i> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p>extinct taxa </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label">†<i><a href="/wiki/Canis_edwardii" title="Canis edwardii">Canis&#160;edwardii</a></i> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"><i><a href="/wiki/Golden_Jackal" class="mw-redirect" title="Golden Jackal">Canis&#160;aureus</a></i>&#160;(Golden&#160;jackal) </td> <td rowspan="2" class="clade-leaf"> <p><span typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_X).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fd/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_X%29.jpg/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_X%29.jpg" decoding="async" width="50" height="32" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fd/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_X%29.jpg/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_X%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fd/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_X%29.jpg/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_X%29.jpg 2x" data-file-width="454" data-file-height="287" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"><i><a href="/wiki/Canis_latrans" class="mw-redirect" title="Canis latrans">Canis&#160;latrans</a></i>&#160;(coyote) </td> <td rowspan="2" class="clade-leaf"> <p><span typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_IX).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/41/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IX%29.jpg/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IX%29.jpg" decoding="async" width="50" height="42" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/41/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IX%29.jpg/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IX%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/41/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IX%29.jpg/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IX%29.jpg 2x" data-file-width="414" data-file-height="347" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label">†<i>Canis&#160;chihliensis</i> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first">†<i><a href="/wiki/Canis_armbrusteri" class="mw-redirect" title="Canis armbrusteri">Canis&#160;armbrusteri</a></i> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first">†<i><a href="/wiki/Canis_dirus" class="mw-redirect" title="Canis dirus">Canis&#160;dirus</a></i> </td> <td rowspan="2" class="clade-leaf"> <p><span typeof="mw:File"><a href="/wiki/File:The_American_Museum_journal_(c1900-(1918))_(Aenocyon_dirus).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fd/The_American_Museum_journal_%28c1900-%281918%29%29_%28Aenocyon_dirus%29.jpg/50px-The_American_Museum_journal_%28c1900-%281918%29%29_%28Aenocyon_dirus%29.jpg" decoding="async" width="50" height="27" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fd/The_American_Museum_journal_%28c1900-%281918%29%29_%28Aenocyon_dirus%29.jpg/75px-The_American_Museum_journal_%28c1900-%281918%29%29_%28Aenocyon_dirus%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fd/The_American_Museum_journal_%28c1900-%281918%29%29_%28Aenocyon_dirus%29.jpg/100px-The_American_Museum_journal_%28c1900-%281918%29%29_%28Aenocyon_dirus%29.jpg 2x" data-file-width="1720" data-file-height="944" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label">†<i><a href="/wiki/Canis_nehringi" title="Canis nehringi">Canis&#160;nehringi</a></i> </td> <td rowspan="2" class="clade-leaf"> <p><span typeof="mw:File"><a href="/wiki/File:Large_sized_canid_icon.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/72/Large_sized_canid_icon.jpg/50px-Large_sized_canid_icon.jpg" decoding="async" width="50" height="31" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/72/Large_sized_canid_icon.jpg/75px-Large_sized_canid_icon.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/72/Large_sized_canid_icon.jpg/100px-Large_sized_canid_icon.jpg 2x" data-file-width="500" data-file-height="314" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label">†<i><a href="/wiki/Canis_nehringi#Canis_gezi" title="Canis nehringi">Canis&#160;gezi</a></i> </td> <td rowspan="2" class="clade-leaf"> <p><span typeof="mw:File"><a href="/wiki/File:Large_sized_canid_icon.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/72/Large_sized_canid_icon.jpg/50px-Large_sized_canid_icon.jpg" decoding="async" width="50" height="31" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/72/Large_sized_canid_icon.jpg/75px-Large_sized_canid_icon.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/72/Large_sized_canid_icon.jpg/100px-Large_sized_canid_icon.jpg 2x" data-file-width="500" data-file-height="314" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"><i><a href="/wiki/Canis_lupus" class="mw-redirect" title="Canis lupus">Canis&#160;lupus</a></i> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"><a href="/wiki/Grey_wolf" class="mw-redirect" title="Grey wolf">Grey&#160;wolf</a> </td> <td rowspan="2" class="clade-leaf"> <p><span typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_I).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c4/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.jpg/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.jpg" decoding="async" width="50" height="31" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c4/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.jpg/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c4/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.jpg/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.jpg 2x" data-file-width="500" data-file-height="314" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"><a href="/wiki/Domestic_dog" class="mw-redirect" title="Domestic dog">Domestic&#160;dog</a> </td> <td rowspan="2" class="clade-leaf"> <p><span style="display: inline-block; transform: scaleX(-1);"><span typeof="mw:File"><a href="/wiki/File:202104_Dog.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f3/202104_Dog.svg/50px-202104_Dog.svg.png" decoding="async" width="50" height="38" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f3/202104_Dog.svg/75px-202104_Dog.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f3/202104_Dog.svg/100px-202104_Dog.svg.png 2x" data-file-width="512" data-file-height="384" /></a></span></span> </p> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label">†<i><a href="/wiki/Canis_falconeri" class="mw-redirect" title="Canis falconeri">Canis&#160;falconeri</a></i> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first">Extinct&#160;taxa </td> <td rowspan="2" class="clade-leaf"> <p><span typeof="mw:File"><a href="/wiki/File:Cynotherium_sardous_restoration.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/73/Cynotherium_sardous_restoration.jpg/50px-Cynotherium_sardous_restoration.jpg" decoding="async" width="50" height="26" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/73/Cynotherium_sardous_restoration.jpg/75px-Cynotherium_sardous_restoration.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/73/Cynotherium_sardous_restoration.jpg/100px-Cynotherium_sardous_restoration.jpg 2x" data-file-width="1984" data-file-height="1048" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"><i>†<a href="/wiki/Xenocyon" title="Xenocyon">Xenocyon</a></i> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"><i><a href="/wiki/Cuon" class="mw-redirect" title="Cuon">Cuon</a></i>&#160;(Dhole) </td> <td rowspan="2" class="clade-leaf"> <p><span typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_XLI).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/7b/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XLI%29.jpg/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XLI%29.jpg" decoding="async" width="50" height="32" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/7b/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XLI%29.jpg/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XLI%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/7b/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XLI%29.jpg/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XLI%29.jpg 2x" data-file-width="471" data-file-height="302" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"><i><a href="/wiki/Lycaon_(genus)" title="Lycaon (genus)">Lycaon</a></i>&#160;(African&#160;hunting&#160;dog) </td> <td rowspan="2" class="clade-leaf"> <p><span typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_XLIV).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/66/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XLIV%29.jpg/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XLIV%29.jpg" decoding="async" width="50" height="31" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/66/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XLIV%29.jpg/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XLIV%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/66/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XLIV%29.jpg/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XLIV%29.jpg 2x" data-file-width="445" data-file-height="277" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td style="text-align: left; font-size: 88%; border: 0; padding: 0;">Previously proposed evolution and branching from <i><a href="/wiki/Eucyon" title="Eucyon">Eucyon</a></i> towards the wolf (2008). <sup id="cite_ref-FOOTNOTEWangTedford2008148_23-2" class="reference"><a href="#cite_note-FOOTNOTEWangTedford2008148-23"><span class="cite-bracket">&#91;</span>23<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-tedford2009_11-5" class="reference"><a href="#cite_note-tedford2009-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: p181">&#58;&#8202;p181&#8202;</span></sup> </td></tr></tbody></table> <p><a href="/wiki/Xiaoming_Wang_(paleontologist)" title="Xiaoming Wang (paleontologist)">Wang</a> and <a href="/wiki/Richard_H._Tedford" title="Richard H. Tedford">Tedford</a> proposed that the genus <i>Canis</i> was the descendant of the coyote-like <i><a href="/wiki/Eucyon" title="Eucyon">Eucyon</a> davisi</i>, and its remains first appeared in the <a href="/wiki/Miocene" title="Miocene">Miocene</a> (6 million YBP) in south-western USA and Mexico. By the <a href="/wiki/Pliocene" title="Pliocene">Pliocene</a> (5 million YBP), the larger <i><a href="/wiki/Canis_lepophagus" title="Canis lepophagus">Canis lepophagus</a></i> appeared in the same region and by the <a href="/wiki/Early_Pleistocene" title="Early Pleistocene">Early Pleistocene</a> (1 million YBP) <i>Canis latrans</i> (the <a href="/wiki/Coyote" title="Coyote">coyote</a>) was in existence. They proposed that the progression from <i>Eucyon davisi</i> to <i>C lepophagus</i> to the coyote was linear evolution.<sup id="cite_ref-FOOTNOTEWangTedford200858_22-1" class="reference"><a href="#cite_note-FOOTNOTEWangTedford200858-22"><span class="cite-bracket">&#91;</span>22<span class="cite-bracket">&#93;</span></a></sup> Additionally, <i>C. edwardii</i>, <i>C. latrans</i> and <i>C. aureus</i> form together a small clade and because <i>C. edwardii</i> appeared earliest spanning the mid-Blancan (late Pliocene) to the close of the Irvingtonian (late Pleistocene) it is proposed as the ancestor.<sup id="cite_ref-tedford2009_11-6" class="reference"><a href="#cite_note-tedford2009-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: p175, 180">&#58;&#8202;p175,&#8202;180&#8202;</span></sup> </p><p>Nowak and Tedford also believed that it was possible for <i>C. lupus</i> to have been derived from a Miocene or Pliocene canid line that preceded and was separate from <i>C. lepophagus</i>.<sup id="cite_ref-nowak1979_28-5" class="reference"><a href="#cite_note-nowak1979-28"><span class="cite-bracket">&#91;</span>28<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-tedford1996_52-0" class="reference"><a href="#cite_note-tedford1996-52"><span class="cite-bracket">&#91;</span>52<span class="cite-bracket">&#93;</span></a></sup> Based on morphology from China, the <a href="/wiki/Pliocene" title="Pliocene">Pliocene</a> wolf <i>C. chihliensis</i> may have been the ancestor for both <i>C. armbrusteri</i> and <i>C. lupus</i> before their migration into North America.<sup id="cite_ref-FOOTNOTEWangTedford2008148_23-3" class="reference"><a href="#cite_note-FOOTNOTEWangTedford2008148-23"><span class="cite-bracket">&#91;</span>23<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-tedford2009_11-7" class="reference"><a href="#cite_note-tedford2009-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: p181">&#58;&#8202;p181&#8202;</span></sup> <i>C. chihliensis</i> appears to be more primitive and smaller than <i>C. lupus</i>, and measurements of its skull and teeth are similar to <i>C. lupus</i> but those of its postcranial elements are smaller.<sup id="cite_ref-tong2012_53-0" class="reference"><a href="#cite_note-tong2012-53"><span class="cite-bracket">&#91;</span>53<span class="cite-bracket">&#93;</span></a></sup> <i>C. armbrusteri</i> appeared in North America in the <a href="/wiki/Middle_Pleistocene" class="mw-redirect" title="Middle Pleistocene">Middle Pleistocene</a> and is a wolf-like form larger than any <i>Canis</i> at that time.<sup id="cite_ref-nowak1979_28-6" class="reference"><a href="#cite_note-nowak1979-28"><span class="cite-bracket">&#91;</span>28<span class="cite-bracket">&#93;</span></a></sup> At the end of the most recent glacial retreat during the past 30,000 years, warming melted the glacial barriers across northern Canada allowing arctic mammals to extend their range into mid-latitude North America, including elk, caribou, bison, and the grey wolf.<sup id="cite_ref-FOOTNOTEWangTedford200861_54-0" class="reference"><a href="#cite_note-FOOTNOTEWangTedford200861-54"><span class="cite-bracket">&#91;</span>54<span class="cite-bracket">&#93;</span></a></sup> </p><p>In Eurasia during the Middle Pleistocene, <i>C. falconeri</i> gave rise to the hypercarnivore genus <i><a href="/wiki/Xenocyon" title="Xenocyon">Xenocyon</a></i>, which then gave rise to genus <i><a href="/wiki/Cuon" class="mw-redirect" title="Cuon">Cuon</a></i> (the dhole) and genus <i><a href="/wiki/Lycaon_(genus)" title="Lycaon (genus)">Lycaon</a></i> (the African hunting dog).<sup id="cite_ref-FOOTNOTEWangTedford2008105_55-0" class="reference"><a href="#cite_note-FOOTNOTEWangTedford2008105-55"><span class="cite-bracket">&#91;</span>55<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-FOOTNOTEWangTedford2008149_56-0" class="reference"><a href="#cite_note-FOOTNOTEWangTedford2008149-56"><span class="cite-bracket">&#91;</span>56<span class="cite-bracket">&#93;</span></a></sup> Just before the appearance of <i>C. dirus</i>, North America was invaded by genus <i>Xenocyon</i> that was as large as <i>C. dirus</i> and more hypercarnivorous. The fossil record shows them as rare and it is assumed that they could not compete with the newly derived <i>C. dirus</i>.<sup id="cite_ref-FOOTNOTEWangTedford200860_57-0" class="reference"><a href="#cite_note-FOOTNOTEWangTedford200860-57"><span class="cite-bracket">&#91;</span>57<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Canis_antonii"><i>Canis antonii</i></h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=6" title="Edit section: Canis antonii"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The large wolf <i>C. antonii</i> from late Pliocene to early Pleistocene China was assessed as being a variation within <i>C. chihliensis</i>,<sup id="cite_ref-tedford2009_11-8" class="reference"><a href="#cite_note-tedford2009-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: p197">&#58;&#8202;p197&#8202;</span></sup> and the large wolf <i>C. falconeri</i> occurred abruptly in Europe in the Early Pleistocene, perhaps representing a westward extension of <i>C. antonii</i>.<sup id="cite_ref-tedford2009_11-9" class="reference"><a href="#cite_note-tedford2009-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: p181">&#58;&#8202;p181&#8202;</span></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Canis_borjgali"><i>Canis borjgali</i></h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=7" title="Edit section: Canis borjgali"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In 2020, researchers named a new species <i>C. borjgali</i> that was found in <a href="/wiki/Dmanisi" title="Dmanisi">Dmanisi</a>, Georgia in a site dated 1.8—1.75 million years ago.<sup id="cite_ref-sotnikova2010_58-0" class="reference"><a href="#cite_note-sotnikova2010-58"><span class="cite-bracket">&#91;</span>58<span class="cite-bracket">&#93;</span></a></sup> This specimen did not show the peculiarities of <i>C. etruscus</i> but appears to be closer to a primitive form of <i>C. mosbachensis</i> and is proposed as the ancestor of <i>C. mosbachensis</i>.<sup id="cite_ref-bartolini2020_59-0" class="reference"><a href="#cite_note-bartolini2020-59"><span class="cite-bracket">&#91;</span>59<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-sotnikova2010_58-1" class="reference"><a href="#cite_note-sotnikova2010-58"><span class="cite-bracket">&#91;</span>58<span class="cite-bracket">&#93;</span></a></sup> The wolf <i>C. borjgali</i> is probably the ancestor of the wolf-like <a href="/wiki/Crown_clade" class="mw-redirect" title="Crown clade">crown-clade</a> species <i><a href="/wiki/Wolf" title="Wolf">C. lupus</a></i>, <i><a href="/wiki/Coyote" title="Coyote">C. latrans</a></i>, and <i><a href="/wiki/African_golden_wolf" class="mw-redirect" title="African golden wolf">C. lupaster</a></i>.<sup id="cite_ref-bartolini2020_59-1" class="reference"><a href="#cite_note-bartolini2020-59"><span class="cite-bracket">&#91;</span>59<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Canis_mosbachensis"><i>Canis mosbachensis</i></h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=8" title="Edit section: Canis mosbachensis"><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:Canis_mosbachensis_Wikipedia_Juandertal.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f7/Canis_mosbachensis_Wikipedia_Juandertal.jpg/220px-Canis_mosbachensis_Wikipedia_Juandertal.jpg" decoding="async" width="220" height="143" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f7/Canis_mosbachensis_Wikipedia_Juandertal.jpg/330px-Canis_mosbachensis_Wikipedia_Juandertal.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f7/Canis_mosbachensis_Wikipedia_Juandertal.jpg/440px-Canis_mosbachensis_Wikipedia_Juandertal.jpg 2x" data-file-width="1265" data-file-height="822" /></a><figcaption>Life restoration of <i>C. mosbachensis</i></figcaption></figure> <p><i><a href="/wiki/Canis_mosbachensis" title="Canis mosbachensis">Canis mosbachensis</a></i>, sometimes known as the Mosbach wolf, is an extinct small wolf that once inhabited Eurasia from the <a href="/wiki/Middle_Pleistocene" class="mw-redirect" title="Middle Pleistocene">Middle</a> to <a href="/wiki/Late_Pleistocene" title="Late Pleistocene">Late Pleistocene</a>.<sup id="cite_ref-kurten1968_60-0" class="reference"><a href="#cite_note-kurten1968-60"><span class="cite-bracket">&#91;</span>60<span class="cite-bracket">&#93;</span></a></sup> The phylogenetic descent of the extant wolf <i>C.&#160;lupus</i> from <i><a href="/wiki/Canis_etruscus" title="Canis etruscus">C.&#160;etruscus</a></i> through <i>C.&#160;mosbachensis</i> is widely accepted.<sup id="cite_ref-kurten1968_60-1" class="reference"><a href="#cite_note-kurten1968-60"><span class="cite-bracket">&#91;</span>60<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-nowak2003_27-13" class="reference"><a href="#cite_note-nowak2003-27"><span class="cite-bracket">&#91;</span>27<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: 239–245">&#58;&#8202;239–245&#8202;</span></sup> In 2010, a study found that the diversity of the <i>Canis</i> group decreased by the end of the <a href="/wiki/Early_Pleistocene" title="Early Pleistocene">Early Pleistocene</a> to <a href="/wiki/Middle_Pleistocene" class="mw-redirect" title="Middle Pleistocene">Middle Pleistocene</a> and was limited in Eurasia to the small wolves of the <i>C.&#160;mosbachensis–C.&#160;variabilis</i> group that were a comparable size to the extant <a href="/wiki/Indian_wolf" title="Indian wolf">Indian wolf</a> (<i>Canis lupus pallipes</i>), and the large hypercarnivorous <a href="/wiki/Xenocyon#Canis_(Xenocyon)_lycaonoides" title="Xenocyon"><i>Canis</i> (<i>Xenocyon</i>) <i>lycanoides</i></a> that was comparable in size to extant northern grey wolves.<sup id="cite_ref-sotnikova2010_58-2" class="reference"><a href="#cite_note-sotnikova2010-58"><span class="cite-bracket">&#91;</span>58<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Canis_lupus"><i>Canis lupus</i></h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=9" title="Edit section: Canis lupus"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The earliest <i>Canis lupus</i> specimen was a fossil tooth discovered at <a href="/wiki/Old_Crow,_Yukon" title="Old Crow, Yukon">Old Crow, Yukon</a>, Canada. The specimen was found in sediment dated 1 million YBP,<sup id="cite_ref-tedford2009_11-10" class="reference"><a href="#cite_note-tedford2009-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup> however the geological attribution of this sediment is questioned.<sup id="cite_ref-tedford2009_11-11" class="reference"><a href="#cite_note-tedford2009-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-westgate2017_61-0" class="reference"><a href="#cite_note-westgate2017-61"><span class="cite-bracket">&#91;</span>61<span class="cite-bracket">&#93;</span></a></sup> Slightly younger specimens were discovered at Cripple Creek Sump, <a href="/wiki/Fairbanks" class="mw-redirect" title="Fairbanks">Fairbanks</a>, Alaska, in strata dated 810,000 YBP. Both discoveries point to an origin of these wolves in east <a href="/wiki/Beringia" title="Beringia">Beringia</a> during the <a href="/wiki/Middle_Pleistocene" class="mw-redirect" title="Middle Pleistocene">Middle Pleistocene</a>.<sup id="cite_ref-tedford2009_11-12" class="reference"><a href="#cite_note-tedford2009-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup> </p><p>In France, the subspecies <i>C. l. lunellensis</i> Bonifay, 1971<sup id="cite_ref-bonifay1971_62-0" class="reference"><a href="#cite_note-bonifay1971-62"><span class="cite-bracket">&#91;</span>62<span class="cite-bracket">&#93;</span></a></sup> discovered at <a href="/wiki/Lunel-Viel" title="Lunel-Viel">Lunel-Viel</a>, <a href="/wiki/H%C3%A9rault" title="Hérault">Hérault</a> dated 400–350,000 YBP, <i>C. l. santenaisiensis</i> Argant, 1991<sup id="cite_ref-argant1991_63-0" class="reference"><a href="#cite_note-argant1991-63"><span class="cite-bracket">&#91;</span>63<span class="cite-bracket">&#93;</span></a></sup> from <a href="/wiki/Santenay,_C%C3%B4te-d%27Or" title="Santenay, Côte-d&#39;Or">Santenay, Côte-d'Or</a> dated to 200,000 YBP, and <i><a href="/wiki/Pleistocene_wolf#Canis_lupus_maximus" title="Pleistocene wolf">C. lupus maximus</a></i> Boudadi-Maligne, 2012<sup id="cite_ref-boudadi2012_64-0" class="reference"><a href="#cite_note-boudadi2012-64"><span class="cite-bracket">&#91;</span>64<span class="cite-bracket">&#93;</span></a></sup> from Jaurens cave, <a href="/wiki/Nespouls" title="Nespouls">Nespouls</a>, <a href="/wiki/Corr%C3%A8ze" title="Corrèze">Corrèze</a> dated 31,000 YBP, show a progressive increase in size and are proposed to be <a href="/wiki/Chronospecies" title="Chronospecies">chrono-subspecies</a>.<sup id="cite_ref-brugal2011_65-0" class="reference"><a href="#cite_note-brugal2011-65"><span class="cite-bracket">&#91;</span>65<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-sardella2014_17-1" class="reference"><a href="#cite_note-sardella2014-17"><span class="cite-bracket">&#91;</span>17<span class="cite-bracket">&#93;</span></a></sup> In Italy, the earliest <i>Canis lupus</i> specimens were found at La Polledrara di Cecanibbio, 20&#160;km north-west of Rome in strata dated 340,000–320,000 YBP.<sup id="cite_ref-anzidei2012_66-0" class="reference"><a href="#cite_note-anzidei2012-66"><span class="cite-bracket">&#91;</span>66<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-gliozzi1997_67-0" class="reference"><a href="#cite_note-gliozzi1997-67"><span class="cite-bracket">&#91;</span>67<span class="cite-bracket">&#93;</span></a></sup> In 2017, a study found that the dimensions of the upper and lower carnassial teeth of the early Holocene Italian wolf are close to those of <i>C. l. maximus</i>. Fluctuations in the size of <i>C. lupus</i> carnassial teeth correlate with the spread of megafauna. The Italian wolf underwent a reduction in body size with the loss of the red deer in Italy during the Renaissance.<sup id="cite_ref-sardella2014_17-2" class="reference"><a href="#cite_note-sardella2014-17"><span class="cite-bracket">&#91;</span>17<span class="cite-bracket">&#93;</span></a></sup> The proposed lineage is: </p> <style data-mw-deduplicate="TemplateStyles:r1244412712">.mw-parser-output .templatequote{overflow:hidden;margin:1em 0;padding:0 32px}.mw-parser-output .templatequotecite{line-height:1.5em;text-align:left;margin-top:0}@media(min-width:500px){.mw-parser-output .templatequotecite{padding-left:1.6em}}</style><blockquote class="templatequote"><p><i>C. etruscus → C. mosbachensis → C. l. lunellensis → C. l. santenaisiensis → C. l. maximus → C. l. lupus</i><sup id="cite_ref-sardella2014_17-3" class="reference"><a href="#cite_note-sardella2014-17"><span class="cite-bracket">&#91;</span>17<span class="cite-bracket">&#93;</span></a></sup></p></blockquote> <div class="mw-heading mw-heading4"><h4 id="Canis_lupus_bohemica"><i>Canis lupus bohemica</i></h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=10" title="Edit section: Canis lupus bohemica"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In 2022, a new subspecies <i>Canis lupus bohemica</i> was <a href="/wiki/Taxonomy_(biology)#Taxonomic_descriptions" title="Taxonomy (biology)">taxonomically described</a> after having been discovered in the Bat Cave system located near <a href="/wiki/Srbsko" title="Srbsko">Srbsko</a>, Central Bohemia, Czech Republic. The Bohemian wolf is an extinct short-legged wolf that first appeared 800,000 years ago (MIS 20, the Rhumian Interglacial of the early <a href="/wiki/Cromerian_stage" class="mw-redirect" title="Cromerian stage">Cromerian stage</a>, Middle Pleistocene) and once inhabited what was part of the mammoth steppe. It is proposed as the ancestor of <i>Canis lupus mosbachensis</i>. In comparison, <i>C. etruscus</i> appears to be the ancestor of the <a href="/wiki/Golden_jackal" title="Golden jackal">Afro-Eurasian jackal</a>.<sup id="cite_ref-Diedrich2022_68-0" class="reference"><a href="#cite_note-Diedrich2022-68"><span class="cite-bracket">&#91;</span>68<span class="cite-bracket">&#93;</span></a></sup> </p><p>In Hungary in 1969, a tooth (the <a href="/wiki/Premolar" title="Premolar">premolar</a> of the <a href="/wiki/Maxilla" title="Maxilla">Maxilla</a>) was found which dated to the Middle Pleistocene, and was assessed as being midway between that of <i>Canis mosbachensis</i> and the <a href="/wiki/Cave_wolf" title="Cave wolf">cave wolf</a> <i>Canis lupus spelaeus</i>, but leaning towards <i>C.l. spelaeus</i>. <sup id="cite_ref-Janossy2012_69-0" class="reference"><a href="#cite_note-Janossy2012-69"><span class="cite-bracket">&#91;</span>69<span class="cite-bracket">&#93;</span></a></sup> During the late Middle Pleistocene around 600,000 years ago, the Bohemian wolf diversified into two wolf lineages that specialized for different environmental and climatic conditions. One was a southern interglacial (warm climate) grey wolf of Europe which was to become the Mosbach wolf, and the other a northern glacial white wolf of Eurasia which was to become <i>C. l. spelaeus</i>.<sup id="cite_ref-Diedrich2022_68-1" class="reference"><a href="#cite_note-Diedrich2022-68"><span class="cite-bracket">&#91;</span>68<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Canis_c.f._familiaris_-_Paleolithic_dog"><i>Canis c.f. familiaris</i> - Paleolithic dog</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=11" title="Edit section: Canis c.f. familiaris - Paleolithic dog"><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/Paleolithic_dog" title="Paleolithic dog">Paleolithic dog</a></div> <p>There are a number of recently discovered specimens which are proposed as being Paleolithic dogs, however their taxonomy is debated. These have been found in either Europe or Siberia and date 40,000-17,000 YBP. They include <a href="/wiki/Hohle_Fels" title="Hohle Fels">Hohle Fels</a> in Germany, <a href="/wiki/Goyet_Caves" title="Goyet Caves">Goyet Caves</a> in Belgium, <a href="/wiki/P%C5%99edmost%C3%AD_u_P%C5%99erova_(archeology)" class="mw-redirect" title="Předmostí u Přerova (archeology)">Predmosti</a> in the Czech Republic, and four sites in Russia: Razboinichya Cave, <a href="/wiki/Kostyonki_(palaeolithic_site)" class="mw-redirect" title="Kostyonki (palaeolithic site)">Kostyonki-8</a>, Ulakhan Sular, and Eliseevichi 1. Paw-prints from <a href="/wiki/Chauvet_Cave" title="Chauvet Cave">Chauvet Cave</a> in France dated 26,000 YBP are suggested as being those of a dog, however these have been challenged as being left by a wolf.<sup id="cite_ref-Thalmann2018_70-0" class="reference"><a href="#cite_note-Thalmann2018-70"><span class="cite-bracket">&#91;</span>70<span class="cite-bracket">&#93;</span></a></sup> Paleolithic dogs were directly associated with human hunting camps in Europe over 30,000 (YBP) and it is proposed that they were domesticated. They are also proposed to be either a proto-dog and the ancestor of the domestic dog or a type of wolf unknown to science.<sup id="cite_ref-germonpre2009_71-0" class="reference"><a href="#cite_note-germonpre2009-71"><span class="cite-bracket">&#91;</span>71<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Canis_familiaris_(domestic_dog)"><span id="Canis_familiaris_.28domestic_dog.29"></span><i>Canis familiaris</i> (domestic dog)</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=12" title="Edit section: Canis familiaris (domestic dog)"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a class="mw-selflink-fragment" href="#Wolf_population_differences">Evolution of the wolf §&#160;Wolf population differences</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Skull_of_bulldog_-_Keulemans.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/52/Skull_of_bulldog_-_Keulemans.png/220px-Skull_of_bulldog_-_Keulemans.png" decoding="async" width="220" height="169" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/52/Skull_of_bulldog_-_Keulemans.png/330px-Skull_of_bulldog_-_Keulemans.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/52/Skull_of_bulldog_-_Keulemans.png/440px-Skull_of_bulldog_-_Keulemans.png 2x" data-file-width="999" data-file-height="769" /></a><figcaption>Bulldog skull – sketch</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Eurowolfskull.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/05/Eurowolfskull.jpg/220px-Eurowolfskull.jpg" decoding="async" width="220" height="256" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/05/Eurowolfskull.jpg/330px-Eurowolfskull.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/05/Eurowolfskull.jpg/440px-Eurowolfskull.jpg 2x" data-file-width="505" data-file-height="587" /></a><figcaption>European wolf skull – sketch</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Skull_of_a_dog_diagram2.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/7d/Skull_of_a_dog_diagram2.svg/220px-Skull_of_a_dog_diagram2.svg.png" decoding="async" width="220" height="156" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/7d/Skull_of_a_dog_diagram2.svg/330px-Skull_of_a_dog_diagram2.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/7d/Skull_of_a_dog_diagram2.svg/440px-Skull_of_a_dog_diagram2.svg.png 2x" data-file-width="1052" data-file-height="744" /></a><figcaption><i>Canis lupus</i> skull: 1 – maxilla, 2 – frontal, 3 – lachrymal, 4 – palatine, 5 – jugal, 6 – nasal, 7 – premaxilla, 8 – parietal, 9 – interparietal, 10 – squamosal, 11 – occipital, 12 – mandible</figcaption></figure> <p>In 2002, a study was undertaken into the fossil skulls of two large canids that had been found buried within meters of the doorway of what was once a mammoth-bone hut at the Eliseevichi-I <a href="/wiki/Upper_Paleolithic" title="Upper Paleolithic">Upper Paleolithic</a> site in the <a href="/wiki/Bryansk_Oblast" title="Bryansk Oblast">Bryansk Region</a> on the Russian Plain, and using an accepted morphologically based definition of domestication declared them to be "Ice Age dogs". The carbon dating gave a calendar-year age estimate that ranged between 16,945 and 13,905 YBP.<sup id="cite_ref-sablin2002_72-0" class="reference"><a href="#cite_note-sablin2002-72"><span class="cite-bracket">&#91;</span>72<span class="cite-bracket">&#93;</span></a></sup> In 2013, a study looked at one of these skulls and its mitochondrial DNA sequence was identified as <i>Canis familiaris</i>.<sup id="cite_ref-thalmann2013_73-0" class="reference"><a href="#cite_note-thalmann2013-73"><span class="cite-bracket">&#91;</span>73<span class="cite-bracket">&#93;</span></a></sup> </p><p>In 2015, a zooarchaeologist stated that "In terms of phenotypes, dogs and wolves are fundamentally different animals."<sup id="cite_ref-morey2015_74-0" class="reference"><a href="#cite_note-morey2015-74"><span class="cite-bracket">&#91;</span>74<span class="cite-bracket">&#93;</span></a></sup> </p><p>In 1986, a study of skull morphology found that the domestic dog is morphologically distinct from all other canids except the wolf-like canids. "The difference in size and proportion between some breeds are as great as those between any wild genera, but all dogs are clearly members of the same species."<sup id="cite_ref-wayne1986_75-0" class="reference"><a href="#cite_note-wayne1986-75"><span class="cite-bracket">&#91;</span>75<span class="cite-bracket">&#93;</span></a></sup> In 2010, a study of dog skull shape compared to extant <a href="/wiki/Carnivorans" class="mw-redirect" title="Carnivorans">carnivorans</a> proposed that "The greatest shape distances between dog breeds clearly surpass the maximum divergence between species in the Carnivora. Moreover, domestic dogs occupy a range of novel shapes outside the domain of wild carnivorans."<sup id="cite_ref-drake2010_76-0" class="reference"><a href="#cite_note-drake2010-76"><span class="cite-bracket">&#91;</span>76<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Comparison_with_modern_wolves">Comparison with modern wolves</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=13" title="Edit section: Comparison with modern wolves"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The domestic dog compared to the modern wolf shows the greatest variation in the size and shape of the skull (Evans 1979) that range from 7 to 28&#160;cm in length (McGreevy 2004). Wolves are <a href="/wiki/Dolichocephaly" title="Dolichocephaly">dolichocephalic</a> (long skulled) but not as extreme as some breeds of such as greyhounds and Russian wolfhounds (McGreevy 2004). Canine <a href="/wiki/Brachycephaly" title="Brachycephaly">brachycephaly</a> (short-skulledness) is found only in domestic dogs and is related to <a href="/wiki/Paedomorphosis" class="mw-redirect" title="Paedomorphosis">paedomorphosis</a> (Goodwin 1997). Puppies are born with short snouts, with the longer skull of dolichocephalic dogs emerging in later development (Coppinger 1995). Other differences in head shape between brachycephalic and dolichocephalic dogs include changes in the craniofacial angle (angle between the <a href="/wiki/Basilar_plexus" class="mw-redirect" title="Basilar plexus">basilar axis</a> and <a href="/wiki/Hard_palate" title="Hard palate">hard palate</a>) (Regodón 1993), morphology of the <a href="/wiki/Temporomandibular_joint" title="Temporomandibular joint">temporomandibular joint</a> (Dickie 2001), and radiographic anatomy of the <a href="/wiki/Cribriform_plate" title="Cribriform plate">cribriform plate</a> (Schwarz 2000).<sup id="cite_ref-roberts2010_77-0" class="reference"><a href="#cite_note-roberts2010-77"><span class="cite-bracket">&#91;</span>77<span class="cite-bracket">&#93;</span></a></sup> </p><p>Nowak indicated that orbital angle of the eye socket is an important characteristic defining the difference between the dog and the wolf, with the wolf having the lower angle. Nowak compared the orbital angles of four North American <a href="/wiki/Canis" title="Canis">canines</a> (including the Indian dog) and produced the following values in degrees: coyote-42.8, wolf-42.8, dog-52.9 dire wolf-53.1. The orbital angle of the eye socket was clearly larger in the dog than in the coyote and the wolf; why it was almost the same as that of the dire wolf was not commented on.<sup id="cite_ref-nowak1979_28-7" class="reference"><a href="#cite_note-nowak1979-28"><span class="cite-bracket">&#91;</span>28<span class="cite-bracket">&#93;</span></a></sup> </p><p>Many authors have concluded that compared to the adult extant wolf, the adult domestic dog has a relatively reduced rostrum (front part of the skull), an elevated <a href="/wiki/Frontal_bone" title="Frontal bone">frontal bone</a>, a wider <a href="/wiki/Palate" title="Palate">palate</a>, a broader <a href="/wiki/Cranium" class="mw-redirect" title="Cranium">cranium</a>, and smaller teeth (Hildebrand1954; Clutton-Brock, Corbet &amp; Hills 1976; Olsen 1985; Wayne 1986; Hemmer 1990; Morey 1990). Other authors have disagreed and have stated that these traits can overlap and vary within the two (Crockford 1999; Harrison 1973). Wolf cubs have similar relative skull proportions as adult dogs and this was proposed as evidence that the domestic dog is a <a href="/wiki/Neoteny" title="Neoteny">neotenic wolf</a>. This was proposed to be due to either human selection for juvenile appearance or due to a <a href="/wiki/Pleiotropy" title="Pleiotropy">pleiotropic</a> effect as a result of selection for juvenile behavior (Clutton-Brock 1977; Belyaev 1979; Wayne 1986; Coppinger and Schneider 1995). Wayne (1986) concluded that his dog samples did not have significant relative shortening of the rostrum compared to wolves, calling this identifying feature into question.<sup id="cite_ref-koler2002_78-0" class="reference"><a href="#cite_note-koler2002-78"><span class="cite-bracket">&#91;</span>78<span class="cite-bracket">&#93;</span></a></sup> A 2004 study that used 310 wolf skulls and over 700 dog skulls representing 100 breeds concluded that the evolution of dog skulls can generally not be described by heterochronic processes such as neoteny although some pedomorphic dog breeds have skulls that resemble the skulls of juvenile wolves.<sup id="cite_ref-drake2004_79-0" class="reference"><a href="#cite_note-drake2004-79"><span class="cite-bracket">&#91;</span>79<span class="cite-bracket">&#93;</span></a></sup> "Dogs are not paedomorphic wolves."<sup id="cite_ref-drake2011_80-0" class="reference"><a href="#cite_note-drake2011-80"><span class="cite-bracket">&#91;</span>80<span class="cite-bracket">&#93;</span></a></sup> </p><p>Compared to the wolf, dog dentition is relatively less robust (Olsen 1985; Hemmer 1990), which is proposed to be due to the relaxation of natural selection when wolves became commensal scavengers, or to artificial selection (Olsen 1985; Clutton-Brock 1995). However, Kieser and Groeneveld (1992) compared the mandibulo-dental measurements of jackals (<i>C. adustus, C. mesomelas</i>) and Cape foxes (<i>Vulpes chama</i>) to equivalent-sized dogs and found that the canines of these other canids tended to be slightly smaller and their second molars larger compared to dogs, otherwise the proportions were essentially the same in all species. They concluded that "...the teeth of canids appear to have evolved in concert with one another and relatively independently of differences in dimorphism, size or functional demands". This calls into question the assumption that dog teeth are relatively small due to recent selection, suggesting that dog dentition is plesiomorphic from an ancestor that was smaller than the wolf.<sup id="cite_ref-koler2002_78-1" class="reference"><a href="#cite_note-koler2002-78"><span class="cite-bracket">&#91;</span>78<span class="cite-bracket">&#93;</span></a></sup> </p><p>The reduced body size of the early dog compared to a wolf is thought due to niche selection (Olsen 1985; Morey 1992; Coppinger &amp; Coppinger 2001). Morey (1992:199) states that "Results...are consistent with a hypothesis that early domestic dogs are evolutionary paedomorphs, products of strong selection for ontogenetically channeled size reduction and alterations of reproductive timing associated with the new domestic way of life."<sup id="cite_ref-koler2002_78-2" class="reference"><a href="#cite_note-koler2002-78"><span class="cite-bracket">&#91;</span>78<span class="cite-bracket">&#93;</span></a></sup> However, in an domestication experiment the domesticated foxes remained the same size as unselected foxes (Trutt 1999:167).<sup id="cite_ref-morey2015_74-1" class="reference"><a href="#cite_note-morey2015-74"><span class="cite-bracket">&#91;</span>74<span class="cite-bracket">&#93;</span></a></sup> </p><p>Wayne (1986) concluded that the dog is closer in skull morphology to <i>C. latrans, C. aureus, C. adustus, C. mesomelas, Cuon alpinus</i> and <i>Lycaon pictus</i> than to the wolf. Dahr (1942) concluded that the shape of the dog brain case is closer to that of the coyote than to that of the wolf. Manwell and Baker (1983) reviewed Dahr's work with the addition of dental data for canids and concluded that the dog ancestor was probably within the range of 13.6–20.5&#160;kg, which is smaller than the range 27–54&#160;kg for extant wolves (Mech 1970) and is comparable with the <a href="/wiki/Dingo" title="Dingo">Dingo</a>.<sup id="cite_ref-koler2002_78-3" class="reference"><a href="#cite_note-koler2002-78"><span class="cite-bracket">&#91;</span>78<span class="cite-bracket">&#93;</span></a></sup> </p><p>The auditory <a href="/wiki/Tympanic_part_of_the_temporal_bone" title="Tympanic part of the temporal bone">bulla</a> of the dog is relatively smaller and flatter than that of the wolf (Harrison 1973; Clutton-Brock, Corbet &amp; Hill 1976; Nowak 1979; Olsen 1985; Wayne 1986), which is proposed to be due to relaxed selection under domestication as the dog no longer required the acute hearing of the wolf. However, bulla shape has been shown to facilitate increased sensitivity to specific frequencies but shape and size may not be correlated with acuity (Ewer 1973). Therefore, the observed difference could be that the dog bulla has retained its ancestral shape.<sup id="cite_ref-koler2002_78-4" class="reference"><a href="#cite_note-koler2002-78"><span class="cite-bracket">&#91;</span>78<span class="cite-bracket">&#93;</span></a></sup> </p><p>The ventral edge of the dog's horizontal <a href="/wiki/Ramus_of_the_mandible" class="mw-redirect" title="Ramus of the mandible">ramus of the mandible</a> has a convex curve that does not exist in the wolf (Olsen 1985; Clutton-Brock 1995), and no discussion of this difference could be found in the literature. However, Biknevicius and Van Valkenburgh (1997) noticed that the horizontal ramus of bone-processing predators is thicker dorso-ventrally at the point <a href="/wiki/Anatomical_terms_of_location#caudal" title="Anatomical terms of location">caudal</a> to the site of bone processing. This thickening may have been a function for niche adaptation by the dog's ancestor.<sup id="cite_ref-koler2002_78-5" class="reference"><a href="#cite_note-koler2002-78"><span class="cite-bracket">&#91;</span>78<span class="cite-bracket">&#93;</span></a></sup> </p><p>A description of the superficial brain morphology of jackals (<i>C. mesomelas, C. aureus</i>), coyotes (<i>C. latrans</i>), wolves (<i>C. lupus, C. rufus</i>), and dogs indicated that the <a href="/wiki/Cerebellum" title="Cerebellum">cerebellum</a> of the dog closely approximates that of the coyote, which is closely aligned with the jackals, and that the wolves show numerous brain traits distinct from the other species (Atkins and Dillon 1971). Wolves also have serological and biochemical traits distinct from dogs (Leone and Wiens 1956; Lauer, Kuyt &amp; Baker 1969).<sup id="cite_ref-koler2002_78-6" class="reference"><a href="#cite_note-koler2002-78"><span class="cite-bracket">&#91;</span>78<span class="cite-bracket">&#93;</span></a></sup> </p><p>The tails of domestic dogs tend to curl upwards, which is not found in other canid.<sup id="cite_ref-Clutton-Brock1995_81-0" class="reference"><a href="#cite_note-Clutton-Brock1995-81"><span class="cite-bracket">&#91;</span>81<span class="cite-bracket">&#93;</span></a></sup> Dogs possess a domed forehead, which is easy to see in a profile view with a bulge above the <a href="/wiki/Orbit_(anatomy)" title="Orbit (anatomy)">orbit</a> (the bony socket) of the eye. The <a href="/wiki/Temporalis_muscle" title="Temporalis muscle">temporalis muscle</a> that closes the jaws is more robust in wolves.<sup id="cite_ref-FOOTNOTEWangTedford2008158_82-0" class="reference"><a href="#cite_note-FOOTNOTEWangTedford2008158-82"><span class="cite-bracket">&#91;</span>82<span class="cite-bracket">&#93;</span></a></sup> Compared with equally sized wolves, dogs possess 20% smaller skulls, and compared with equally weighted wolves dogs possess 20% smaller brains.<sup id="cite_ref-serpell1995_83-0" class="reference"><a href="#cite_note-serpell1995-83"><span class="cite-bracket">&#91;</span>83<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: 47">&#58;&#8202;47&#8202;</span></sup> The <a href="/wiki/Auditory_bulla" class="mw-redirect" title="Auditory bulla">tympanic bullae</a> are large, convex, and almost spherical in wolves, while the bullae of dogs are smaller, compressed, and slightly crumpled.<sup id="cite_ref-mech1974_84-0" class="reference"><a href="#cite_note-mech1974-84"><span class="cite-bracket">&#91;</span>84<span class="cite-bracket">&#93;</span></a></sup> Dogs have a distinctive "stop" between the forehead and nose.<sup id="cite_ref-85" class="reference"><a href="#cite_note-85"><span class="cite-bracket">&#91;</span>85<span class="cite-bracket">&#93;</span></a></sup> Domestic dogs are distinguishable from wolves by starch gel <a href="/wiki/Electrophoresis" title="Electrophoresis">electrophoresis</a> of red blood cell acid <a href="/wiki/Phosphatase" title="Phosphatase">phosphatase</a>.<sup id="cite_ref-86" class="reference"><a href="#cite_note-86"><span class="cite-bracket">&#91;</span>86<span class="cite-bracket">&#93;</span></a></sup> </p><p>During the Last Glacial Maximum, there was greater wolf <a href="/wiki/Genetic_diversity" title="Genetic diversity">genetic diversity</a> than there is today,<sup id="cite_ref-freedman2014_12-1" class="reference"><a href="#cite_note-freedman2014-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-thalmann2013_73-1" class="reference"><a href="#cite_note-thalmann2013-73"><span class="cite-bracket">&#91;</span>73<span class="cite-bracket">&#93;</span></a></sup> and within the Pleistocene grey wolf population the variations between local environments would have encouraged a range of wolf ecotypes that were genetically, morphologically and ecologically distinct from one another.<sup id="cite_ref-perri2016_87-0" class="reference"><a href="#cite_note-perri2016-87"><span class="cite-bracket">&#91;</span>87<span class="cite-bracket">&#93;</span></a></sup> One author has proposed that the most likely explanation for the different morphological characteristics of the dog compared to the wolf is that the dog's ancestor was adapted to a different niche than the wolf.<sup id="cite_ref-koler2002_78-7" class="reference"><a href="#cite_note-koler2002-78"><span class="cite-bracket">&#91;</span>78<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Genetic_record">Genetic record</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=14" title="Edit section: Genetic record"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="DNA_sequences">DNA sequences</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=15" title="Edit section: DNA sequences"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The <a href="/wiki/Mitochondria" class="mw-redirect" title="Mitochondria">mitochondria</a> within each <a href="/wiki/Cell_(biology)" title="Cell (biology)">cell</a> contain many copies of a small circular <a href="/wiki/DNA_genome" class="mw-redirect" title="DNA genome">DNA genome</a> and in mammals it is 16,000–18,000 <a href="/wiki/Base_pairs" class="mw-redirect" title="Base pairs">base pairs</a> in length. A cell contains hundreds or thousands of mitochondria and therefore the <a href="/wiki/Genes" class="mw-redirect" title="Genes">genes</a> contained within those mitochondria are more abundant than the genes that occur in the nucleus of the cell.<sup id="cite_ref-rand2001_88-0" class="reference"><a href="#cite_note-rand2001-88"><span class="cite-bracket">&#91;</span>88<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-wayne2006_89-0" class="reference"><a href="#cite_note-wayne2006-89"><span class="cite-bracket">&#91;</span>89<span class="cite-bracket">&#93;</span></a></sup> The abundance of <a href="/wiki/Mitochondrial_DNA" title="Mitochondrial DNA">mitochondrial DNA</a> (mDNA) is useful for the genetic analysis of ancient remains where the DNA has degraded.<sup id="cite_ref-wayne2006_89-1" class="reference"><a href="#cite_note-wayne2006-89"><span class="cite-bracket">&#91;</span>89<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-hofreiter2001_90-0" class="reference"><a href="#cite_note-hofreiter2001-90"><span class="cite-bracket">&#91;</span>90<span class="cite-bracket">&#93;</span></a></sup> </p><p>Mitochondrial <a href="/wiki/DNA_sequences" class="mw-redirect" title="DNA sequences">DNA sequences</a> have a higher <a href="/wiki/Mutation_rate" title="Mutation rate">mutation rate</a> than the mutation rate of nuclear genes and for mammals this rate is 5–10 times faster.<sup id="cite_ref-wayne2006_89-2" class="reference"><a href="#cite_note-wayne2006-89"><span class="cite-bracket">&#91;</span>89<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-li1997_91-0" class="reference"><a href="#cite_note-li1997-91"><span class="cite-bracket">&#91;</span>91<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-persole1999_92-0" class="reference"><a href="#cite_note-persole1999-92"><span class="cite-bracket">&#91;</span>92<span class="cite-bracket">&#93;</span></a></sup> The mitochondrial protein-coding genes evolve much faster and are powerful markers for inferring evolution history at category levels such as families, genera, and species. However, they have evolved at a faster rate than other DNA markers and there is a timing difference in its molecular clock that needs to be validated against other sources. The taxonomic status of uncertain species is better resolved through using <a href="/wiki/Nuclear_DNA" title="Nuclear DNA">nuclear DNA</a> from the nucleus of the cell, which is more suitable for analyzing the recent history.<sup id="cite_ref-wan2003_93-0" class="reference"><a href="#cite_note-wan2003-93"><span class="cite-bracket">&#91;</span>93<span class="cite-bracket">&#93;</span></a></sup> In most cases, mDNA is inherited from the maternal ancestor.<sup id="cite_ref-wayne2006_89-3" class="reference"><a href="#cite_note-wayne2006-89"><span class="cite-bracket">&#91;</span>89<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-birky2001_94-0" class="reference"><a href="#cite_note-birky2001-94"><span class="cite-bracket">&#91;</span>94<span class="cite-bracket">&#93;</span></a></sup> Therefore, <a href="/wiki/Phylogenetic_analysis" class="mw-redirect" title="Phylogenetic analysis">phylogenetic analysis</a> of mDNA sequences within species provides a history of maternal lineages that can be represented as a <a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">phylogenetic tree</a>.<sup id="cite_ref-wayne2006_89-4" class="reference"><a href="#cite_note-wayne2006-89"><span class="cite-bracket">&#91;</span>89<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-avice1994_95-0" class="reference"><a href="#cite_note-avice1994-95"><span class="cite-bracket">&#91;</span>95<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-avice2000_96-0" class="reference"><a href="#cite_note-avice2000-96"><span class="cite-bracket">&#91;</span>96<span class="cite-bracket">&#93;</span></a></sup> </p> <style data-mw-deduplicate="TemplateStyles:r1248256098">@media all and (max-width:720px){.mw-parser-output .mod-gallery{width:100%!important}}.mw-parser-output .mod-gallery{display:table}.mw-parser-output .mod-gallery-default{background:transparent;margin-top:4px}.mw-parser-output .mod-gallery-center{margin-left:auto;margin-right:auto}.mw-parser-output .mod-gallery-left{float:left}.mw-parser-output .mod-gallery-right{float:right}.mw-parser-output .mod-gallery-none{float:none}.mw-parser-output .mod-gallery-collapsible{width:100%}.mw-parser-output .mod-gallery .title,.mw-parser-output .mod-gallery .main,.mw-parser-output .mod-gallery .footer{display:table-row}.mw-parser-output .mod-gallery .title>div{display:table-cell;padding:0 4px 4px;text-align:center;font-weight:bold}.mw-parser-output .mod-gallery .main>div{display:table-cell}.mw-parser-output .mod-gallery .gallery{line-height:1.35em}.mw-parser-output .mod-gallery .footer>div{display:table-cell;padding:4px;text-align:right;font-size:85%;line-height:1em}.mw-parser-output .mod-gallery .title>div *,.mw-parser-output .mod-gallery .footer>div *{overflow:visible}.mw-parser-output .mod-gallery .gallerybox img{background:none!important}.mw-parser-output .mod-gallery .bordered-images .thumb img{border:solid var(--background-color-neutral,#eaecf0)1px}.mw-parser-output .mod-gallery .whitebg .thumb{background:var(--background-color-base,#fff)!important}</style><div class="mod-gallery mod-gallery-default mod-gallery-left"><div class="title"><div>From specimen to phylogenetic tree</div></div><div class="main"><div><ul class="gallery mw-gallery-traditional nochecker bordered-images whitebg"> <li class="gallerybox" style="width: 165px"> <div class="thumb" style="width: 160px; height: 250px;"><span typeof="mw:File"><a href="/wiki/File:Razboinichya_canid.tif" class="mw-file-description" title="The 33,500 year old skull of the &quot;Altai dog&quot;"><img alt="The 33,500 year old skull of the &quot;Altai dog&quot;" src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b9/Razboinichya_canid.tif/lossy-page1-130px-Razboinichya_canid.tif.jpg" decoding="async" width="130" height="79" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b9/Razboinichya_canid.tif/lossy-page1-195px-Razboinichya_canid.tif.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b9/Razboinichya_canid.tif/lossy-page1-260px-Razboinichya_canid.tif.jpg 2x" data-file-width="1781" data-file-height="1078" /></a></span></div> <div class="gallerytext">The 33,500 year old skull of the <a href="/wiki/Paleolithic_dog#Altai_dog_–_33,000_YBP" title="Paleolithic dog">"Altai dog"</a></div> </li> <li class="gallerybox" style="width: 165px"> <div class="thumb" style="width: 160px; height: 250px;"><span typeof="mw:File"><a href="/wiki/File:Eukaryote_DNA-en.svg" class="mw-file-description" title="Location of nuclear DNA within the chromosomes of a cell nucleus"><img alt="Location of nuclear DNA within the chromosomes of a cell nucleus" src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e2/Eukaryote_DNA-en.svg/130px-Eukaryote_DNA-en.svg.png" decoding="async" width="130" height="80" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e2/Eukaryote_DNA-en.svg/195px-Eukaryote_DNA-en.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e2/Eukaryote_DNA-en.svg/260px-Eukaryote_DNA-en.svg.png 2x" data-file-width="1189" data-file-height="734" /></a></span></div> <div class="gallerytext">Location of <a href="/wiki/Nuclear_DNA" title="Nuclear DNA">nuclear DNA</a> within the <a href="/wiki/Chromosomes" class="mw-redirect" title="Chromosomes">chromosomes</a> of a <a href="/wiki/Cell_nucleus" title="Cell nucleus">cell nucleus</a></div> </li> <li class="gallerybox" style="width: 165px"> <div class="thumb" style="width: 160px; height: 250px;"><span typeof="mw:File"><a href="/wiki/File:DNA_animation.gif" class="mw-file-description" title="The structure of part of a DNA double helix"><img alt="The structure of part of a DNA double helix" src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0c/DNA_animation.gif/127px-DNA_animation.gif" decoding="async" width="127" height="220" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/0/0c/DNA_animation.gif 1.5x" data-file-width="181" data-file-height="313" /></a></span></div> <div class="gallerytext">The structure of part of a DNA <a href="/wiki/Double_helix" class="mw-redirect" title="Double helix">double helix</a></div> </li> <li class="gallerybox" style="width: 165px"> <div class="thumb" style="width: 160px; height: 250px;"><span typeof="mw:File"><a href="/wiki/File:Radioactive_Fluorescent_Seq.jpg" class="mw-file-description" title="The results of DNA sequencing listing the DNA molecule&#39;s nucleobases (coded as G, C, A or T)"><img alt="The results of DNA sequencing listing the DNA molecule&#39;s nucleobases (coded as G, C, A or T)" src="//upload.wikimedia.org/wikipedia/commons/thumb/3/3d/Radioactive_Fluorescent_Seq.jpg/130px-Radioactive_Fluorescent_Seq.jpg" decoding="async" width="130" height="185" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/3d/Radioactive_Fluorescent_Seq.jpg/195px-Radioactive_Fluorescent_Seq.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/3d/Radioactive_Fluorescent_Seq.jpg/260px-Radioactive_Fluorescent_Seq.jpg 2x" data-file-width="417" data-file-height="594" /></a></span></div> <div class="gallerytext">The results of <a href="/wiki/DNA_sequencing" title="DNA sequencing">DNA sequencing</a> listing the DNA molecule's <a href="/wiki/Nucleobases" class="mw-redirect" title="Nucleobases">nucleobases</a> (coded as <a href="/wiki/Guanine" title="Guanine">G</a>, <a href="/wiki/Cytosine" title="Cytosine">C</a>, <a href="/wiki/Adenine" title="Adenine">A</a> or <a href="/wiki/Thymine" title="Thymine">T</a>)</div> </li> <li class="gallerybox" style="width: 165px"> <div class="thumb" style="width: 160px; height: 250px;"><span typeof="mw:File"><a href="/wiki/File:Dna-SNP.svg" class="mw-file-description" title="DNA molecule 1 differs from DNA molecule 2 at a single base pair location, called a single-nucleotide polymorphism (a SNP mutation)"><img alt="DNA molecule 1 differs from DNA molecule 2 at a single base pair location, called a single-nucleotide polymorphism (a SNP mutation)" src="//upload.wikimedia.org/wikipedia/commons/thumb/2/2e/Dna-SNP.svg/130px-Dna-SNP.svg.png" decoding="async" width="130" height="83" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/2e/Dna-SNP.svg/195px-Dna-SNP.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/2e/Dna-SNP.svg/260px-Dna-SNP.svg.png 2x" data-file-width="520" data-file-height="333" /></a></span></div> <div class="gallerytext">DNA molecule 1 differs from DNA molecule 2 at a single <a href="/wiki/Base_pair" title="Base pair">base pair</a> location, called a <a href="/wiki/Single-nucleotide_polymorphism" title="Single-nucleotide polymorphism">single-nucleotide polymorphism</a> (a SNP <a href="/wiki/Mutation" title="Mutation">mutation</a>)</div> </li> <li class="gallerybox" style="width: 165px"> <div class="thumb" style="width: 160px; height: 250px;"><span typeof="mw:File"><a href="/wiki/File:Phylogenetic_tree_for_wolves.jpg" class="mw-file-description" title="Phylogenetic tree of wolves under study"><img alt="Phylogenetic tree of wolves under study" src="//upload.wikimedia.org/wikipedia/commons/thumb/2/2f/Phylogenetic_tree_for_wolves.jpg/130px-Phylogenetic_tree_for_wolves.jpg" decoding="async" width="130" height="167" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/2f/Phylogenetic_tree_for_wolves.jpg/195px-Phylogenetic_tree_for_wolves.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/2f/Phylogenetic_tree_for_wolves.jpg/260px-Phylogenetic_tree_for_wolves.jpg 2x" data-file-width="743" data-file-height="952" /></a></span></div> <div class="gallerytext"><a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">Phylogenetic tree</a> of wolves under study</div> </li> </ul></div></div></div> <div style="clear:left; margin-left: 5px; width: 900px; border: #C0C0C0 solid 4px; padding: 5px; overflow-x:auto;max-width:100%;box-sizing:border-box;"> <table style="border: 1px solid #ccc; vertical-align: middle; width: 248px; margin:0.2em 0 0.2em 1em; float:right; clear:right;"> <tbody><tr> <td style="text-align: center; font-weight: bold; border: 0; padding: 0;">Phylogenetic tree </td></tr> <tr> <td style="padding: 5px"> <div class="clade"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade" style="font-size: 85%;line-height:85%;width:325px;"> <tbody><tr> <td class="clade-label first">common&#160;ancestor </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p><a href="/wiki/Golden_jackal" title="Golden jackal">Golden jackal</a> <span typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_X).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fd/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_X%29.jpg/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_X%29.jpg" decoding="async" width="50" height="32" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fd/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_X%29.jpg/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_X%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fd/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_X%29.jpg/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_X%29.jpg 2x" data-file-width="454" data-file-height="287" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p><a href="/wiki/Coyote" title="Coyote">Coyote</a> <span typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_IX).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/41/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IX%29.jpg/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IX%29.jpg" decoding="async" width="50" height="42" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/41/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IX%29.jpg/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IX%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/41/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IX%29.jpg/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IX%29.jpg 2x" data-file-width="414" data-file-height="347" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p><a href="/wiki/Grey_wolf" class="mw-redirect" title="Grey wolf">Grey wolf</a> <span typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_I).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c4/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.jpg/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.jpg" decoding="async" width="50" height="31" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c4/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.jpg/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c4/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.jpg/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.jpg 2x" data-file-width="500" data-file-height="314" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <p><a href="/wiki/Dog" title="Dog">Dog</a> <span style="display: inline-block; transform: scaleX(-1);"><span typeof="mw:File"><a href="/wiki/File:202104_Dog.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f3/202104_Dog.svg/50px-202104_Dog.svg.png" decoding="async" width="50" height="38" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f3/202104_Dog.svg/75px-202104_Dog.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f3/202104_Dog.svg/100px-202104_Dog.svg.png 2x" data-file-width="512" data-file-height="384" /></a></span></span> </p> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td style="text-align: left; font-size: 88%; border: 0; padding: 0;">Phylogenetic relationship between four canids.<sup id="cite_ref-lindblad2005_97-0" class="reference"><a href="#cite_note-lindblad2005-97"><span class="cite-bracket">&#91;</span>97<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-koepfli2015_98-0" class="reference"><a href="#cite_note-koepfli2015-98"><span class="cite-bracket">&#91;</span>98<span class="cite-bracket">&#93;</span></a></sup> </td></tr></tbody></table> <p>The mutations that are different in these 4 <a href="/wiki/Sequence_alignment" title="Sequence alignment">sequences</a> have been numbered and bolded. These mutations can then be used to build a <a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">phylogenetic tree</a> for the four canids. In this example, the dog and grey wolf differ by two substitutions (highlighted in red), and each of them differs from the coyote by four substitutions.<sup id="cite_ref-wayne2006_89-5" class="reference"><a href="#cite_note-wayne2006-89"><span class="cite-bracket">&#91;</span>89<span class="cite-bracket">&#93;</span></a></sup> </p><p><span style="padding-left:7.5em;">&#160;</span><code><b>1</b><span style="padding-left:1.30em;">&#160;</span><b>2</b><span style="padding-left:1.15em;">&#160;</span><b>3</b><span style="padding-left:1.1em;">&#160;</span><b>4</b><span style="padding-left:1.65em;">&#160;</span><b>5</b><span style="padding-left:1.7em;">&#160;</span><b>67</b></code> </p><p>Golden Jackal<span style="padding-left:1.0em;">&#160;</span><code><b>A</b>-G-<b>C</b>-T-<b>G</b>-T-<b>C</b>-GA-<b>T</b>-TC-<b>CA</b></code> </p><p>Coyote <span style="padding-left:3.8em;">&#160;</span><code><b>A</b>-G-<b>C</b>-T-<b>A</b>-T-<b>C</b>-GA-<b>A</b>-TC-<b>GA</b></code> </p><p>Wolf <span style="padding-left:5.0em;">&#160;</span><code><b>T</b>-G-<b>C</b>-T-<b>A</b>-T-<b>G</b>-GA-<b>T</b>-TC-<b>CT</b></code> </p><p>Dog <span style="padding-left:5.15em;">&#160;</span><code><b>T</b>-G-<span style="color: red"><b>G</b></span>-T-<b>A</b>-T-<b>G</b>-GA-<b>T</b>-TC-<b>C</b><span style="color: red"><b>A</b></span></code> </p> </div> <p>The mDNA sequences of the dog and wolf differ by only 0–12 substitutions within 261 base-pairs, whereas dogs always differed from coyotes and jackals by at least 20 substitutions.<sup id="cite_ref-wayne2006_89-6" class="reference"><a href="#cite_note-wayne2006-89"><span class="cite-bracket">&#91;</span>89<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-vila1997_99-0" class="reference"><a href="#cite_note-vila1997-99"><span class="cite-bracket">&#91;</span>99<span class="cite-bracket">&#93;</span></a></sup> This finding implies that the dog derived from the wolf and that there has been repeated back-crossing,<sup id="cite_ref-vila1997_99-1" class="reference"><a href="#cite_note-vila1997-99"><span class="cite-bracket">&#91;</span>99<span class="cite-bracket">&#93;</span></a></sup> or that the dog may have descended from a now extinct species of canid whose closest living relative is the modern wolf.<sup id="cite_ref-wayne1999_100-0" class="reference"><a href="#cite_note-wayne1999-100"><span class="cite-bracket">&#91;</span>100<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Marker_issue">Marker issue</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=16" title="Edit section: Marker issue"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Different DNA studies may give conflicting results because of the specimens selected, the technology used, and the assumptions made by the researchers.<sup id="cite_ref-boyko2009_101-0" class="reference"><a href="#cite_note-boyko2009-101"><span class="cite-bracket">&#91;</span>101<span class="cite-bracket">&#93;</span></a></sup> Any one from a panel of <a href="/wiki/Genetic_marker" title="Genetic marker">genetic markers</a> can be chosen for use in a study. The techniques used to <a href="/wiki/DNA_extraction" title="DNA extraction">extract</a>, <a href="/wiki/DNA_sequencing" title="DNA sequencing">locate</a> and <a href="/wiki/DNA_analysis" class="mw-redirect" title="DNA analysis">compare</a> genetic sequences can be applied using advances in technology, which allows researchers to observe longer lengths of <a href="/wiki/Base_pair#Length_measurements" title="Base pair">base pairs</a> that provide more data to give better <a href="/wiki/Phylogenetic" class="mw-redirect" title="Phylogenetic">phylogenetic</a> resolution.<sup id="cite_ref-pang2009_102-0" class="reference"><a href="#cite_note-pang2009-102"><span class="cite-bracket">&#91;</span>102<span class="cite-bracket">&#93;</span></a></sup> <a href="/wiki/Phylogenetic_trees" class="mw-redirect" title="Phylogenetic trees">Phylogenetic trees</a> compiled using different <a href="/wiki/Genetic_markers" class="mw-redirect" title="Genetic markers">genetic markers</a> have given conflicting results on the relationship between the wolf, dog and coyote. One study based on <a href="/wiki/Single-nucleotide_polymorphism" title="Single-nucleotide polymorphism">SNPs</a><sup id="cite_ref-cronin2014_103-0" class="reference"><a href="#cite_note-cronin2014-103"><span class="cite-bracket">&#91;</span>103<span class="cite-bracket">&#93;</span></a></sup> (a single <a href="/wiki/Mutation" title="Mutation">mutation</a>), and another based on <a href="/wiki/Nuclear_gene" title="Nuclear gene">nuclear gene</a> sequences<sup id="cite_ref-bardeleben2005_104-0" class="reference"><a href="#cite_note-bardeleben2005-104"><span class="cite-bracket">&#91;</span>104<span class="cite-bracket">&#93;</span></a></sup> (taken from the <a href="/wiki/Cell_nucleus" title="Cell nucleus">cell nucleus</a>), showed dogs clustering with coyotes and separate from wolves. Another study based on SNPS showed wolves clustering with coyotes and separate from dogs.<sup id="cite_ref-gray2010_105-0" class="reference"><a href="#cite_note-gray2010-105"><span class="cite-bracket">&#91;</span>105<span class="cite-bracket">&#93;</span></a></sup> Other studies based on a number of markers show the more widely accepted result of wolves clustering with dogs separate from coyotes.<sup id="cite_ref-vila1997_99-2" class="reference"><a href="#cite_note-vila1997-99"><span class="cite-bracket">&#91;</span>99<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-wayne2012_106-0" class="reference"><a href="#cite_note-wayne2012-106"><span class="cite-bracket">&#91;</span>106<span class="cite-bracket">&#93;</span></a></sup> These results demonstrate that caution is needed when interpreting the results provided by genetic markers.<sup id="cite_ref-cronin2014_103-1" class="reference"><a href="#cite_note-cronin2014-103"><span class="cite-bracket">&#91;</span>103<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Timing_issue">Timing issue</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=17" title="Edit section: Timing issue"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>There are two key assumptions that are made for dating the divergence time for species: the <a href="/wiki/Generation_time" title="Generation time">generation time</a> and the genetic <a href="/wiki/Mutation_rate" title="Mutation rate">mutation rate</a> per generation. The time between generations for wolves is assumed to be three years based on the extant grey wolf, and two years for the dog based on the extant dog.<sup id="cite_ref-lindblad2005_97-1" class="reference"><a href="#cite_note-lindblad2005-97"><span class="cite-bracket">&#91;</span>97<span class="cite-bracket">&#93;</span></a></sup> One recent major study assumed a generation time of 2 years for the dog for as far back as 10,000 years ago, and then assumed a generation time of 3 years (the same as the wolf) before that to calculate a proposed divergence time between the two.<sup id="cite_ref-freedman2014_12-2" class="reference"><a href="#cite_note-freedman2014-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup> In 2017, the wolf research scientist <a href="/wiki/L._David_Mech" title="L. David Mech">L. David Mech</a> queried why evolutionary biologists were calculating the approximate time of the dog diverging from the wolf through using a wolf generation time of three years when published works using large data sets demonstrate a figure of 4.2–4.7 years. They were encouraged to recalculate their divergence dates accordingly.<sup id="cite_ref-mech2017_107-0" class="reference"><a href="#cite_note-mech2017-107"><span class="cite-bracket">&#91;</span>107<span class="cite-bracket">&#93;</span></a></sup> </p><p>DNA studies are conducted but with "the mutation rate as the dominant source of uncertainty."<sup id="cite_ref-freedman2014_12-3" class="reference"><a href="#cite_note-freedman2014-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup> In 2005, Lindblad-Toh sequenced the first draft genome of the extant dog, and calculated a proposed mutation rate of 1x10<sup>−8</sup> mutations per generation.<sup id="cite_ref-lindblad2005_97-2" class="reference"><a href="#cite_note-lindblad2005-97"><span class="cite-bracket">&#91;</span>97<span class="cite-bracket">&#93;</span></a></sup> In 2015, Skoglund was able to sequence the first draft genome of the 35,000 YBP <a href="/wiki/Pleistocene_wolf#Taimyr_wolf" title="Pleistocene wolf">Taimyr wolf</a> and used its radio-carbon date to validate a proposed genetic mutation rate of 0.4x10<sup>−8</sup> mutations per generation.<sup id="cite_ref-skoglund2015_14-1" class="reference"><a href="#cite_note-skoglund2015-14"><span class="cite-bracket">&#91;</span>14<span class="cite-bracket">&#93;</span></a></sup> The difference is a timing factor of 2.5, however another study stated that because only one Pleistocene wolf specimen has so far been sequenced, then the result should be treated with caution, with that study then providing both estimates to calculate the proposed divergence times between the wolf and dog.<sup id="cite_ref-fan2016_13-1" class="reference"><a href="#cite_note-fan2016-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> However, in 2016 the mutation rate of the 4,800 YBP <a href="/wiki/Origin_of_the_domestic_dog#Late_Pleistocene_wolf_admixture" class="mw-redirect" title="Origin of the domestic dog">Newgrange dog</a> matched that of the Taimyr wolf.<sup id="cite_ref-frantz2016_108-0" class="reference"><a href="#cite_note-frantz2016-108"><span class="cite-bracket">&#91;</span>108<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Wolf-like_canids">Wolf-like canids</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=18" title="Edit section: Wolf-like canids"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Canidae#Phylogenetic_relationships" title="Canidae">Canidae §&#160;Phylogenetic relationships</a></div> <p>The wolf-like canids (the canid subfamily <a href="/wiki/Caninae" title="Caninae">Caninae</a>) are a group of large carnivores that are genetically closely related because their <a href="/wiki/Chromosome" title="Chromosome">chromosomes</a> number 78. The group includes genus <i>Canis</i>, <i>Cuon</i> and <i>Lycaon</i>. The members are the <a href="/wiki/Dog" title="Dog">dog</a> <i>(C. familiaris)</i>, <a href="/wiki/Grey_wolf" class="mw-redirect" title="Grey wolf">grey wolf</a> (<i>C. lupus</i>), <a href="/wiki/Coyote" title="Coyote">coyote</a> (<i>C. latrans</i>), <a href="/wiki/Golden_jackal" title="Golden jackal">golden jackal</a> (<i>C. aureus</i>), <a href="/wiki/Ethiopian_wolf" title="Ethiopian wolf">Ethiopian wolf</a> (<i>C. simensis</i>), <a href="/wiki/Black-backed_jackal" title="Black-backed jackal">black-backed jackal</a> (<i>Lupulella mesomelas</i>), <a href="/wiki/Side-striped_jackal" title="Side-striped jackal">side-striped jackal</a> (<i>Lupulella adusta</i>), <a href="/wiki/Dhole" title="Dhole">dhole</a> (<i>Cuon alpinus</i>), and <a href="/wiki/African_wild_dog" title="African wild dog">African wild dog</a> (<i>Lycaon pictus</i>).<sup id="cite_ref-wayne1993_109-0" class="reference"><a href="#cite_note-wayne1993-109"><span class="cite-bracket">&#91;</span>109<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-wayne1989_110-0" class="reference"><a href="#cite_note-wayne1989-110"><span class="cite-bracket">&#91;</span>110<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-vila2012_111-0" class="reference"><a href="#cite_note-vila2012-111"><span class="cite-bracket">&#91;</span>111<span class="cite-bracket">&#93;</span></a></sup> Newly proposed members include the <a href="/wiki/Red_wolf" title="Red wolf">red wolf</a> (<i>Canis rufus</i>), <a href="/wiki/Eastern_wolf" title="Eastern wolf">eastern wolf</a> (<i>Canis lycaon</i>), and <a href="/wiki/African_golden_wolf" class="mw-redirect" title="African golden wolf">African golden wolf</a> (<i>C. anthus</i>). As they possess 78 chromosomes, all members of the genus <i>Canis</i> (coyotes, wolves, jackals) are <a href="/wiki/Karyology" class="mw-redirect" title="Karyology">karyologically</a> indistinguishable from each other, and from the dhole and the African hunting dog.<sup id="cite_ref-wayne2006_89-7" class="reference"><a href="#cite_note-wayne2006-89"><span class="cite-bracket">&#91;</span>89<span class="cite-bracket">&#93;</span></a></sup><sup class="reference nowrap"><span title="Page / location: p279">&#58;&#8202;p279&#8202;</span></sup><sup id="cite_ref-wurster1982_112-0" class="reference"><a href="#cite_note-wurster1982-112"><span class="cite-bracket">&#91;</span>112<span class="cite-bracket">&#93;</span></a></sup> The members of <i>Canis</i> can potentially <a href="/wiki/Hybrid_(biology)" title="Hybrid (biology)">interbreed</a><sup id="cite_ref-wayne1999_100-1" class="reference"><a href="#cite_note-wayne1999-100"><span class="cite-bracket">&#91;</span>100<span class="cite-bracket">&#93;</span></a></sup> and there is evidence that the Ethiopian wolf has hybridized with dogs.<sup id="cite_ref-gottelli1994_113-0" class="reference"><a href="#cite_note-gottelli1994-113"><span class="cite-bracket">&#91;</span>113<span class="cite-bracket">&#93;</span></a></sup> According to zoologist <a href="/wiki/Reginald_Innes_Pocock" title="Reginald Innes Pocock">Reginald Pocock</a>, a dhole interbred with a golden jackal.<sup id="cite_ref-pocock1941_114-0" class="reference"><a href="#cite_note-pocock1941-114"><span class="cite-bracket">&#91;</span>114<span class="cite-bracket">&#93;</span></a></sup> The African hunting dog is large, highly mobile, known to disperse over large distances and are rare throughout much of their geographical range,<sup id="cite_ref-girman2001_115-0" class="reference"><a href="#cite_note-girman2001-115"><span class="cite-bracket">&#91;</span>115<span class="cite-bracket">&#93;</span></a></sup> making opportunities for hybridization difficult. A study of the maternal <a href="/wiki/Mitochondrial_DNA" title="Mitochondrial DNA">mitochondrial DNA</a> of the black-backed jackal could find no evidence of <a href="/wiki/Genotypes" class="mw-redirect" title="Genotypes">genotypes</a> from the most likely mates – the side-striped jackal nor the golden jackal – indicating that male black-backed jackals had not bred with these.<sup id="cite_ref-wayne1990_116-0" class="reference"><a href="#cite_note-wayne1990-116"><span class="cite-bracket">&#91;</span>116<span class="cite-bracket">&#93;</span></a></sup> A search of the scientific literature could not find evidence of hybridization for the rare side-striped jackal. </p><p>A DNA sequence alignment for the wolf-like canids gave a phylogenetic tree with the grey wolf and dog being the most closely related, followed by a close affiliation with the coyote, golden jackal and Ethiopian wolf, and the dog can hybridize in the wild with these three species. Next closest to this group are the dhole and African wild dog that both have unique meat-slicing teeth, suggesting that this adaptation was later lost by the other members.<sup id="cite_ref-lindblad2005_97-3" class="reference"><a href="#cite_note-lindblad2005-97"><span class="cite-bracket">&#91;</span>97<span class="cite-bracket">&#93;</span></a></sup> The two African jackals are shown as the most <a href="/wiki/Basal_(phylogenetics)" title="Basal (phylogenetics)">basal</a> members of this clade, which means that this tree is indicating an African origin for the clade.<sup id="cite_ref-lindblad2005_97-4" class="reference"><a href="#cite_note-lindblad2005-97"><span class="cite-bracket">&#91;</span>97<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-kaminski2014_117-0" class="reference"><a href="#cite_note-kaminski2014-117"><span class="cite-bracket">&#91;</span>117<span class="cite-bracket">&#93;</span></a></sup> The tree illustrates the <a href="/wiki/Genotype%E2%80%93phenotype_distinction" title="Genotype–phenotype distinction">genotype–phenotype distinction</a>, where a <a href="/wiki/Genotype" title="Genotype">genotype</a> is an organism's full <a href="/wiki/Hereditary" class="mw-redirect" title="Hereditary">hereditary</a> information and a <a href="/wiki/Phenotype" title="Phenotype">phenotype</a> is an organism's actual observed properties, such as <a href="/wiki/Morphology_(biology)" title="Morphology (biology)">morphology</a>, <a href="/wiki/Developmental_biology" title="Developmental biology">development</a>, or <a href="/wiki/Behavior" title="Behavior">behavior</a>. By phenotype, the dhole (genus <i>Cuon</i>) and the African hunting dog (genus <i>Lycaon</i>) are not classified as members of the genus <i>Canis</i>, but by genotype they are closer to dogs, wolves and coyotes than are the two genus <i>Canis</i> jackals – the Side-striped jackal (<i>C. adustus</i>) and the Black-backed jackal (<i>C. mesomelas</i>). </p><p>In 2015, a study of mitochondrial genome sequences and nuclear genome sequences of African and Eurasian canids indicated that extant wolf-like canids had colonized Africa from Eurasia at least 5 times throughout the Pliocene and Pleistocene, which is consistent with fossil evidence suggesting that much of the African canid diversity resulted from the immigration of Eurasian ancestors, likely coincident with Plio-Pleistocene climatic oscillations between arid and humid conditions.<sup id="cite_ref-koepfli2015_98-1" class="reference"><a href="#cite_note-koepfli2015-98"><span class="cite-bracket">&#91;</span>98<span class="cite-bracket">&#93;</span></a></sup> </p><p>The phylogenetic tree for the wolf-like canids may give conflicting positions for the black-backed jackal and the side-striped jackal relative to the genus <i>Canis</i> members depending on whether the genetic markers were based on mitochondrial DNA or nuclear DNA. The explanation proposed for this mito-nuclear discord is that mitochondrial DNA <a href="/wiki/Introgression" title="Introgression">introgression</a> occurred from an ancient ancestor of genus <i>Canis</i> into the lineage that led to the black-backed jackal around 6.2–5.2 million years ago.<sup id="cite_ref-Hassanin2021_118-0" class="reference"><a href="#cite_note-Hassanin2021-118"><span class="cite-bracket">&#91;</span>118<span class="cite-bracket">&#93;</span></a></sup> </p><p><b><a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">Phylogenetic tree</a> of the extant wolf-like canids,<sup id="cite_ref-mtdna_seq_desc_132-0" class="reference"><a href="#cite_note-mtdna_seq_desc-132"><span class="cite-bracket">&#91;</span>a<span class="cite-bracket">&#93;</span></a></sup> with the pink shading representing the species <i>Canis lupus</i>.</b> </p> <style data-mw-deduplicate="TemplateStyles:r934431750">.mw-parser-output table.clade{border-spacing:0;margin:0;font-size:100%;line-height:100%;border-collapse:separate;width:auto}.mw-parser-output table.clade table.clade{width:100%}.mw-parser-output table.clade td.clade-label{width:0.7em;padding:0 0.15em;vertical-align:bottom;text-align:center;border-left:1px solid;border-bottom:1px solid;white-space:nowrap}.mw-parser-output table.clade td.clade-fixed-width{overflow:hidden;text-overflow:ellipsis}.mw-parser-output table.clade td.clade-fixed-width:hover{overflow:visible}.mw-parser-output table.clade td.clade-label.first{border-left:none;border-right:none}.mw-parser-output table.clade td.clade-label.reverse{border-left:none;border-right:1px solid}.mw-parser-output table.clade td.clade-slabel{padding:0 0.15em;vertical-align:top;text-align:center;border-left:1px solid;white-space:nowrap}.mw-parser-output table.clade td.clade-slabel:hover{overflow:visible}.mw-parser-output table.clade td.clade-slabel.last{border-left:none;border-right:none}.mw-parser-output table.clade td.clade-slabel.reverse{border-left:none;border-right:1px solid}.mw-parser-output table.clade td.clade-bar{vertical-align:middle;text-align:left;padding:0 0.5em;position:relative}.mw-parser-output table.clade td.clade-bar.reverse{text-align:right;position:relative}.mw-parser-output table.clade td.clade-leaf{border:0;padding:0;text-align:left}.mw-parser-output table.clade td.clade-leafR{border:0;padding:0;text-align:right}.mw-parser-output table.clade td.clade-leaf.reverse{text-align:right}.mw-parser-output table.clade:hover span.linkA{background-color:yellow}.mw-parser-output table.clade:hover span.linkB{background-color:green}</style> <table class="clade" style="background-color:#ccccff;font-size:85%;"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:50px;max-width:50px;padding:0em;overflow:visible;"><div style="position:relative;"><div style="position:absolute;white‑space:nowrap;right:‑25px;;"></div><br /></div> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:50px;max-width:50px;padding:0em;overflow:visible;"><div style="position:relative;"><div style="position:absolute;white‑space:nowrap;right:25px;;">3.5 mya</div><br /></div> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:50px;max-width:50px;padding:0em;overflow:visible;"><div style="position:relative;"><div style="position:absolute;white‑space:nowrap;right:25px;;">3.0 mya</div><br /></div> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:50px;max-width:50px;padding:0em;overflow:visible;"><div style="position:relative;"><div style="position:absolute;white‑space:nowrap;right:25px;;">2.5 mya</div><br /></div> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:50px;max-width:50px;padding:0em;overflow:visible;"><div style="position:relative;"><div style="position:absolute;white‑space:nowrap;right:25px;;">2.0 mya</div><br /></div> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:50px;max-width:50px;padding:0em;overflow:visible;"><div style="position:relative;"><div style="position:absolute;white‑space:nowrap;right:25px;;">1.5 mya</div><br /></div> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:50px;max-width:50px;padding:0em;overflow:visible;"><div style="position:relative;"><div style="position:absolute;white‑space:nowrap;right:25px;;">1.0 mya</div><br /></div> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:50px;max-width:50px;padding:0em;overflow:visible;"><div style="position:relative;"><div style="position:absolute;white‑space:nowrap;right:25px;;">0.5 mya</div><br /></div> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:150px;max-width:150px;padding:0em;overflow:visible;text-align:left;"><div style="position:relative;"><div style="position:absolute;white‑space:nowrap;right:‑25px;;">Present</div><br /></div> </td> <td rowspan="2" class="clade-leaf"> <p><span></span> </p> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:150px;padding:0em;border-left:1px solid black;"><br /> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:50px;padding:0em;border-left:1px solid black;background-color:#aaaaff;"><br /> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:50px;padding:0em;border-left:1px solid black;"><br /> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:50px;padding:0em;border-left:1px solid black;background-color:#aaaaff;"><br /> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:50px;padding:0em;border-left:1px solid black;"><br /> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:50px;padding:0em;border-left:1px solid black;background-color:#aaaaff;"><br /> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:50px;padding:0em;border-left:1px solid black;"><br /> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:50px;padding:0em;border-left:1px solid black;background-color:#aaaaff;"><br /> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:50px;padding:0em;border-left:1px solid black;"><br /> </td></tr></tbody></table> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade" style="font-size:85%;line-height:85%;"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:50px;max-width:50px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:50px;max-width:50px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:38px;max-width:38px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:262px;max-width:262px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <table style="width:100%;border-spacing:0;"> <tbody><tr> <td style="border:0;padding:0;"> <p><a href="/wiki/Side-striped_jackal" title="Side-striped jackal">Side-striped jackal</a> </p> </td> <td style="border:0;padding:0;"> <figure class="mw-halign-right" typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_XIII).png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a8/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XIII%29.png/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XIII%29.png" decoding="async" width="50" height="31" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a8/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XIII%29.png/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XIII%29.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/a8/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XIII%29.png/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XIII%29.png 2x" data-file-width="489" data-file-height="302" /></a><figcaption></figcaption></figure> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel clade-fixed-width" style="max-width:262px;padding:0em;"><br /> </td></tr> <tr> <td class="clade-label clade-fixed-width" style="max-width:262px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <table style="width:100%;border-spacing:0;"> <tbody><tr> <td style="border:0;padding:0;"> <p><a href="/wiki/Black-backed_jackal" title="Black-backed jackal">Black-backed jackal</a> </p> </td> <td style="border:0;padding:0;"> <figure class="mw-halign-right" typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_XII).png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/ca/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XII%29.png/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XII%29.png" decoding="async" width="50" height="34" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/ca/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XII%29.png/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XII%29.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/ca/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XII%29.png/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XII%29.png 2x" data-file-width="430" data-file-height="290" /></a><figcaption></figcaption></figure> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:262px;padding:0em;"><br /> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:38px;padding:0em;overflow:visible;"><div style="position:relative;"><div style="position:absolute;white‑space:nowrap;right:0px;"><i>2.62 mya</i></div><br /></div> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel clade-fixed-width" style="max-width:50px;padding:0em;overflow:visible;"><div style="position:relative;"><div style="position:absolute;white‑space:nowrap;right:0px;"></div><br /></div> </td></tr> <tr> <td class="clade-label clade-fixed-width" style="max-width:50px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:50px;max-width:50px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:250px;max-width:250px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <table style="width:100%;border-spacing:0;"> <tbody><tr> <td style="border:0;padding:0;"> <p><a href="/wiki/African_wild_dog" title="African wild dog">African wild dog</a> </p> </td> <td style="border:0;padding:0;"> <figure class="mw-halign-right" typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_XLIV).png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a5/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XLIV%29.png/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XLIV%29.png" decoding="async" width="50" height="31" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a5/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XLIV%29.png/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XLIV%29.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/a5/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XLIV%29.png/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XLIV%29.png 2x" data-file-width="445" data-file-height="277" /></a><figcaption></figcaption></figure> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:250px;padding:0em;"><br /> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel clade-fixed-width" style="max-width:50px;padding:0em;"><br /> </td></tr> <tr> <td class="clade-label clade-fixed-width" style="max-width:50px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:50px;max-width:50px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:200px;max-width:200px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <table style="width:100%;border-spacing:0;"> <tbody><tr> <td style="border:0;padding:0;"> <p><a href="/wiki/Dhole" title="Dhole">Dhole</a> </p> </td> <td style="border:0;padding:0;"> <figure class="mw-halign-right" typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_XLI).png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c8/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XLI%29.png/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XLI%29.png" decoding="async" width="50" height="32" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c8/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XLI%29.png/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XLI%29.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c8/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XLI%29.png/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XLI%29.png 2x" data-file-width="471" data-file-height="302" /></a><figcaption></figcaption></figure> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:200px;padding:0em;"><br /> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel clade-fixed-width" style="max-width:50px;padding:0em;"><br /> </td></tr> <tr> <td class="clade-label clade-fixed-width" style="max-width:50px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:104px;max-width:104px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:96px;max-width:96px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <table style="width:100%;border-spacing:0;"> <tbody><tr> <td style="border:0;padding:0;"> <p><a href="/wiki/Ethiopian_wolf" title="Ethiopian wolf">Ethiopian wolf</a> </p> </td> <td style="border:0;padding:0;"> <figure class="mw-halign-right" typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_VI).png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d6/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_VI%29.png/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_VI%29.png" decoding="async" width="50" height="33" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d6/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_VI%29.png/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_VI%29.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d6/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_VI%29.png/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_VI%29.png 2x" data-file-width="471" data-file-height="310" /></a><figcaption></figcaption></figure> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:96px;padding:0em;"><br /> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel clade-fixed-width" style="max-width:104px;padding:0em;"><br /> </td></tr> <tr> <td class="clade-label clade-fixed-width" style="max-width:104px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:36px;max-width:36px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <div class="clade"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:60px;max-width:60px;padding:0em;"> </td> <td rowspan="2" class="clade-leaf"> <table style="width:100%;border-spacing:0"> <tbody><tr> <td style="border:0;padding:0"> <p><a href="/wiki/Golden_jackal" title="Golden jackal">Golden jackal</a> </p> </td> <td style="border:0;padding:0"> <figure class="mw-halign-right" typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_X).png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/7a/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_X%29.png/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_X%29.png" decoding="async" width="50" height="32" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/7a/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_X%29.png/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_X%29.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/7a/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_X%29.png/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_X%29.png 2x" data-file-width="454" data-file-height="287" /></a><figcaption></figcaption></figure> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="width:60px;max-width:60px;padding:0em;"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel clade-fixed-width" style="max-width:36px;padding:0em;"><br /> </td></tr> <tr> <td class="clade-label clade-fixed-width" style="max-width:36px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:22px;max-width:22px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:38px;max-width:38px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <table style="width:100%;border-spacing:0;"> <tbody><tr> <td style="border:0;padding:0;"> <p><a href="/wiki/African_golden_wolf" class="mw-redirect" title="African golden wolf">African golden wolf</a> </p> </td> <td style="border:0;padding:0;"> <figure class="mw-halign-right" typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_XI).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/be/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XI%29.jpg/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XI%29.jpg" decoding="async" width="50" height="32" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/be/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XI%29.jpg/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XI%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/be/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XI%29.jpg/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XI%29.jpg 2x" data-file-width="480" data-file-height="308" /></a><figcaption></figcaption></figure> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:38px;padding:0em;"><br /> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel clade-fixed-width" style="max-width:22px;padding:0em;"><br /> </td></tr> <tr> <td class="clade-label clade-fixed-width" style="max-width:22px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:14px;max-width:14px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:24px;max-width:24px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <table style="width:100%;border-spacing:0;"> <tbody><tr> <td style="border:0;padding:0;"> <p><a href="/wiki/Coyote" title="Coyote">Coyote</a> </p> </td> <td style="border:0;padding:0;"> <figure class="mw-halign-right" typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_IX).png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/9d/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IX%29.png/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IX%29.png" decoding="async" width="50" height="42" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/9d/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IX%29.png/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IX%29.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/9d/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IX%29.png/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IX%29.png 2x" data-file-width="414" data-file-height="347" /></a><figcaption></figcaption></figure> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:24px;padding:0em;"><br /> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel clade-fixed-width" style="max-width:14px;padding:0em;"><br /> </td></tr> <tr> <td class="clade-label clade-fixed-width" style="max-width:14px;padding:0em;background-color:#ffcccc;"><br /> </td> <td rowspan="2" class="clade-leaf" style="background-color:#ffcccc;"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:12px;max-width:12px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:12px;max-width:12px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <table style="width:100%;border-spacing:0;"> <tbody><tr> <td style="border:0;padding:0;"> <p><a href="/wiki/Himalayan_wolf" title="Himalayan wolf">Himalayan wolf</a> </p> </td> <td style="border:0;padding:0;"> <figure class="mw-halign-right" typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_I).png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/ce/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.png/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.png" decoding="async" width="50" height="31" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/ce/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.png/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/ce/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.png/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.png 2x" data-file-width="500" data-file-height="314" /></a><figcaption></figcaption></figure> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:12px;padding:0em;"><br /> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel clade-fixed-width" style="max-width:12px;padding:0em;"><br /> </td></tr> <tr> <td class="clade-label clade-fixed-width" style="max-width:12px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:4px;max-width:4px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:8px;max-width:8px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <table style="width:100%;border-spacing:0;"> <tbody><tr> <td style="border:0;padding:0;"> <p><a href="/wiki/Indian_wolf#Canis_indica" title="Indian wolf">Indian<br />plains wolf</a> </p> </td> <td style="border:0;padding:0;"> <figure class="mw-halign-right" typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_I).png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/ce/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.png/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.png" decoding="async" width="50" height="31" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/ce/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.png/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/ce/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.png/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.png 2x" data-file-width="500" data-file-height="314" /></a><figcaption></figcaption></figure> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:8px;padding:0em;"><br /> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel clade-fixed-width" style="max-width:4px;padding:0em;"><br /> </td></tr> <tr> <td class="clade-label clade-fixed-width" style="max-width:4px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:5px;max-width:5px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:1px;max-width:1px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <table style="width:100%;border-spacing:0;"> <tbody><tr> <td style="border:0;padding:0;"> <p>†<a href="/wiki/Pleistocene_wolf" title="Pleistocene wolf">Late Pleistocene<br />&#160;&#160;wolf</a> </p> </td> <td style="border:0;padding:0;"> <figure class="mw-halign-right" typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_I).png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/ce/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.png/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.png" decoding="async" width="50" height="31" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/ce/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.png/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/ce/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.png/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.png 2x" data-file-width="500" data-file-height="314" /></a><figcaption></figcaption></figure> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:1px;padding:0em;"><br /> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel clade-fixed-width" style="max-width:5px;padding:0em;"><br /> </td></tr> <tr> <td class="clade-label clade-fixed-width" style="max-width:5px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r934431750"> <table class="clade"> <tbody><tr> <td class="clade-label first clade-fixed-width" style="width:3px;max-width:3px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <table style="width:100%;border-spacing:0;"> <tbody><tr> <td style="border:0;padding:0;"> <p><a href="/wiki/Wolf" title="Wolf">Modern grey wolf</a> </p> </td> <td style="border:0;padding:0;"> <figure class="mw-halign-right" typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_I).png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/ce/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.png/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.png" decoding="async" width="50" height="31" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/ce/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.png/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/ce/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.png/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.png 2x" data-file-width="500" data-file-height="314" /></a><figcaption></figcaption></figure> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel clade-fixed-width" style="max-width:3px;padding:0em;"><br /> </td></tr> <tr> <td class="clade-label clade-fixed-width" style="max-width:3px;padding:0em;"><br /> </td> <td rowspan="2" class="clade-leaf"> <table style="width:100%;border-spacing:0;"> <tbody><tr> <td style="border:0;padding:0;"> <p><a href="/wiki/Dog" title="Dog">Dog</a> </p> </td> <td style="border:0;padding:0;"> <span style="display: inline-block; transform: scaleX(-1);"><figure class="mw-halign-right" typeof="mw:File"><a href="/wiki/File:202104_Dog.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f3/202104_Dog.svg/50px-202104_Dog.svg.png" decoding="async" width="50" height="38" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f3/202104_Dog.svg/75px-202104_Dog.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f3/202104_Dog.svg/100px-202104_Dog.svg.png 2x" data-file-width="512" data-file-height="384" /></a><figcaption></figcaption></figure></span> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:3px;padding:0em;"><br /> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:5px;padding:0em;overflow:visible;"><div style="position:relative;"><div style="position:absolute;white‑space:nowrap;right:0px;">25 kya</div><br /></div> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:4px;padding:0em;overflow:visible;"><div style="position:relative;"><div style="position:absolute;white‑space:nowrap;right:0px;"><i>80 kya</i> </div><br /></div> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:12px;padding:0em;overflow:visible;"><div style="position:relative;"><div style="position:absolute;white‑space:nowrap;right:0px;"><i>120 kya</i> </div><br /></div> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:14px;padding:0em;overflow:visible;background-color:#ffcccc;"><div style="position:relative;"><div style="position:absolute;white‑space:nowrap;right:0px;"><i>240 kya</i> </div><br /></div> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:22px;padding:0em;overflow:visible;"><div style="position:relative;"><div style="position:absolute;white‑space:nowrap;right:0px;"><i>380 kya</i> </div><br /></div> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:36px;padding:0em;overflow:visible;"><div style="position:relative;"><div style="position:absolute;white‑space:nowrap;right:0px;"><i>600 kya</i> </div><br /></div> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:104px;padding:0em;overflow:visible;"><div style="position:relative;"><div style="position:absolute;white‑space:nowrap;right:0px;"><i>960 kya</i> </div><br /></div> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:50px;padding:0em;overflow:visible;"><div style="position:relative;"><div style="position:absolute;white‑space:nowrap;right:0px;"><i>2.0 mya</i> </div><br /></div> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:50px;padding:0em;overflow:visible;"><div style="position:relative;"><div style="position:absolute;white‑space:nowrap;right:0px;"><i>2.5 mya</i> </div><br /></div> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:50px;padding:0em;overflow:visible;"><div style="position:relative;"><div style="position:absolute;white‑space:nowrap;right:0px;"><i>3.0 mya</i> </div><br /></div> </td></tr></tbody></table> </td></tr> <tr> <td class="clade-slabel last clade-fixed-width" style="max-width:50px;padding:0em;overflow:visible;"><div style="position:relative;"><div style="position:absolute;white‑space:nowrap;right:0px;"><i>3.5 mya</i></div><br /></div> </td></tr></tbody></table> <div style="clear:both;" class=""></div> <div class="mw-heading mw-heading4"><h4 id="Admixture_with_an_extinct_unknown_canid">Admixture with an extinct unknown canid</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=19" title="Edit section: Admixture with an extinct unknown canid"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Canis_early_hybridisation.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/ef/Canis_early_hybridisation.jpg/400px-Canis_early_hybridisation.jpg" decoding="async" width="400" height="300" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/ef/Canis_early_hybridisation.jpg/600px-Canis_early_hybridisation.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/ef/Canis_early_hybridisation.jpg/800px-Canis_early_hybridisation.jpg 2x" data-file-width="960" data-file-height="720" /></a><figcaption>Canis hybridisation in the distant past<sup id="cite_ref-gopalakrishnan2018_119-1" class="reference"><a href="#cite_note-gopalakrishnan2018-119"><span class="cite-bracket">&#91;</span>119<span class="cite-bracket">&#93;</span></a></sup></figcaption></figure> <p>In 2018, <a href="/wiki/Whole_genome_sequencing" title="Whole genome sequencing">whole genome sequencing</a> was used to compare members of genus <i>Canis</i>, along with the dhole (<i>Cuon alpinus</i>) and the African hunting dog (<i>Lycaon pictus</i>). There is evidence of gene flow between <a href="/wiki/African_golden_wolves" class="mw-redirect" title="African golden wolves">African golden wolves</a>, <a href="/wiki/Golden_jackals" class="mw-redirect" title="Golden jackals">golden jackals</a>, and grey wolves. The study suggests that the African golden wolf is a descendant of a genetically <a href="/wiki/Genetic_admixture" title="Genetic admixture">admixed</a> canid of 72% grey <a href="/wiki/Wolf" title="Wolf">wolf</a> and 28% <a href="/wiki/Ethiopian_wolf" title="Ethiopian wolf">Ethiopian wolf</a> ancestry, and that the Ethiopian wolf once had a wider range in Africa. One African golden wolf from the Egyptian <a href="/wiki/Sinai_Peninsula" title="Sinai Peninsula">Sinai Peninsula</a> showed high admixture with the Middle Eastern grey wolves and dogs, highlighting the role of the land bridge between the African and Eurasian continents in canid evolution. There is evidence of gene flow between golden jackals and Middle Eastern wolves, less so with European and Asian wolves, and least with North American wolves. The study proposes that the golden jackal ancestry found in North American wolves may have occurred before the divergence of the Eurasian and North American grey wolves. The study indicates that the common ancestor of the <a href="/wiki/Coyote" title="Coyote">coyote</a> and grey wolf has genetically <a href="/wiki/Genetic_admixture" title="Genetic admixture">admixed</a> with a <a href="/wiki/Ghost_population" title="Ghost population">ghost population</a> of an extinct unidentified canid. The canid is genetically close to the <a href="/wiki/Dhole" title="Dhole">dhole</a> and has evolved after the divergence of the African hunting dog from the other canid species. The <a href="/wiki/Basal_(phylogenetics)" title="Basal (phylogenetics)">basal</a> position of the coyote compared to the wolf is proposed to be due to the coyote retaining more of the mitochondrial genome of this unknown canid.<sup id="cite_ref-gopalakrishnan2018_119-2" class="reference"><a href="#cite_note-gopalakrishnan2018-119"><span class="cite-bracket">&#91;</span>119<span class="cite-bracket">&#93;</span></a></sup> </p><p>A genomic study on the wolves of China included museum specimens of wolves from southern China that were collected between 1963 and 1988. The wolves in the study formed 3 clades: north Asian wolves that included those from northern China and eastern Russia, <a href="/wiki/Himalayan_wolf" title="Himalayan wolf">Himalayan wolves</a> from the Tibetan Plateau, and a unique population from southern China. One specimen located as far southeast as <a href="/wiki/Jiangxi" title="Jiangxi">Jiangxi</a> province shows evidence of being admixed between Tibetan-related wolves and other wolves in China. One specimen from <a href="/wiki/Zhejiang" title="Zhejiang">Zhejiang</a> province in eastern China shared gene flow with the wolves from southern China, however its genome was 12-14 percent admixed with a canid that may be the dhole or an unknown canid that predates the genetic divergence of the dhole. The wolf population from southern China is believed to be still existing in that region.<sup id="cite_ref-Wang2019_133-0" class="reference"><a href="#cite_note-Wang2019-133"><span class="cite-bracket">&#91;</span>132<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Out_of_Beringia">Out of Beringia</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=20" title="Edit section: Out of Beringia"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Grey wolves suffered a species-wide <a href="/wiki/Population_bottleneck" title="Population bottleneck">population bottleneck</a> (reduction) approximately 25,000 YBP during the Last Glacial Maximum. This was followed by a single population of modern wolves expanding out of a <a href="/wiki/Beringia#Refugium" title="Beringia">Beringia refuge</a> to repopulate the wolf's former range, replacing the remaining Late Pleistocene wolf populations across Eurasia and North America as they did so.<sup id="cite_ref-Loog2018_15-3" class="reference"><a href="#cite_note-Loog2018-15"><span class="cite-bracket">&#91;</span>15<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Werhahn2018_130-1" class="reference"><a href="#cite_note-Werhahn2018-130"><span class="cite-bracket">&#91;</span>130<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Schweizer2020_131-1" class="reference"><a href="#cite_note-Schweizer2020-131"><span class="cite-bracket">&#91;</span>131<span class="cite-bracket">&#93;</span></a></sup> This source population probably did not give rise to dogs, but admixed with dogs which allowed them to gain coat colour genes that are also related to immunity, and provided dogs with genes which allowed them to adapt to high-altitude environments (e.g. Tibet). This suggests that the genetic divergence of European and East Asian dogs could be based on admixture with different sub-populations of wolves.<sup id="cite_ref-Schweizer2020_131-2" class="reference"><a href="#cite_note-Schweizer2020-131"><span class="cite-bracket">&#91;</span>131<span class="cite-bracket">&#93;</span></a></sup> </p><p>There is little genetic information available on the ancient wolves that existed prior to the bottleneck. However, studies show that one or more of these ancient populations is more directly ancestral to dogs than are modern wolves, and conceivably these were more prone to domestication by the first humans to invade Eurasia.<sup id="cite_ref-Schweizer2020_131-3" class="reference"><a href="#cite_note-Schweizer2020-131"><span class="cite-bracket">&#91;</span>131<span class="cite-bracket">&#93;</span></a></sup> </p><p>As of 2020, the oldest known intact wolf remains belongs to a mummified pup dated 56,000 YBP that was recovered from the permafrost along a small tributary of Last Chance Creek near <a href="/wiki/Dawson_City" title="Dawson City">Dawson City</a>, Yukon, Canada. A DNA analysis showed that it belonged to the Beringian wolf clade, that the <a href="/wiki/Most_recent_common_ancestor" title="Most recent common ancestor">most recent common ancestor</a> of this clade dates to 86,700–67,500 YBP, and that this clade was basal to all other wolves except for the Himalayan wolf.<sup id="cite_ref-meachen2020_134-0" class="reference"><a href="#cite_note-meachen2020-134"><span class="cite-bracket">&#91;</span>133<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Into_America_and_Japan">Into America and Japan</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=21" title="Edit section: Into America and Japan"><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">See also: <a href="/wiki/Japanese_wolf" title="Japanese wolf">Japanese wolf</a> and <a href="/wiki/Ezo_wolf" class="mw-redirect" title="Ezo wolf">Ezo wolf</a></div> <figure class="mw-halign-right" typeof="mw:File"><a href="/wiki/File:Beringia_land_bridge-noaagov.gif" class="mw-file-description" title="Shrinking of the Bering land bridge"><img alt="Shrinking of the Bering land bridge" src="//upload.wikimedia.org/wikipedia/commons/thumb/5/52/Beringia_land_bridge-noaagov.gif/320px-Beringia_land_bridge-noaagov.gif" decoding="async" width="320" height="209" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/52/Beringia_land_bridge-noaagov.gif/480px-Beringia_land_bridge-noaagov.gif 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/52/Beringia_land_bridge-noaagov.gif/640px-Beringia_land_bridge-noaagov.gif 2x" data-file-width="700" data-file-height="458" /></a><figcaption>Shrinking of the Bering land bridge</figcaption></figure> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Japan_glaciation.gif" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d9/Japan_glaciation.gif/220px-Japan_glaciation.gif" decoding="async" width="220" height="248" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d9/Japan_glaciation.gif/330px-Japan_glaciation.gif 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d9/Japan_glaciation.gif/440px-Japan_glaciation.gif 2x" data-file-width="1010" data-file-height="1137" /></a><figcaption>Japanese archipelago 20,000 years ago with Hokkaido island bridged to the mainland, thin black line indicates present-day shoreline</figcaption></figure> <p>In 2016, a study built on the work of another major study<sup id="cite_ref-thalmann2013_73-3" class="reference"><a href="#cite_note-thalmann2013-73"><span class="cite-bracket">&#91;</span>73<span class="cite-bracket">&#93;</span></a></sup> and analyzed the sequences of 12 genes that are located on the <a href="/wiki/Heavy_strand" title="Heavy strand">heavy strand</a> of the mitochondrial genome of extinct and modern <i>C. lupus</i>. The study excluded the sequences of the divergent <a href="/wiki/Himalayan_wolf" title="Himalayan wolf">Himalayan wolf</a> and the <a href="/wiki/Indian_wolf#Canis_indica" title="Indian wolf">Indian grey wolf</a>. The ancient specimens were <a href="/wiki/Radiocarbon_dated" class="mw-redirect" title="Radiocarbon dated">radiocarbon dated</a> and <a href="/wiki/Stratigraphy" title="Stratigraphy">stratagraphically</a> dated, and together with the sequences generated a time-based phylogenetic tree. From the tree, the study was able to infer the most recent common ancestor for all other <i>C. lupus</i> specimens – modern and extinct – was 80,000 YBP and this date concurred with the earlier study.<sup id="cite_ref-thalmann2013_73-4" class="reference"><a href="#cite_note-thalmann2013-73"><span class="cite-bracket">&#91;</span>73<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-koblmuller2016_129-1" class="reference"><a href="#cite_note-koblmuller2016-129"><span class="cite-bracket">&#91;</span>129<span class="cite-bracket">&#93;</span></a></sup> The study could find no evidence of a population bottleneck for wolves until a few thousand years ago.<sup id="cite_ref-koblmuller2016_129-2" class="reference"><a href="#cite_note-koblmuller2016-129"><span class="cite-bracket">&#91;</span>129<span class="cite-bracket">&#93;</span></a></sup> </p><p>The phylogenetic tree showed the <a href="/wiki/Polyphyly" title="Polyphyly">polyphyly</a> of American wolves, the Mexican wolf was divergent from other North American wolves, and these other North American wolves formed two closely related clades. A scenario consistent with the phylogenetic, ice sheet and sea-level data was that during the Ice Age when sea levels were at their lowest, there was a single wave of wolf colonization into North America starting with the opening of the <a href="/wiki/Bering_land_bridge" class="mw-redirect" title="Bering land bridge">Bering land bridge</a> 70,000 YBP and closing during the <a href="/wiki/Late_Glacial_Maximum" class="mw-redirect" title="Late Glacial Maximum">Late Glacial Maximum</a> of the Yukon corridor that ran through the division between the <a href="/wiki/Laurentide_Ice_Sheet" class="mw-redirect" title="Laurentide Ice Sheet">Laurentide Ice Sheet</a> and the <a href="/wiki/Cordilleran_Ice_Sheet" class="mw-redirect" title="Cordilleran Ice Sheet">Cordilleran Ice Sheet</a> 23,000 YBP. Mexican wolves were part of the single wave and either diverged from the other wolves before entering North America or once in North America due to the change in its environment. </p><p>As wolves had been in the fossil record of North America but modern wolves could trace their ancestry back only 80,000 years, the wolf haplotypes that were already in North America were replaced by these invaders, either through competitive displacement or through admixture.<sup id="cite_ref-koblmuller2016_129-3" class="reference"><a href="#cite_note-koblmuller2016-129"><span class="cite-bracket">&#91;</span>129<span class="cite-bracket">&#93;</span></a></sup> The replacement in North America of a basal population of wolves by a more recent one supported the findings of earlier studies.<sup id="cite_ref-pilot2010_126-1" class="reference"><a href="#cite_note-pilot2010-126"><span class="cite-bracket">&#91;</span>126<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-randi2011_135-0" class="reference"><a href="#cite_note-randi2011-135"><span class="cite-bracket">&#91;</span>134<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-leonard2007_125-1" class="reference"><a href="#cite_note-leonard2007-125"><span class="cite-bracket">&#91;</span>125<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-koblmuller2016_129-4" class="reference"><a href="#cite_note-koblmuller2016-129"><span class="cite-bracket">&#91;</span>129<span class="cite-bracket">&#93;</span></a></sup> There possibly existed a panmictic wolf population with gene flow spanning Eurasia and North America until the closing of the ice sheets.<sup id="cite_ref-pilot2010_126-2" class="reference"><a href="#cite_note-pilot2010-126"><span class="cite-bracket">&#91;</span>126<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-hofreiter2007_136-0" class="reference"><a href="#cite_note-hofreiter2007-136"><span class="cite-bracket">&#91;</span>135<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-koblmuller2016_129-5" class="reference"><a href="#cite_note-koblmuller2016-129"><span class="cite-bracket">&#91;</span>129<span class="cite-bracket">&#93;</span></a></sup> Once the sheets closed, the southern wolves were isolated and north of the sheets only the <a href="/wiki/Beringian_wolf" title="Beringian wolf">Beringian wolf</a> existed. The land bridge became inundated by the sea 10,000 YBP, the sheets receded 12,000–6,000 YBP, the Beringian wolf went extinct and the southern wolves expanded to recolonize the rest of North America. All North American wolves are descended from those that were once isolated south of the ice sheets. However, much of their diversity was later lost during the twentieth century.<sup id="cite_ref-koblmuller2016_129-6" class="reference"><a href="#cite_note-koblmuller2016-129"><span class="cite-bracket">&#91;</span>129<span class="cite-bracket">&#93;</span></a></sup> </p><p>Studies using <a href="/wiki/Mitochondrial_DNA" title="Mitochondrial DNA">mitochondrial DNA</a> have indicated that the wolves of coastal south-east Alaska are genetically distinct from inland grey wolves, reflecting a pattern also observed in other taxa. They show a phylogenetic relationship with extirpated wolves from the south (Oklahoma), indicating that these wolves are the last remains of a once widespread group that has been largely extirpated during the last century, and that the wolves of northern North America had originally expanded from southern refuges below the <a href="/wiki/Wisconsin_glaciation" title="Wisconsin glaciation">Wisconsin glaciation</a> after the ice had melted at the end of the <a href="/wiki/Last_Glacial_Maximum" title="Last Glacial Maximum">Last Glacial Maximum</a>.<sup id="cite_ref-weckworth2005_137-0" class="reference"><a href="#cite_note-weckworth2005-137"><span class="cite-bracket">&#91;</span>136<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-weckworth2010_138-0" class="reference"><a href="#cite_note-weckworth2010-138"><span class="cite-bracket">&#91;</span>137<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-weckworth2011_139-0" class="reference"><a href="#cite_note-weckworth2011-139"><span class="cite-bracket">&#91;</span>138<span class="cite-bracket">&#93;</span></a></sup> A <a href="/wiki/Whole_genome_sequencing" title="Whole genome sequencing">whole-genome</a> DNA study indicated that all North American wolves were <a href="/wiki/Monophyletic" class="mw-redirect" title="Monophyletic">monophyletic</a> and therefore are the descendants of a common ancestor.<sup id="cite_ref-vonholdt2016_140-0" class="reference"><a href="#cite_note-vonholdt2016-140"><span class="cite-bracket">&#91;</span>139<span class="cite-bracket">&#93;</span></a></sup> </p><p>During the same period, the <a href="/wiki/Soya_Strait" class="mw-redirect" title="Soya Strait">Soya Strait</a> between <a href="/wiki/Hokkaido" title="Hokkaido">Hokkaido</a> and <a href="/wiki/Sakhalin_Island" class="mw-redirect" title="Sakhalin Island">Sakhalin Island</a> was dry for 75,000 years and it was proposed that the extinct <a href="/wiki/Ezo_Wolf" class="mw-redirect" title="Ezo Wolf">Ezo wolf</a> (<i>C. l. hattai</i>) arrived on Hokkaido from Sakhalin.<sup id="cite_ref-matsumura2014_141-0" class="reference"><a href="#cite_note-matsumura2014-141"><span class="cite-bracket">&#91;</span>140<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-koblmuller2016_129-7" class="reference"><a href="#cite_note-koblmuller2016-129"><span class="cite-bracket">&#91;</span>129<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-abe1999_142-0" class="reference"><a href="#cite_note-abe1999-142"><span class="cite-bracket">&#91;</span>141<span class="cite-bracket">&#93;</span></a></sup> However, the sequences indicated that it arrived in Hokkaido less than 10,000 YBP. The Ezo wolf was closely related to one of the North American clades,<sup id="cite_ref-matsumura2014_141-1" class="reference"><a href="#cite_note-matsumura2014-141"><span class="cite-bracket">&#91;</span>140<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-koblmuller2016_129-8" class="reference"><a href="#cite_note-koblmuller2016-129"><span class="cite-bracket">&#91;</span>129<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-ishiguro2010_143-0" class="reference"><a href="#cite_note-ishiguro2010-143"><span class="cite-bracket">&#91;</span>142<span class="cite-bracket">&#93;</span></a></sup> but different from the more southerly <a href="/wiki/Japanese_wolf" title="Japanese wolf">Japanese wolf</a> (<i>C. l. hodophilax</i>) that was basal to modern wolves.<sup id="cite_ref-matsumura2014_141-2" class="reference"><a href="#cite_note-matsumura2014-141"><span class="cite-bracket">&#91;</span>140<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-koblmuller2016_129-9" class="reference"><a href="#cite_note-koblmuller2016-129"><span class="cite-bracket">&#91;</span>129<span class="cite-bracket">&#93;</span></a></sup> The Japanese wolf inhabited Kyushu, Shikoku, and Honshu islands<sup id="cite_ref-walker2008_144-0" class="reference"><a href="#cite_note-walker2008-144"><span class="cite-bracket">&#91;</span>143<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-ishiguro2009_145-0" class="reference"><a href="#cite_note-ishiguro2009-145"><span class="cite-bracket">&#91;</span>144<span class="cite-bracket">&#93;</span></a></sup> but not Hokkaido Island.<sup id="cite_ref-ishiguro2009_145-1" class="reference"><a href="#cite_note-ishiguro2009-145"><span class="cite-bracket">&#91;</span>144<span class="cite-bracket">&#93;</span></a></sup> This indicates that its ancestor may have migrated from the Asian continent through the Korean Peninsula into Japan.<sup id="cite_ref-matsumura2014_141-3" class="reference"><a href="#cite_note-matsumura2014-141"><span class="cite-bracket">&#91;</span>140<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-ishiguro2009_145-2" class="reference"><a href="#cite_note-ishiguro2009-145"><span class="cite-bracket">&#91;</span>144<span class="cite-bracket">&#93;</span></a></sup> The past sea levels of the <a href="/wiki/Korean_Strait" class="mw-redirect" title="Korean Strait">Korean Strait</a> together with the timing of the Japanese wolf sequences indicated that it arrived to the southern islands less than 20,000 YBP.<sup id="cite_ref-koblmuller2016_129-10" class="reference"><a href="#cite_note-koblmuller2016-129"><span class="cite-bracket">&#91;</span>129<span class="cite-bracket">&#93;</span></a></sup> In 2020, a genomic study found that the Japanese wolf was the last of the Siberian Pleistocene wolves, which were thought to have gone extinct at the end of the Late Pleistocene (11,700 years ago). Some of these had survived into the 20th Century and had admixed with Japanese dogs.<sup id="cite_ref-Niemann2020_146-0" class="reference"><a href="#cite_note-Niemann2020-146"><span class="cite-bracket">&#91;</span>145<span class="cite-bracket">&#93;</span></a></sup> </p><p>The dog was a very successful invader of North America and had established a widespread <a href="/wiki/Ecological_niche" title="Ecological niche">ecological niche</a> by the Early–Middle Holocene. There was no overlap in niche between the dog and the wolf in comparison to the dog and other North American canids. By the Late Holocene, the dog's niche area was less in size than researchers had expected to find, indicating that it was limited by <a href="/wiki/Biotic_component" class="mw-redirect" title="Biotic component">biotic factors</a>. These regions include the northeast and northwest of the United States that correlate with the greatest densities of early human occupation, indicating that the dog had "defected" from the wolf niche to the human niche and explains why the dog's niche area was not as large as expected. The separation between dog and wolf may reflect the rapid rate in which domestication occurred,<sup id="cite_ref-pardi2016_147-0" class="reference"><a href="#cite_note-pardi2016-147"><span class="cite-bracket">&#91;</span>146<span class="cite-bracket">&#93;</span></a></sup> including the possibility of a second domestication event occurring in North America.<sup id="cite_ref-witt2015_148-0" class="reference"><a href="#cite_note-witt2015-148"><span class="cite-bracket">&#91;</span>147<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-pardi2016_147-1" class="reference"><a href="#cite_note-pardi2016-147"><span class="cite-bracket">&#91;</span>146<span class="cite-bracket">&#93;</span></a></sup> Packs of wolves and hunter-gatherers hunt similar prey in a similar way within a similar group social structure that may have facilitated wolf domestication.<sup id="cite_ref-hall1978_149-0" class="reference"><a href="#cite_note-hall1978-149"><span class="cite-bracket">&#91;</span>148<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-olsen1985_150-0" class="reference"><a href="#cite_note-olsen1985-150"><span class="cite-bracket">&#91;</span>149<span class="cite-bracket">&#93;</span></a></sup> </p><p>The wolf was exterminated in the southern part of their historic geographical range in North America by the middle of the 20th century. An mDNA study of 34 wolf remains from North America dated between 1856 and 1915 found their genetic diversity to be twice that of modern wolves in these regions, and two thirds of the haplotypes identified were unique. These results indicate that a historic population of several hundred thousand wolves once existed in Mexico and the western US.<sup id="cite_ref-leonard2004_151-0" class="reference"><a href="#cite_note-leonard2004-151"><span class="cite-bracket">&#91;</span>150<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-miklosi2015_128-1" class="reference"><a href="#cite_note-miklosi2015-128"><span class="cite-bracket">&#91;</span>128<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Divergence_with_the_coyote">Divergence with the coyote</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=22" title="Edit section: Divergence with the coyote"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In 1993, a study proposed that the wolves of North America display skull traits more similar towards the coyote than those wolves from Eurasia.<sup id="cite_ref-goulet1993_47-1" class="reference"><a href="#cite_note-goulet1993-47"><span class="cite-bracket">&#91;</span>47<span class="cite-bracket">&#93;</span></a></sup> In 2016, a <a href="/wiki/Whole_genome_sequencing" title="Whole genome sequencing">whole-genome</a> DNA study proposed, based on the assumptions made, that all of the North American wolves and coyotes diverged from a common ancestor less than 6,000–117,000 years ago. The study also indicated that all North American wolves have a significant amount of coyote ancestry and all coyotes some degree of wolf ancestry, and that the <a href="/wiki/Red_wolf" title="Red wolf">red wolf</a> and <a href="/wiki/Eastern_wolf" title="Eastern wolf">eastern wolf</a> are highly <a href="/wiki/Genetic_admixture" title="Genetic admixture">admixed</a> with different proportions of grey wolf and coyote ancestry. However, the test that was used cannot differentiate between ancient and recent hybridization. One test indicated a wolf/coyote divergence time of 51,000 years before present that matched other studies indicating that the extant wolf came into being around this time. Another test indicated that the red wolf diverged from the coyote between 55,000 and 117,000 years before present and the Great Lakes region wolf 32,000 years before present. But a 2000 study on the haplotypes of North American canids showed that the divergences between coyotes, grey wolves, red wolves, and Algonquin (eastern) wolves happened much earlier. This study showed that there was a 3.2&#160;% divergence between the eastern Canadian wolf and coyote gene sequences, and a 2.3&#160;% sequence divergence between the red wolf and coyote haplotypes. However, the divergence was a much greater 8.0% between gray wolf (C. lupus) mtDNA and eastern/red wolf haplotypes, and 10.0% between gray wolf and coyote haplotypes. The sequence difference observed between eastern Canadian wolf sequences and coyote sequences is consistent with a separation of 150 000 – 300 000 years, using a divergence rate of 1–2% per 100 000 years.<sup id="cite_ref-152" class="reference"><a href="#cite_note-152"><span class="cite-bracket">&#91;</span>151<span class="cite-bracket">&#93;</span></a></sup> Additionally, conflicting with the recent divergent hypothesis, the fossil record that indicates a coyote-like specimen dated to 1 million years before present.<sup id="cite_ref-wang2008_36-1" class="reference"><a href="#cite_note-wang2008-36"><span class="cite-bracket">&#91;</span>36<span class="cite-bracket">&#93;</span></a></sup> </p><p>The modern grey wolf expanded out of Beringia 25,000 years ago.<sup id="cite_ref-Schweizer2020_131-4" class="reference"><a href="#cite_note-Schweizer2020-131"><span class="cite-bracket">&#91;</span>131<span class="cite-bracket">&#93;</span></a></sup> The modern coyote appeared around 10,000 years ago, and most likely descended from the <a href="/wiki/Pleistocene_coyote" title="Pleistocene coyote">Pleistocene coyote</a>. The most genetically basal coyote mDNA clade pre-dates the <a href="/wiki/Last_Glacial_Maximum" title="Last Glacial Maximum">Last Glacial Maximum</a> and is a haplotype that can only be found in the Eastern wolf. This implies that the large, wolf-like <a href="/wiki/Pleistocene_coyote" title="Pleistocene coyote">Pleistocene coyote</a> was the ancestor of the <a href="/wiki/Eastern_wolf" title="Eastern wolf">Eastern wolf</a> and <a href="/wiki/Red_wolf" title="Red wolf">Red wolf</a>, which diverged from each other later in time. </p> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Pleistocene_coyote" title="Pleistocene coyote">Pleistocene coyote</a></div> <div class="mw-heading mw-heading3"><h3 id="Domestic_dog">Domestic dog</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=23" title="Edit section: Domestic dog"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Origin_of_the_domestic_dog" class="mw-redirect" title="Origin of the domestic dog">Origin of the domestic dog</a></div> <p>The domestic dog (<i>Canis lupus familiaris</i>) is the most widely abundant large carnivore.<sup id="cite_ref-thalmann2013_73-5" class="reference"><a href="#cite_note-thalmann2013-73"><span class="cite-bracket">&#91;</span>73<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-fan2016_13-2" class="reference"><a href="#cite_note-fan2016-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-vila1999_153-0" class="reference"><a href="#cite_note-vila1999-153"><span class="cite-bracket">&#91;</span>152<span class="cite-bracket">&#93;</span></a></sup> Over the past million years, numerous wolf-like forms existed but their turnover has been high, and modern wolves are not the lineal ancestors of dogs.<sup id="cite_ref-thalmann2013_73-6" class="reference"><a href="#cite_note-thalmann2013-73"><span class="cite-bracket">&#91;</span>73<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-freedman2014_12-4" class="reference"><a href="#cite_note-freedman2014-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-fan2016_13-3" class="reference"><a href="#cite_note-fan2016-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-leonard2004_151-1" class="reference"><a href="#cite_note-leonard2004-151"><span class="cite-bracket">&#91;</span>150<span class="cite-bracket">&#93;</span></a></sup> Although research had suggested that dogs and wolves were genetically very close relatives,<sup id="cite_ref-vila1997_99-3" class="reference"><a href="#cite_note-vila1997-99"><span class="cite-bracket">&#91;</span>99<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-wayne1999_100-2" class="reference"><a href="#cite_note-wayne1999-100"><span class="cite-bracket">&#91;</span>100<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-wayne1993_109-1" class="reference"><a href="#cite_note-wayne1993-109"><span class="cite-bracket">&#91;</span>109<span class="cite-bracket">&#93;</span></a></sup> later <a href="/wiki/Phylogenetic_analysis" class="mw-redirect" title="Phylogenetic analysis">phylogenetic analysis</a> strongly supported the hypothesis that dogs and wolves are reciprocally <a href="/wiki/Monophyly" title="Monophyly">monophylic</a> <a href="/wiki/Taxa" class="mw-redirect" title="Taxa">taxa</a> that form two sister <a href="/wiki/Clades" class="mw-redirect" title="Clades">clades</a>.<sup id="cite_ref-vila1997_99-4" class="reference"><a href="#cite_note-vila1997-99"><span class="cite-bracket">&#91;</span>99<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-freedman2014_12-5" class="reference"><a href="#cite_note-freedman2014-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-larson2014_154-0" class="reference"><a href="#cite_note-larson2014-154"><span class="cite-bracket">&#91;</span>153<span class="cite-bracket">&#93;</span></a></sup> This suggests that none of the modern wolf populations are related to the wolves that were first domesticated and the wolf ancestor of dogs is therefore presumed extinct.<sup id="cite_ref-freedman2014_12-6" class="reference"><a href="#cite_note-freedman2014-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-larson2014_154-1" class="reference"><a href="#cite_note-larson2014-154"><span class="cite-bracket">&#91;</span>153<span class="cite-bracket">&#93;</span></a></sup> Recent <a href="/wiki/Mitochondrial_DNA" title="Mitochondrial DNA">mitochondrial DNA</a> analyses of ancient and modern grey wolf specimens supports a pattern of population reduction and turnover.<sup id="cite_ref-thalmann2013_73-7" class="reference"><a href="#cite_note-thalmann2013-73"><span class="cite-bracket">&#91;</span>73<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-pilot2010_126-3" class="reference"><a href="#cite_note-pilot2010-126"><span class="cite-bracket">&#91;</span>126<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-leonard2007_125-2" class="reference"><a href="#cite_note-leonard2007-125"><span class="cite-bracket">&#91;</span>125<span class="cite-bracket">&#93;</span></a></sup> An alternate proposal is that during the ecological upheavals of the Late Pleistocene all of the remaining members of a dwindling lineage joined humans.<sup id="cite_ref-morey2016_155-0" class="reference"><a href="#cite_note-morey2016-155"><span class="cite-bracket">&#91;</span>154<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-morey2015_74-2" class="reference"><a href="#cite_note-morey2015-74"><span class="cite-bracket">&#91;</span>74<span class="cite-bracket">&#93;</span></a></sup> </p><p>In 2016, a study investigated for the first time the population subdivisions, demography, and the relationships of grey wolves based on their <a href="/wiki/Whole_genome_sequencing" title="Whole genome sequencing">whole-genome sequences</a>. The study indicated that the dog was a divergent subspecies of the grey wolf and was derived from a now-extinct <a href="/wiki/Ghost_population" title="Ghost population">ghost population</a> of Late Pleistocene wolves,<sup id="cite_ref-thalmann2013_73-8" class="reference"><a href="#cite_note-thalmann2013-73"><span class="cite-bracket">&#91;</span>73<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-freedman2014_12-7" class="reference"><a href="#cite_note-freedman2014-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-fan2016_13-4" class="reference"><a href="#cite_note-fan2016-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> and the dog and the dingo are not separate species.<sup id="cite_ref-fan2016_13-5" class="reference"><a href="#cite_note-fan2016-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> The genome-wide <a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">phylogenetic tree</a> indicated a <a href="/wiki/Genetic_divergence" title="Genetic divergence">genetic divergence</a> between New World and Old World wolves, which was then followed by a divergence between the dog and Old World wolves 27,000YBP<sup id="cite_ref-skoglund2015_14-2" class="reference"><a href="#cite_note-skoglund2015-14"><span class="cite-bracket">&#91;</span>14<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-fan2016_13-6" class="reference"><a href="#cite_note-fan2016-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> – 29,000 YBP.<sup id="cite_ref-fan2016_13-7" class="reference"><a href="#cite_note-fan2016-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> The dog forms a sister taxon with Eurasian grey wolves but not North American wolves. The dog had considerable pre-ancestry after its divergence from the Old World wolves before it separated into distinct lineages that are nearly as distinct from one another as they are from wolves.<sup id="cite_ref-fan2016_13-8" class="reference"><a href="#cite_note-fan2016-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> The study suggested that previous datings based on the divergence between wolves and coyotes of one million years ago using fossils of what appeared to be coyote-like specimens may not reflect the ancestry of the modern forms.<sup id="cite_ref-koepfli2015_98-6" class="reference"><a href="#cite_note-koepfli2015-98"><span class="cite-bracket">&#91;</span>98<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-freedman2014_12-8" class="reference"><a href="#cite_note-freedman2014-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-skoglund2015_14-3" class="reference"><a href="#cite_note-skoglund2015-14"><span class="cite-bracket">&#91;</span>14<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-fan2016_13-9" class="reference"><a href="#cite_note-fan2016-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> </p> <table style="border: 1px solid #ccc; vertical-align: middle; width: 248px; margin:0.2em 0 0.2em 1em; float:right; clear:right;"> <tbody><tr> <td style="text-align: center; font-weight: bold; border: 0; padding: 0;">Grey wolf divergence and timing in years before present </td></tr> <tr> <td style="padding: 5px"> <div class="clade"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade" style="font-size:75%; line-height:75%;width:550px;"> <tbody><tr> <td class="clade-label first">wolf/dog&#160;ancestor </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first">New&#160;World&#160;clade </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p>Mexico <span typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_IV)_C._l._baileyi_mod.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e2/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IV%29_C._l._baileyi_mod.jpg/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IV%29_C._l._baileyi_mod.jpg" decoding="async" width="50" height="39" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e2/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IV%29_C._l._baileyi_mod.jpg/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IV%29_C._l._baileyi_mod.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e2/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IV%29_C._l._baileyi_mod.jpg/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IV%29_C._l._baileyi_mod.jpg 2x" data-file-width="465" data-file-height="363" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel">13,000 </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <p>North America/Hokkaido <span typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_V).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/3c/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_V%29.jpg/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_V%29.jpg" decoding="async" width="50" height="35" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/3c/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_V%29.jpg/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_V%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/3c/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_V%29.jpg/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_V%29.jpg 2x" data-file-width="491" data-file-height="344" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel">31,000 </td></tr> <tr> <td class="clade-label">Old&#160;World&#160;clade </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first">Old&#160;World&#160;wolves </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p>Asian highland <span typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_III).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f9/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_III%29.jpg/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_III%29.jpg" decoding="async" width="50" height="39" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f9/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_III%29.jpg/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_III%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f9/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_III%29.jpg/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_III%29.jpg 2x" data-file-width="465" data-file-height="363" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <p>Asian lowland <span typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_IV).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/ca/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IV%29.jpg/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IV%29.jpg" decoding="async" width="50" height="39" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/ca/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IV%29.jpg/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IV%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/ca/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IV%29.jpg/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IV%29.jpg 2x" data-file-width="459" data-file-height="357" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel">27,500 </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p>Middle East <span typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_IV)_C._l._arabs_mod.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/45/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IV%29_C._l._arabs_mod.jpg/40px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IV%29_C._l._arabs_mod.jpg" decoding="async" width="40" height="31" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/45/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IV%29_C._l._arabs_mod.jpg/60px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IV%29_C._l._arabs_mod.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/45/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IV%29_C._l._arabs_mod.jpg/80px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_IV%29_C._l._arabs_mod.jpg 2x" data-file-width="459" data-file-height="357" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <p>Europe <span typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_I).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c4/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.jpg/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.jpg" decoding="async" width="50" height="31" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c4/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.jpg/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c4/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.jpg/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_I%29.jpg 2x" data-file-width="500" data-file-height="314" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label">Ancestral&#160;dog </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p>Dog <span style="display: inline-block; transform: scaleX(-1);"><span typeof="mw:File"><a href="/wiki/File:202104_Dog.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f3/202104_Dog.svg/50px-202104_Dog.svg.png" decoding="async" width="50" height="38" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f3/202104_Dog.svg/75px-202104_Dog.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f3/202104_Dog.svg/100px-202104_Dog.svg.png 2x" data-file-width="512" data-file-height="384" /></a></span></span> </p> </td></tr> <tr> <td class="clade-slabel">25,700 </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <p>Dingo <span typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_XXXVII).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/1e/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XXXVII%29.jpg/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XXXVII%29.jpg" decoding="async" width="50" height="41" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/1e/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XXXVII%29.jpg/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XXXVII%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/1e/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XXXVII%29.jpg/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XXXVII%29.jpg 2x" data-file-width="474" data-file-height="388" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <p>Basenji <span typeof="mw:File"><a href="/wiki/File:Dogs,_jackals,_wolves,_and_foxes_(Plate_XXXVII).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/1e/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XXXVII%29.jpg/50px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XXXVII%29.jpg" decoding="async" width="50" height="41" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/1e/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XXXVII%29.jpg/75px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XXXVII%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/1e/Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XXXVII%29.jpg/100px-Dogs%2C_jackals%2C_wolves%2C_and_foxes_%28Plate_XXXVII%29.jpg 2x" data-file-width="474" data-file-height="388" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last">29,000 </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td style="text-align: left; font-size: 88%; border: 0; padding: 0;">Whole-genome <a href="/wiki/Phylogenetic_analysis" class="mw-redirect" title="Phylogenetic analysis">phylogenetic</a> tree of modern grey wolf populations.<sup id="cite_ref-fan2016_13-10" class="reference"><a href="#cite_note-fan2016-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> </td></tr></tbody></table> <p>The study indicated that the <a href="/wiki/Mexican_wolf" title="Mexican wolf">Mexican wolf</a> was also a divergent form of grey wolf, suggesting that may have been part of an early invasion into North America.<sup id="cite_ref-fan2016_13-11" class="reference"><a href="#cite_note-fan2016-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-leonard2004_151-2" class="reference"><a href="#cite_note-leonard2004-151"><span class="cite-bracket">&#91;</span>150<span class="cite-bracket">&#93;</span></a></sup> The Tibetan wolf was found to be the most highly divergent of the Old World wolves, had suffered a historical population bottleneck and had only recently recolonized the Tibetan Plateau. Glaciation may have caused its habitat loss, genetic isolation then local adaption.<sup id="cite_ref-fan2016_13-12" class="reference"><a href="#cite_note-fan2016-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> </p><p>The study indicated that there has been extensive <a href="/wiki/Genetic_admixture" title="Genetic admixture">genetic admixture</a> between domestic dogs and wolves, with up to 25% of the genome of Old World wolves showing signs of dog ancestry, possibly as the result of <a href="/wiki/Gene_flow" title="Gene flow">gene flow</a> from dogs into wolves that were ancestral to all modern wolves. There was evidence of significant gene flow between the European wolves plus the Israeli wolf with the <a href="/wiki/Basenji" title="Basenji">basenji</a> and <a href="/wiki/Boxer_(Dog)" class="mw-redirect" title="Boxer (Dog)">boxer</a>, which suggests admixture between the lineages ancestral to these breeds and wolf populations.<sup id="cite_ref-freedman2014_12-9" class="reference"><a href="#cite_note-freedman2014-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-fan2016_13-13" class="reference"><a href="#cite_note-fan2016-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> For the lowland Asian wolves: the Central Russian and East Russian wolves and all of the lowland Chinese wolves had significant gene flow with the Chinese indigenous dogs, the <a href="/wiki/Tibetan_Mastiff" title="Tibetan Mastiff">Tibetan Mastiff</a> and the <a href="/wiki/Dingo" title="Dingo">dingo</a>. For the highland Asian wolves: The Tibetan wolves did not show significant admixture with dogs; however, the <a href="/wiki/Qinghai" title="Qinghai">Qinghai</a> wolves had gene flow with the dingo and one of them had gene flow with the Chinese dogs. The New World wolves did not show any gene flow with the boxer, dingo or Chinese indigenous dogs but there was indication of gene flow between the Mexican wolf and the African basenji.<sup id="cite_ref-fan2016_13-14" class="reference"><a href="#cite_note-fan2016-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> All species within the genus <i>Canis</i>, the <a href="/wiki/Canidae#Phylogenetic_relationships" title="Canidae">wolf-like canids</a>, are phylogenetically closely related with 78 <a href="/wiki/Chromosome" title="Chromosome">chromosomes</a> and can potentially <a href="/wiki/Hybrid_(biology)" title="Hybrid (biology)">interbreed</a>.<sup id="cite_ref-wayne1999_100-3" class="reference"><a href="#cite_note-wayne1999-100"><span class="cite-bracket">&#91;</span>100<span class="cite-bracket">&#93;</span></a></sup> There was indication of gene flow into the <a href="/wiki/Golden_jackal" title="Golden jackal">golden jackal</a> from the population ancestral to all wolves and dogs (11.3%–13.6%) and much lower rates (up to 2.8%) from extant wolf populations.<sup id="cite_ref-freedman2014_12-10" class="reference"><a href="#cite_note-freedman2014-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-fan2016_13-15" class="reference"><a href="#cite_note-fan2016-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> </p><p>The data indicated that all wolves shared similar population trajectories, followed by population decline that coincided with the expansion of modern humans worldwide and their technology for capturing large game.<sup id="cite_ref-fan2016_13-16" class="reference"><a href="#cite_note-fan2016-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-vanvalkenburgh2016_156-0" class="reference"><a href="#cite_note-vanvalkenburgh2016-156"><span class="cite-bracket">&#91;</span>155<span class="cite-bracket">&#93;</span></a></sup> Late Pleistocene carnivores would have been social living in large prides, clans and packs in order to hunt the larger game available at that time, and these larger groups would have been more conspicuous targets for human persecutors.<sup id="cite_ref-vanvalkenburgh2016_156-1" class="reference"><a href="#cite_note-vanvalkenburgh2016-156"><span class="cite-bracket">&#91;</span>155<span class="cite-bracket">&#93;</span></a></sup> Large dogs accompanying the humans may have accelerated the rate of decline of carnivores that competed for game,<sup id="cite_ref-fan2016_13-17" class="reference"><a href="#cite_note-fan2016-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-shipman2015_157-0" class="reference"><a href="#cite_note-shipman2015-157"><span class="cite-bracket">&#91;</span>156<span class="cite-bracket">&#93;</span></a></sup> therefore humans expanded across Eurasia, encountered wolves, domesticated some and possibly caused the decline of others.<sup id="cite_ref-fan2016_13-18" class="reference"><a href="#cite_note-fan2016-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> </p><p>The study concluded that admixture had confounded the ability to make inferences about the place of dog domestication. Past studies based on SNPs, genome-wide similarities with Chinese wolves, and lower <a href="/wiki/Linkage_disequilibrium" title="Linkage disequilibrium">linkage disequilibrium</a> might reflect regional admixture between dogs with wolves and gene flow between dog populations, with divergent dog breeds possibly maintaining more wolf ancestry in their genome. The study proposed that analysis of ancient DNA might be a better approach.<sup id="cite_ref-fan2016_13-19" class="reference"><a href="#cite_note-fan2016-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> </p><p>In the same year, a study found that there were only 11 fixed genes that showed variation between wolves and dogs. These genes are thought to affect tameness and emotional processing ability.<sup id="cite_ref-cagan2016_158-0" class="reference"><a href="#cite_note-cagan2016-158"><span class="cite-bracket">&#91;</span>157<span class="cite-bracket">&#93;</span></a></sup> Another study provided a listing of all of the grey wolf and dog mDNA haplotypes combined in the one phylogenetic tree.<sup id="cite_ref-song2016_159-0" class="reference"><a href="#cite_note-song2016-159"><span class="cite-bracket">&#91;</span>158<span class="cite-bracket">&#93;</span></a></sup> </p><p>In 2018, a study compared the sequences of 61,000 <a href="/wiki/Single-nucleotide_polymorphisms" class="mw-redirect" title="Single-nucleotide polymorphisms">Single-nucleotide polymorphisms</a> (<a href="/wiki/Mutations" class="mw-redirect" title="Mutations">mutations</a>) taken from across the genome of grey wolves. The study indicated that there exists individual wolves of dog/wolf ancestry in most of the wolf populations of Eurasia but less so in North America. The hybridization has been occurring across different time scales and was not a recent event. Low-level hybridization did not reduce the wolf distinctiveness.<sup id="cite_ref-pilot2018_160-0" class="reference"><a href="#cite_note-pilot2018-160"><span class="cite-bracket">&#91;</span>159<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Dingo">Dingo</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=24" title="Edit section: Dingo"><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/Canis_lupus_dingo#Lineage" title="Canis lupus dingo">Canis lupus dingo §&#160;Lineage</a></div> <p>The dingo (<i>Canis familiaris dingo</i>) refers to the dog found in <a href="/wiki/Australia" title="Australia">Australia</a>. The dingo is a divergent subspecies of the grey wolf and is not a separate species,<sup id="cite_ref-fan2016_13-20" class="reference"><a href="#cite_note-fan2016-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> and is considered genetically to be a <a href="/wiki/Basal_(phylogenetics)" title="Basal (phylogenetics)">basal</a> member of the domestic dog clade.<sup id="cite_ref-fan2016_13-21" class="reference"><a href="#cite_note-fan2016-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-koepfli2015_98-7" class="reference"><a href="#cite_note-koepfli2015-98"><span class="cite-bracket">&#91;</span>98<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-freedman2014_12-11" class="reference"><a href="#cite_note-freedman2014-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup> The genetic evidence indicates that the dingo originated from East Asian domestic dogs and was introduced through the South-East Asian archipelago into Australia,<sup id="cite_ref-savolainen2004_161-0" class="reference"><a href="#cite_note-savolainen2004-161"><span class="cite-bracket">&#91;</span>160<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-oskarsson2011_162-0" class="reference"><a href="#cite_note-oskarsson2011-162"><span class="cite-bracket">&#91;</span>161<span class="cite-bracket">&#93;</span></a></sup> with a common ancestry between the Australian dingo and the <a href="/wiki/New_Guinea_Singing_Dog" class="mw-redirect" title="New Guinea Singing Dog">New Guinea Singing Dog</a>.<sup id="cite_ref-oskarsson2011_162-1" class="reference"><a href="#cite_note-oskarsson2011-162"><span class="cite-bracket">&#91;</span>161<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-ardalan2012_163-0" class="reference"><a href="#cite_note-ardalan2012-163"><span class="cite-bracket">&#91;</span>162<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Canis_variabilis"><i>Canis variabilis</i></h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=25" title="Edit section: Canis variabilis"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In 2015, a study looked at the <a href="/wiki/MtDNA_control_region" title="MtDNA control region">mitochondrial control region</a> sequences of 13 ancient canid remains and one modern wolf from five sites across Arctic north-east Siberia. The fourteen canids revealed nine mitochondrial <a href="/wiki/Haplotype" title="Haplotype">haplotypes</a>, three of which were on record and the others not reported before. The phylogentic tree generated from the sequences showed that four of the Siberian canids dated 28,000 YBP and one <a href="/wiki/Canis_lupus_variabilis" class="mw-redirect" title="Canis lupus variabilis"><i>Canis c.f. variabilis</i></a> dated 360,000 YBP were highly divergent. The haplotype designated as S805 (28,000 YBP) from the <a href="/wiki/Yana_River" class="mw-redirect" title="Yana River">Yana River</a> was one mutation away from another haplotype S902 (8,000 YBP) that represents Clade A of the modern wolf and domestic dog lineages. Closely related to this haplotype was one that was found in the recently extinct <a href="/wiki/Japanese_wolf" title="Japanese wolf">Japanese wolf</a>. Several ancient haplotypes were oriented around S805, including <i>Canis c.f. variabilis</i> (360,000 YBP), Belgium (36,000 YBP – the "Goyet dog"), Belgium (30,000 YBP), and Konsteki, Russia (22,000 YBP). Given the position of the S805 haplotype on the phylogenetic tree, it may potentially represent a direct link from the <a href="/wiki/Progenitor" title="Progenitor">progenitor</a> (including <i>Canis c.f. variabilis</i>) to the domestic dog and modern wolf lineages. The grey wolf is thought to be ancestral to the domestic dog, however its relationship to <i>C. variabilis</i>, and the genetic contribution of <i>C. variabilis</i> to the dog, is the subject of debate.<sup id="cite_ref-lee2015_164-0" class="reference"><a href="#cite_note-lee2015-164"><span class="cite-bracket">&#91;</span>163<span class="cite-bracket">&#93;</span></a></sup> </p><p>The <a href="/wiki/Zhokhov_Island" title="Zhokhov Island">Zhokhov Island</a> (8,700 YBP) and Aachim (1,700 YBP) canid haplotypes fell within the domestic dog clade, cluster with S805, and also share their haplotypes with – or are one mutation away from – the <a href="/wiki/Tibetan_wolf" class="mw-redirect" title="Tibetan wolf">Tibetan wolf</a> (<i>C. l. filchneri</i>) and the recently extinct Japanese wolf (<i>C. l. hodophilax</i>). This may indicate that these canids retained the genetic signature of admixture with regional wolf populations. Another haplotype designated as S504 (47,000 YBP) from <a href="/wiki/Silene_stenophylla#Duvanny_Yar" title="Silene stenophylla">Duvanny Yar</a> appeared on the phylogenetic tree as not being connected to wolves (both ancient and modern) yet ancestral to dogs, and may represent a genetic source for regional dogs.<sup id="cite_ref-lee2015_164-1" class="reference"><a href="#cite_note-lee2015-164"><span class="cite-bracket">&#91;</span>163<span class="cite-bracket">&#93;</span></a></sup> </p><p>The authors concluded that the structure of the modern dog <a href="/wiki/Gene_pool" title="Gene pool">gene pool</a> was contributed to from ancient Siberian wolves and possibly from <i>Canis c.f. variabilis</i>.<sup id="cite_ref-lee2015_164-2" class="reference"><a href="#cite_note-lee2015-164"><span class="cite-bracket">&#91;</span>163<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Irizarry2018_165-0" class="reference"><a href="#cite_note-Irizarry2018-165"><span class="cite-bracket">&#91;</span>164<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Rise_to_dominant_predator">Rise to dominant predator</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=26" title="Edit section: Rise to dominant predator"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In 2015, a study looked at the paleoecology of large carnivores across the <a href="/wiki/Mammoth_steppe" title="Mammoth steppe">Mammoth steppe</a> during the <a href="/wiki/Late_Pleistocene" title="Late Pleistocene">Late Pleistocene</a> by using stable isotope analysis of their fossil <a href="/wiki/Collagen" title="Collagen">collagen</a> to reconstruct their diets. Based on testing in Belgium, around 40,000 YBP the <a href="/wiki/Cave_hyena" title="Cave hyena">Cave hyenas</a> preyed on mammoth, woolly rhinoceros, horses and reindeer, with <a href="/wiki/Panthera_leo_spelaea" class="mw-redirect" title="Panthera leo spelaea">cave lions</a> taking reindeer and young <a href="/wiki/Cave_bear" title="Cave bear">cave bears</a>. Wolves appear to have been out-competed by <a href="/wiki/Cave_hyena" title="Cave hyena">cave hyenas</a> and had their diet restricted to chamois, giant deer and red deer. However, after the Last Glacial Maximum around 14,000 YBP, wolves had access to all prey species, the cave lion was restricted to reindeer, and the cave hyena had gone extinct.<sup id="cite_ref-bocherens2011_166-0" class="reference"><a href="#cite_note-bocherens2011-166"><span class="cite-bracket">&#91;</span>165<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-yeakel2013_167-0" class="reference"><a href="#cite_note-yeakel2013-167"><span class="cite-bracket">&#91;</span>166<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-bocherens2015_168-0" class="reference"><a href="#cite_note-bocherens2015-168"><span class="cite-bracket">&#91;</span>167<span class="cite-bracket">&#93;</span></a></sup> The data suggests that the extinction of the cave hyena allowed the wolf to become the dominant predator rather than the cave lion, just before the cave lion's extinction.<sup id="cite_ref-bocherens2015_168-1" class="reference"><a href="#cite_note-bocherens2015-168"><span class="cite-bracket">&#91;</span>167<span class="cite-bracket">&#93;</span></a></sup> Another study indicated that the wolf thrived compared to the cave hyena when there was greater snow cover.<sup id="cite_ref-dinnis2016_169-0" class="reference"><a href="#cite_note-dinnis2016-169"><span class="cite-bracket">&#91;</span>168<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Wolf_population_differences">Wolf population differences</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=27" title="Edit section: Wolf population differences"><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">See also: <a href="/wiki/Beringian_wolf" title="Beringian wolf">Beringian wolf</a>, <a href="/wiki/Cave_wolf" title="Cave wolf">Cave wolf</a>, <a href="/wiki/Pleistocene_wolf" title="Pleistocene wolf">Pleistocene wolf</a>, and <a href="/wiki/Paleolithic_dog" title="Paleolithic dog">Paleolithic dog</a></div> <p>The grey wolf <i>Canis lupus</i> is a highly adaptable species that is able to exist in a range of environments and which possesses a wide distribution across the <a href="/wiki/Holarctic" class="mw-redirect" title="Holarctic">Holarctic</a>. Studies of modern grey wolves have identified distinct sub-populations that live in close proximity to each other.<sup id="cite_ref-musiani2007_4-1" class="reference"><a href="#cite_note-musiani2007-4"><span class="cite-bracket">&#91;</span>4<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-schweizer2016a_5-1" class="reference"><a href="#cite_note-schweizer2016a-5"><span class="cite-bracket">&#91;</span>5<span class="cite-bracket">&#93;</span></a></sup> This variation in sub-populations is closely linked to differences in habitat – precipitation, temperature, vegetation, and prey specialization – which affect cranio-dental plasticity.<sup id="cite_ref-geffen2004_6-1" class="reference"><a href="#cite_note-geffen2004-6"><span class="cite-bracket">&#91;</span>6<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-pilot2006_7-1" class="reference"><a href="#cite_note-pilot2006-7"><span class="cite-bracket">&#91;</span>7<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-flower2014_8-1" class="reference"><a href="#cite_note-flower2014-8"><span class="cite-bracket">&#91;</span>8<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-leonard2014_9-1" class="reference"><a href="#cite_note-leonard2014-9"><span class="cite-bracket">&#91;</span>9<span class="cite-bracket">&#93;</span></a></sup> The archaeological and paleontological records show their continuous presence for at least the last 300,000 years.<sup id="cite_ref-sotnikova2010_58-3" class="reference"><a href="#cite_note-sotnikova2010-58"><span class="cite-bracket">&#91;</span>58<span class="cite-bracket">&#93;</span></a></sup> This continuous wolf presence contrasts with <a href="/wiki/Genome" title="Genome">genomic</a> studies, which suggest that all modern wolves and dogs descend from a common ancestral wolf population<sup id="cite_ref-freedman2014_12-12" class="reference"><a href="#cite_note-freedman2014-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-skoglund2015_14-4" class="reference"><a href="#cite_note-skoglund2015-14"><span class="cite-bracket">&#91;</span>14<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-fan2016_13-22" class="reference"><a href="#cite_note-fan2016-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup> that existed as recently as 20,000 years ago.<sup id="cite_ref-freedman2014_12-13" class="reference"><a href="#cite_note-freedman2014-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup> These studies indicate that a <a href="/wiki/Population_bottleneck" title="Population bottleneck">population bottleneck</a> was followed by a rapid radiation from an ancestral population at a time during, or just after, the <a href="/wiki/Last_Glacial_Maximum" title="Last Glacial Maximum">Last Glacial Maximum</a>. This implies that the original wolf populations were out-competed by a new type of wolf which replaced them.<sup id="cite_ref-koblmuller2016_129-11" class="reference"><a href="#cite_note-koblmuller2016-129"><span class="cite-bracket">&#91;</span>129<span class="cite-bracket">&#93;</span></a></sup> However, the geographic origin of this radiation is not known. </p><p>Apart from domestication, humans have harmed the wolf by restricting its habitat through persecution. This has caused a dramatic decrease in its population size over the last two centuries.<sup id="cite_ref-boitani2003_170-0" class="reference"><a href="#cite_note-boitani2003-170"><span class="cite-bracket">&#91;</span>169<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-leonard2005_171-0" class="reference"><a href="#cite_note-leonard2005-171"><span class="cite-bracket">&#91;</span>170<span class="cite-bracket">&#93;</span></a></sup> The shrinking of its habitats that overlap with those of close-relatives such as dogs and coyotes have led to numerous occurrences of hybridization.<sup id="cite_ref-vonholdt2011_172-0" class="reference"><a href="#cite_note-vonholdt2011-172"><span class="cite-bracket">&#91;</span>171<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-vonholdt2013_173-0" class="reference"><a href="#cite_note-vonholdt2013-173"><span class="cite-bracket">&#91;</span>172<span class="cite-bracket">&#93;</span></a></sup> These events, in addition to recent turnovers (extinctions and repopulations by other geneotypes), has made the unravelling of the <a href="/wiki/Phylogeography" title="Phylogeography">phylogeographic</a> history of the wolf difficult.<sup id="cite_ref-ersmark2016_120-1" class="reference"><a href="#cite_note-ersmark2016-120"><span class="cite-bracket">&#91;</span>120<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Ecotypes">Ecotypes</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=28" title="Edit section: Ecotypes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>An <a href="/wiki/Ecotype" title="Ecotype">ecotype</a> is a variant in which the <a href="/wiki/Phenotype" title="Phenotype">phenotypic</a> differences are too few or too subtle to warrant being classified as a subspecies. These can occur in the same geographic region where distinct habitats such as meadow, forest, swamp, and sand dunes provide ecological niches. Where similar ecological conditions occur in widely separated places it is possible for a similar ecotype to occur. This is different from a subspecies, which may exist across a number of different habitats. In animals, ecotypes can be regarded as micro-subspecies that owe their differing characteristics to the effects of a very local environment.<sup id="cite_ref-mayr1999_174-0" class="reference"><a href="#cite_note-mayr1999-174"><span class="cite-bracket">&#91;</span>173<span class="cite-bracket">&#93;</span></a></sup> Ecotypes have no <a href="/wiki/Taxonomic_rank" title="Taxonomic rank">taxonomic rank</a>. </p><p>Grey wolves have a wide, natural distribution across the <a href="/wiki/Holarctic" class="mw-redirect" title="Holarctic">Holarctic</a> that includes many different habitats, which can vary from the high arctic to dense forests, open steppe and deserts. The genetic differences between different populations of grey wolves is tightly linked to the type of habitat in which they live.<sup id="cite_ref-leonard2014_9-2" class="reference"><a href="#cite_note-leonard2014-9"><span class="cite-bracket">&#91;</span>9<span class="cite-bracket">&#93;</span></a></sup> Differences in genetic markers among the Scandinavian wolf population has arisen in only just over a decade due to their small population size,<sup id="cite_ref-leonard2014_9-3" class="reference"><a href="#cite_note-leonard2014-9"><span class="cite-bracket">&#91;</span>9<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-vila2003_175-0" class="reference"><a href="#cite_note-vila2003-175"><span class="cite-bracket">&#91;</span>174<span class="cite-bracket">&#93;</span></a></sup> which indicates that these differences are not dependent on a long time spent in isolation and that larger population patterns can evolve in just a few thousand years.<sup id="cite_ref-leonard2014_9-4" class="reference"><a href="#cite_note-leonard2014-9"><span class="cite-bracket">&#91;</span>9<span class="cite-bracket">&#93;</span></a></sup> These differences can also include fur color and density, and body size.<sup id="cite_ref-leonard2014_9-5" class="reference"><a href="#cite_note-leonard2014-9"><span class="cite-bracket">&#91;</span>9<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-munoz2009_176-0" class="reference"><a href="#cite_note-munoz2009-176"><span class="cite-bracket">&#91;</span>175<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-musiani2007_4-2" class="reference"><a href="#cite_note-musiani2007-4"><span class="cite-bracket">&#91;</span>4<span class="cite-bracket">&#93;</span></a></sup> The differences can also include behavior, as coastal wolves eat fish<sup id="cite_ref-leonard2014_9-6" class="reference"><a href="#cite_note-leonard2014-9"><span class="cite-bracket">&#91;</span>9<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-munoz2009_176-1" class="reference"><a href="#cite_note-munoz2009-176"><span class="cite-bracket">&#91;</span>175<span class="cite-bracket">&#93;</span></a></sup> and tundra wolves migrate.<sup id="cite_ref-leonard2014_9-7" class="reference"><a href="#cite_note-leonard2014-9"><span class="cite-bracket">&#91;</span>9<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-musiani2007_4-3" class="reference"><a href="#cite_note-musiani2007-4"><span class="cite-bracket">&#91;</span>4<span class="cite-bracket">&#93;</span></a></sup> These differences have been observed between two wolf populations that are living in close proximity. It has been shown that mountain wolves do not interbreed with nearby coastal wolves, and the Alps of France and Switzerland have been repopulated with wolves from the mountains of nearby Italy<sup id="cite_ref-leonard2014_9-8" class="reference"><a href="#cite_note-leonard2014-9"><span class="cite-bracket">&#91;</span>9<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-valiere2003_177-0" class="reference"><a href="#cite_note-valiere2003-177"><span class="cite-bracket">&#91;</span>176<span class="cite-bracket">&#93;</span></a></sup> and from the far away mountains of Croatia<sup id="cite_ref-leonard2014_9-9" class="reference"><a href="#cite_note-leonard2014-9"><span class="cite-bracket">&#91;</span>9<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-fabbri2014_178-0" class="reference"><a href="#cite_note-fabbri2014-178"><span class="cite-bracket">&#91;</span>177<span class="cite-bracket">&#93;</span></a></sup> rather than from the nearer lowlands, which indicates that distance is not the driving force in differences between the two <a href="/wiki/Ecomorphology" title="Ecomorphology">ecomorphs</a>.<sup id="cite_ref-leonard2014_9-10" class="reference"><a href="#cite_note-leonard2014-9"><span class="cite-bracket">&#91;</span>9<span class="cite-bracket">&#93;</span></a></sup> </p><p>In 2013, a genetic study found that the wolf population in Europe was divided along a north–south axis and formed five major clusters. Three clusters were identified occupying southern and central Europe in Italy, the Carpathians, and the Dinaric-Balkans. Another two clusters were identified occupying north-central Europe and the Ukrainian steppe. The Italian wolf consisted of an isolated population with low genetic diversity. Wolves from Croatia, Bulgaria, and Greece formed the Dinaric-Balkans cluster. Wolves from Finland, Latvia, Belarus, Poland and Russia formed the north-central Europe cluster with wolves from the Carpathians cluster a mixture of wolves from the north-central cluster and the Dinaric-Balkans cluster. The wolves from the Carpathians were more similar to the wolves from the Ukrainian Steppe than they were to wolves from north-central Europe. These clusters may have been the result of expansion from glacial refugia, an adaptation to local environments, and landscape fragmentation and the killing of wolves in some areas by humans.<sup id="cite_ref-stronen2013_179-0" class="reference"><a href="#cite_note-stronen2013-179"><span class="cite-bracket">&#91;</span>178<span class="cite-bracket">&#93;</span></a></sup> </p><p>In 2016, two studies compared the sequences of 42,000 <a href="/wiki/Single_nucleotide_polymorphisms" class="mw-redirect" title="Single nucleotide polymorphisms">single nucleotide polymorphisms</a> in North American grey wolves and found that they formed six ecotypes. These six wolf ecotypes were named West Forest, Boreal Forest, Arctic, High Arctic, Baffin, and British Columbia. The studies found that precipitation and mean diurnal temperature range were the most influential variables on sequence variation.<sup id="cite_ref-schweizer2016a_5-2" class="reference"><a href="#cite_note-schweizer2016a-5"><span class="cite-bracket">&#91;</span>5<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-schweizer2016b_180-0" class="reference"><a href="#cite_note-schweizer2016b-180"><span class="cite-bracket">&#91;</span>179<span class="cite-bracket">&#93;</span></a></sup> These findings were in accord with previous findings that precipitation influenced morphology,<sup id="cite_ref-okeefe2013_181-0" class="reference"><a href="#cite_note-okeefe2013-181"><span class="cite-bracket">&#91;</span>180<span class="cite-bracket">&#93;</span></a></sup> and that vegetation<sup id="cite_ref-geffen2004_6-2" class="reference"><a href="#cite_note-geffen2004-6"><span class="cite-bracket">&#91;</span>6<span class="cite-bracket">&#93;</span></a></sup> and habitat type<sup id="cite_ref-munoz2009_176-2" class="reference"><a href="#cite_note-munoz2009-176"><span class="cite-bracket">&#91;</span>175<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-carmichael2007_182-0" class="reference"><a href="#cite_note-carmichael2007-182"><span class="cite-bracket">&#91;</span>181<span class="cite-bracket">&#93;</span></a></sup> influenced wolf differences. One of these studies found that the variation in 11 key genes affected wolf vision, sense of smell, hearing, coat color, metabolism, and immunity. The study identified 1,040 genes that are potentially under selection due to habitat variation, and therefore that there was evidence of local adaption of the wolf ecotypes at a molecular level. Most notable was the positive selection of genes that influence vision, coat color, metabolism and immunity in the Arctic and High Arctic ecotypes, and that the British Columbia ecotype also has a unique set of adaptions.<sup id="cite_ref-schweizer2016b_180-1" class="reference"><a href="#cite_note-schweizer2016b-180"><span class="cite-bracket">&#91;</span>179<span class="cite-bracket">&#93;</span></a></sup> The local adaptation of a wolf ecotype most likely reflects a wolf's preference to remain in the type of habitat that it was born into.<sup id="cite_ref-schweizer2016a_5-3" class="reference"><a href="#cite_note-schweizer2016a-5"><span class="cite-bracket">&#91;</span>5<span class="cite-bracket">&#93;</span></a></sup> </p><p>Ecological factors including habitat type, climate, prey specialization and predatory competition will greatly influence grey wolf <a href="/wiki/Population_genetics" title="Population genetics">genetic population structure</a> and cranio-dental <a href="/wiki/Phenotypic_plasticity" title="Phenotypic plasticity">plasticity</a>.<sup id="cite_ref-hofreiter2010_183-0" class="reference"><a href="#cite_note-hofreiter2010-183"><span class="cite-bracket">&#91;</span>182<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-perri2016_87-1" class="reference"><a href="#cite_note-perri2016-87"><span class="cite-bracket">&#91;</span>87<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-leonard2014_9-11" class="reference"><a href="#cite_note-leonard2014-9"><span class="cite-bracket">&#91;</span>9<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-musiani2007_4-4" class="reference"><a href="#cite_note-musiani2007-4"><span class="cite-bracket">&#91;</span>4<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-carmichael2001_184-0" class="reference"><a href="#cite_note-carmichael2001-184"><span class="cite-bracket">&#91;</span>183<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-carmichael2006_185-0" class="reference"><a href="#cite_note-carmichael2006-185"><span class="cite-bracket">&#91;</span>184<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-geffen2004_6-3" class="reference"><a href="#cite_note-geffen2004-6"><span class="cite-bracket">&#91;</span>6<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-pilot2006_7-2" class="reference"><a href="#cite_note-pilot2006-7"><span class="cite-bracket">&#91;</span>7<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-flower2014_8-2" class="reference"><a href="#cite_note-flower2014-8"><span class="cite-bracket">&#91;</span>8<span class="cite-bracket">&#93;</span></a></sup> During the Last Glacial Maximum, there was greater wolf <a href="/wiki/Genetic_diversity" title="Genetic diversity">genetic diversity</a> than there is today,<sup id="cite_ref-freedman2014_12-14" class="reference"><a href="#cite_note-freedman2014-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-thalmann2013_73-9" class="reference"><a href="#cite_note-thalmann2013-73"><span class="cite-bracket">&#91;</span>73<span class="cite-bracket">&#93;</span></a></sup> and within the Pleistocene grey wolf population the variations between local environments would have encouraged a range of wolf ecotypes that were genetically, morphologically and ecologically distinct from one another.<sup id="cite_ref-perri2016_87-2" class="reference"><a href="#cite_note-perri2016-87"><span class="cite-bracket">&#91;</span>87<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Pleistocene_wolves">Pleistocene wolves</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=29" title="Edit section: Pleistocene wolves"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Pleistocene_wolf" title="Pleistocene wolf">Pleistocene wolf</a></div> <p>During the <a href="/wiki/Last_Glacial_Maximum" title="Last Glacial Maximum">Last Glacial Maximum</a> 20,000 YBP, the <a href="/wiki/Mammoth_steppe" title="Mammoth steppe">Pleistocene steppe</a> stretched across northern and central Eurasia and through <a href="/wiki/Beringia" title="Beringia">Beringia</a> into North America. The Pleistocene <a href="/wiki/Beringian_wolf" title="Beringian wolf">wolves of Beringia</a>, and perhaps those across the steppe, were adapted to this habitat. Their tooth and skull morphology indicates that they specialized in preying on now-extinct <a href="/wiki/Pleistocene_megafauna" class="mw-redirect" title="Pleistocene megafauna">Pleistocene megafauna</a>, and their tooth wear indicates that their behavior was different from modern wolves.<sup id="cite_ref-leonard2007_125-3" class="reference"><a href="#cite_note-leonard2007-125"><span class="cite-bracket">&#91;</span>125<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-leonard2014_9-12" class="reference"><a href="#cite_note-leonard2014-9"><span class="cite-bracket">&#91;</span>9<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-fox2008_186-0" class="reference"><a href="#cite_note-fox2008-186"><span class="cite-bracket">&#91;</span>185<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-baryshnikov2009_187-0" class="reference"><a href="#cite_note-baryshnikov2009-187"><span class="cite-bracket">&#91;</span>186<span class="cite-bracket">&#93;</span></a></sup> This highlights the success of <i>C. lupus</i> as a species in adapting to different environmental conditions.<sup id="cite_ref-flower2014_8-3" class="reference"><a href="#cite_note-flower2014-8"><span class="cite-bracket">&#91;</span>8<span class="cite-bracket">&#93;</span></a></sup> This grey wolf ecomorph became extinct at the end of the glaciation, along with the horse and other species on which it depended, and was replaced by wolves from southern North America. This indicates that specialized wolf ecomorphs can become extinct when their environment changes even though the habitat may still support other wolves.<sup id="cite_ref-leonard2014_9-13" class="reference"><a href="#cite_note-leonard2014-9"><span class="cite-bracket">&#91;</span>9<span class="cite-bracket">&#93;</span></a></sup> Wolves went through a <a href="/wiki/Population_bottleneck" title="Population bottleneck">population bottleneck</a> 20,000 YBP that coincides with the Last Glacial Maximum,<sup id="cite_ref-freedman2014_12-15" class="reference"><a href="#cite_note-freedman2014-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-pilot2014_188-0" class="reference"><a href="#cite_note-pilot2014-188"><span class="cite-bracket">&#91;</span>187<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-perri2016_87-3" class="reference"><a href="#cite_note-perri2016-87"><span class="cite-bracket">&#91;</span>87<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-leonard2014_9-14" class="reference"><a href="#cite_note-leonard2014-9"><span class="cite-bracket">&#91;</span>9<span class="cite-bracket">&#93;</span></a></sup> which indicates that many wolf populations may have gone extinct at the same time as the Beringian wolves.<sup id="cite_ref-leonard2014_9-15" class="reference"><a href="#cite_note-leonard2014-9"><span class="cite-bracket">&#91;</span>9<span class="cite-bracket">&#93;</span></a></sup> </p><p>There are a small number of <i>Canis</i> remains that have been found at Goyet Cave, Belgium (36,500 YBP)<sup id="cite_ref-germonpre2009_71-1" class="reference"><a href="#cite_note-germonpre2009-71"><span class="cite-bracket">&#91;</span>71<span class="cite-bracket">&#93;</span></a></sup> Razboinichya Cave, Russia (33,500 YBP)<sup id="cite_ref-ovodov2011_189-0" class="reference"><a href="#cite_note-ovodov2011-189"><span class="cite-bracket">&#91;</span>188<span class="cite-bracket">&#93;</span></a></sup> Kostenki 8, Russia (33,500–26,500 YBP)<sup id="cite_ref-germonpre2015_190-0" class="reference"><a href="#cite_note-germonpre2015-190"><span class="cite-bracket">&#91;</span>189<span class="cite-bracket">&#93;</span></a></sup> Predmosti, Czech Republic (31,000 YBP)<sup id="cite_ref-germonpre2012_191-0" class="reference"><a href="#cite_note-germonpre2012-191"><span class="cite-bracket">&#91;</span>190<span class="cite-bracket">&#93;</span></a></sup> and Eliseevichi 1, Russia (17,000 YBP).<sup id="cite_ref-sablin2002_72-1" class="reference"><a href="#cite_note-sablin2002-72"><span class="cite-bracket">&#91;</span>72<span class="cite-bracket">&#93;</span></a></sup> Based on cranial morphometric study of the characteristics thought to be associated with the domestication process, these have been proposed as early Paleolithic dogs.<sup id="cite_ref-germonpre2015_190-1" class="reference"><a href="#cite_note-germonpre2015-190"><span class="cite-bracket">&#91;</span>189<span class="cite-bracket">&#93;</span></a></sup> These characteristics of shortened rostrum, tooth crowding, and absence or rotation of premolars have been documented in both ancient and modern wolves.<sup id="cite_ref-leonard2007_125-4" class="reference"><a href="#cite_note-leonard2007-125"><span class="cite-bracket">&#91;</span>125<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-perri2016_87-4" class="reference"><a href="#cite_note-perri2016-87"><span class="cite-bracket">&#91;</span>87<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-flower2014_8-4" class="reference"><a href="#cite_note-flower2014-8"><span class="cite-bracket">&#91;</span>8<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-boudadi2010_192-0" class="reference"><a href="#cite_note-boudadi2010-192"><span class="cite-bracket">&#91;</span>191<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-dimitrijevic2015_193-0" class="reference"><a href="#cite_note-dimitrijevic2015-193"><span class="cite-bracket">&#91;</span>192<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-crockford2012_194-0" class="reference"><a href="#cite_note-crockford2012-194"><span class="cite-bracket">&#91;</span>193<span class="cite-bracket">&#93;</span></a></sup> Rather than representing early dogs, these specimens may represent "a morphologically distinct local, now extinct, population of wolves".<sup id="cite_ref-perri2016_87-5" class="reference"><a href="#cite_note-perri2016-87"><span class="cite-bracket">&#91;</span>87<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-larson2012_195-0" class="reference"><a href="#cite_note-larson2012-195"><span class="cite-bracket">&#91;</span>194<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Notes">Notes</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=30" title="Edit section: Notes"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-lower-alpha"> <div class="mw-references-wrap"><ol class="references"> <li id="cite_note-mtdna_seq_desc-132"><span class="mw-cite-backlink"><b><a href="#cite_ref-mtdna_seq_desc_132-0">^</a></b></span> <span class="reference-text"><a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">Phylogenetic relationships</a> and time of <a href="/wiki/Genetic_divergence" title="Genetic divergence">genetic divergence</a> for the extant <a href="#Wolf-like_canids">wolf-like canids</a> based on DNA of the entire <a href="/wiki/Cell_nucleus" title="Cell nucleus">cell nucleus</a>.<sup id="cite_ref-gopalakrishnan2018_119-0" class="reference"><a href="#cite_note-gopalakrishnan2018-119"><span class="cite-bracket">&#91;</span>119<span class="cite-bracket">&#93;</span></a></sup> Exception 1: Side-striped jackal and Black-backed jackal added based on segments of DNA taken from the <a href="/wiki/Cell_nucleus" title="Cell nucleus">cell nucleus</a><sup id="cite_ref-lindblad2005_97-5" class="reference"><a href="#cite_note-lindblad2005-97"><span class="cite-bracket">&#91;</span>97<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-koepfli2015_98-2" class="reference"><a href="#cite_note-koepfli2015-98"><span class="cite-bracket">&#91;</span>98<span class="cite-bracket">&#93;</span></a></sup> with proposed divergence times in millions of years.<sup id="cite_ref-koepfli2015_98-3" class="reference"><a href="#cite_note-koepfli2015-98"><span class="cite-bracket">&#91;</span>98<span class="cite-bracket">&#93;</span></a></sup> Exception 2: The position of the Himalayan wolf is based on segments of <a href="/wiki/Mitochondrial_DNA" title="Mitochondrial DNA">mitochondrial DNA</a><sup id="cite_ref-koepfli2015_98-4" class="reference"><a href="#cite_note-koepfli2015-98"><span class="cite-bracket">&#91;</span>98<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-ersmark2016_120-0" class="reference"><a href="#cite_note-ersmark2016-120"><span class="cite-bracket">&#91;</span>120<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-werhahn2017_121-0" class="reference"><a href="#cite_note-werhahn2017-121"><span class="cite-bracket">&#91;</span>121<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-werhahn2018_122-0" class="reference"><a href="#cite_note-werhahn2018-122"><span class="cite-bracket">&#91;</span>122<span class="cite-bracket">&#93;</span></a></sup> and segments from the cell nucleus.<sup id="cite_ref-koepfli2015_98-5" class="reference"><a href="#cite_note-koepfli2015-98"><span class="cite-bracket">&#91;</span>98<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-werhahn2017_121-1" class="reference"><a href="#cite_note-werhahn2017-121"><span class="cite-bracket">&#91;</span>121<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-werhahn2018_122-1" class="reference"><a href="#cite_note-werhahn2018-122"><span class="cite-bracket">&#91;</span>122<span class="cite-bracket">&#93;</span></a></sup> Exception 3: The position of the Indian plains wolf is based on mDNA.<sup id="cite_ref-aggarwal2007_123-0" class="reference"><a href="#cite_note-aggarwal2007-123"><span class="cite-bracket">&#91;</span>123<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-sharma2004_124-0" class="reference"><a href="#cite_note-sharma2004-124"><span class="cite-bracket">&#91;</span>124<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-leonard2007_125-0" class="reference"><a href="#cite_note-leonard2007-125"><span class="cite-bracket">&#91;</span>125<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-pilot2010_126-0" class="reference"><a href="#cite_note-pilot2010-126"><span class="cite-bracket">&#91;</span>126<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-rueness2011_127-0" class="reference"><a href="#cite_note-rueness2011-127"><span class="cite-bracket">&#91;</span>127<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-miklosi2015_128-0" class="reference"><a href="#cite_note-miklosi2015-128"><span class="cite-bracket">&#91;</span>128<span class="cite-bracket">&#93;</span></a></sup> and its timing 120,000 years ago.<sup id="cite_ref-Loog2018_15-1" class="reference"><a href="#cite_note-Loog2018-15"><span class="cite-bracket">&#91;</span>15<span class="cite-bracket">&#93;</span></a></sup> Exception 4: <a href="/wiki/Late_Pleistocene" title="Late Pleistocene">Late Pleistocene</a> wolf included for time-line purposes only - the <a href="/wiki/Most_recent_common_ancestor" title="Most recent common ancestor">most recent common ancestor</a> for all other <i>Canis lupus</i> specimens – modern and extinct but excluding the Indian plains wolf and the Himalayan wolf – was 80,000 years before present.<sup id="cite_ref-thalmann2013_73-2" class="reference"><a href="#cite_note-thalmann2013-73"><span class="cite-bracket">&#91;</span>73<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-koblmuller2016_129-0" class="reference"><a href="#cite_note-koblmuller2016-129"><span class="cite-bracket">&#91;</span>129<span class="cite-bracket">&#93;</span></a></sup> Exception 5: Grey wolf and dog timing 25,000 years ago.<sup id="cite_ref-Loog2018_15-2" class="reference"><a href="#cite_note-Loog2018-15"><span class="cite-bracket">&#91;</span>15<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Werhahn2018_130-0" class="reference"><a href="#cite_note-Werhahn2018-130"><span class="cite-bracket">&#91;</span>130<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-Schweizer2020_131-0" class="reference"><a href="#cite_note-Schweizer2020-131"><span class="cite-bracket">&#91;</span>131<span class="cite-bracket">&#93;</span></a></sup></span> </li> </ol></div></div> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=31" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239543626"><div class="reflist reflist-columns references-column-width" style="column-width: 30em;"> <ol class="references"> <li id="cite_note-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</a></b></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output 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Archived from <a rel="nofollow" class="external text" href="https://pdfs.semanticscholar.org/d7c7/212ab77c711f6c0db4cde4e8daa855bc8ab2.pdf?_ga=2.31998270.1689261500.1568962364-317290015.1568962364">the original</a> <span class="cs1-format">(PDF)</span> on 2019-09-20<span class="reference-accessdate">. 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Doctoral Dissertations Available from Proquest. Paper AAI3136721. <a rel="nofollow" class="external text" href="http://scholarworks.umass.edu/dissertations/AAI3136721">link</a></span> </li> <li id="cite_note-drake2011-80"><span class="mw-cite-backlink"><b><a href="#cite_ref-drake2011_80-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFDrake2011" class="citation journal cs1">Drake, Abby Grace (2011). 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"Fossil dogs and wolves from Palaeolithic sites in Belgium, the Ukraine and Russia: osteometry, ancient DNA and stable isotopes". <i>Journal of Archaeological Science</i>. <b>36</b> (2): 473–490. <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/2009JArSc..36..473G">2009JArSc..36..473G</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.jas.2008.09.033">10.1016/j.jas.2008.09.033</a>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.jtitle=Journal+of+Archaeological+Science&amp;rft.atitle=Fossil+dogs+and+wolves+from+Palaeolithic+sites+in+Belgium%2C+the+Ukraine+and+Russia%3A+osteometry%2C+ancient+DNA+and+stable+isotopes&amp;rft.volume=36&amp;rft.issue=2&amp;rft.pages=473-490&amp;rft.date=2008&amp;rft_id=info%3Adoi%2F10.1016%2Fj.jas.2008.09.033&amp;rft_id=info%3Abibcode%2F2009JArSc..36..473G&amp;rft.aulast=Germonpr%C3%A9a&amp;rft.aufirst=Mietje&amp;rft.au=Sablinb%2C+Mikhail+V.&amp;rft.au=Stevensc%2C+Rhiannon+E.&amp;rft.au=Hedgesd%2C+Robert+E.+M.&amp;rft.au=Hofreitere%2C+Michael&amp;rft.au=Stillere%2C+Mathias&amp;rft.au=Despr%C3%A9se%2C+Viviane+R.&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+the+wolf" class="Z3988"></span></span> </li> <li id="cite_note-larson2012-195"><span class="mw-cite-backlink"><b><a href="#cite_ref-larson2012_195-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLarson_G2012" class="citation journal cs1">Larson G (2012). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3384140">"Rethinking dog domestication by integrating genetics, archeology, and biogeography"</a>. <i>PNAS</i>. <b>109</b> (23): 8878–8883. <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/2012PNAS..109.8878L">2012PNAS..109.8878L</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.1203005109">10.1073/pnas.1203005109</a></span>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a>&#160;<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3384140">3384140</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a>&#160;<a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/22615366">22615366</a>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.jtitle=PNAS&amp;rft.atitle=Rethinking+dog+domestication+by+integrating+genetics%2C+archeology%2C+and+biogeography&amp;rft.volume=109&amp;rft.issue=23&amp;rft.pages=8878-8883&amp;rft.date=2012&amp;rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3384140%23id-name%3DPMC&amp;rft_id=info%3Apmid%2F22615366&amp;rft_id=info%3Adoi%2F10.1073%2Fpnas.1203005109&amp;rft_id=info%3Abibcode%2F2012PNAS..109.8878L&amp;rft.au=Larson+G&amp;rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3384140&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+the+wolf" class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="Works">Works</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_the_wolf&amp;action=edit&amp;section=32" title="Edit section: Works"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWangTedford2008" class="citation book cs1"><a href="/wiki/Xiaoming_Wang_(paleontologist)" title="Xiaoming Wang (paleontologist)">Wang, Xiaoming</a>; <a href="/wiki/Richard_H._Tedford" title="Richard H. Tedford">Tedford, Richard H.</a> (2008). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=LnWdpK7ctI0C"><i>Dogs: Their Fossil Relatives and Evolutionary History</i></a>. <a href="/wiki/Columbia_University_Press" title="Columbia University Press">Columbia University Press</a>, New York. pp.&#160;1–232. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a>&#160;<a href="/wiki/Special:BookSources/978-0-231-13529-0" title="Special:BookSources/978-0-231-13529-0"><bdi>978-0-231-13529-0</bdi></a>. <a href="/wiki/OCLC_(identifier)" class="mw-redirect" title="OCLC (identifier)">OCLC</a>&#160;<a rel="nofollow" class="external text" href="https://search.worldcat.org/oclc/502410693">502410693</a>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&amp;rft.genre=book&amp;rft.btitle=Dogs%3A+Their+Fossil+Relatives+and+Evolutionary+History&amp;rft.pages=1-232&amp;rft.pub=Columbia+University+Press%2C+New+York&amp;rft.date=2008&amp;rft_id=info%3Aoclcnum%2F502410693&amp;rft.isbn=978-0-231-13529-0&amp;rft.aulast=Wang&amp;rft.aufirst=Xiaoming&amp;rft.au=Tedford%2C+Richard+H.&amp;rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DLnWdpK7ctI0C&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+the+wolf" class="Z3988"></span></li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline ol,.mw-parser-output .hlist.inline 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title="Evolutionary taxonomy">Evolutionary taxonomy</a></li> <li><a href="/wiki/Systematics" title="Systematics">Systematics</a></li></ul> </div></td><td class="noviewer navbox-image" rowspan="7" style="width:1px;padding:0 0 0 2px"><div><ul role="navigation" aria-label="Portals" class="noprint portalbox portalborder portalright"> <li class="portalbox-entry"><span class="portalbox-image"><span class="noviewer" typeof="mw:File"><a href="/wiki/File:Tree_of_life.svg" class="mw-file-description"><img alt="icon" src="//upload.wikimedia.org/wikipedia/commons/thumb/0/09/Tree_of_life.svg/32px-Tree_of_life.svg.png" decoding="async" width="32" height="27" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/09/Tree_of_life.svg/48px-Tree_of_life.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/09/Tree_of_life.svg/64px-Tree_of_life.svg.png 2x" data-file-width="915" data-file-height="760" /></a></span></span><span class="portalbox-link"><a href="/wiki/Portal:Evolutionary_biology" title="Portal:Evolutionary biology">Evolutionary biology portal</a></span></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-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Phylogenesis" title="Phylogenesis">Phylogenesis</a> <ul><li><a href="/wiki/Cladogenesis" title="Cladogenesis">Cladogenesis</a></li></ul></li> <li><a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">Phylogenetic tree</a> <ul><li><a href="/wiki/Cladogram" title="Cladogram">Cladogram</a></li></ul></li> <li><a href="/wiki/Phylogenetic_network" title="Phylogenetic network">Phylogenetic network</a></li> <li><a href="/wiki/Long_branch_attraction" title="Long branch attraction">Long branch attraction</a></li> <li><a href="/wiki/Clade" title="Clade">Clade</a> vs <a href="/wiki/Evolutionary_grade" title="Evolutionary grade">Grade</a></li> <li><a href="/wiki/Lineage_(evolution)" title="Lineage (evolution)">Lineage</a> <ul><li><a href="/wiki/Ghost_lineage" title="Ghost lineage">Ghost lineage</a></li></ul></li> <li><a href="/wiki/Ghost_population" title="Ghost population">Ghost population</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Inference methods</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/Maximum_parsimony_(phylogenetics)" title="Maximum parsimony (phylogenetics)">Maximum parsimony</a></li> <li><a href="/wiki/Phylogenetic_reconciliation" title="Phylogenetic reconciliation">Phylogenetic reconciliation</a></li> <li>Probabilistic methods <ul><li><a href="/wiki/Maximum_likelihood" class="mw-redirect" title="Maximum likelihood">Maximum likelihood</a></li> <li><a href="/wiki/Bayesian_inference_in_phylogeny" title="Bayesian inference in phylogeny">Bayesian inference</a></li></ul></li> <li><a href="/wiki/Distance_matrices_in_phylogeny" title="Distance matrices in phylogeny">Distance-matrix methods</a> <ul><li><a href="/wiki/Neighbor-joining" class="mw-redirect" title="Neighbor-joining">Neighbor-joining</a></li> <li><a href="/wiki/UPGMA" title="UPGMA">UPGMA</a></li> <li><a href="/wiki/Least_squares_inference_in_phylogeny" title="Least squares inference in phylogeny">Least squares</a></li></ul></li> <li><a href="/wiki/Three-taxon_analysis" title="Three-taxon analysis">Three-taxon analysis</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Current topics</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><i><a href="/wiki/PhyloCode" title="PhyloCode">PhyloCode</a></i></li> <li><a href="/wiki/DNA_barcoding" title="DNA barcoding">DNA barcoding</a></li> <li><a href="/wiki/Molecular_phylogenetics" title="Molecular phylogenetics">Molecular phylogenetics</a></li> <li><a href="/wiki/Phylogenetic_comparative_methods" title="Phylogenetic comparative methods">Phylogenetic comparative methods</a></li> <li><a href="/wiki/Phylogenetic_niche_conservatism" title="Phylogenetic niche conservatism">Phylogenetic niche conservatism</a></li> <li><a href="/wiki/Phylogenetic_signal" title="Phylogenetic signal">Phylogenetic signal</a></li> <li><a href="/wiki/List_of_phylogenetics_software" title="List of phylogenetics software">Phylogenetics software</a></li> <li><a href="/wiki/Phylogenomics" title="Phylogenomics">Phylogenomics</a></li> <li><a href="/wiki/Phylogeography" title="Phylogeography">Phylogeography</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Group traits</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/Primitive_(phylogenetics)" title="Primitive (phylogenetics)">Primitive</a> <ul><li><a href="/wiki/Plesiomorphy" class="mw-redirect" title="Plesiomorphy">Plesiomorphy</a></li> <li><a href="/wiki/Symplesiomorphy" class="mw-redirect" title="Symplesiomorphy">Symplesiomorphy</a></li></ul></li> <li>Derived <ul><li><a href="/wiki/Synapomorphy" class="mw-redirect" title="Synapomorphy">Apomorphy</a></li> <li><a href="/wiki/Synapomorphy" class="mw-redirect" title="Synapomorphy">Synapomorphy</a></li> <li><a href="/wiki/Autapomorphy" title="Autapomorphy">Autapomorphy</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Group types</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/Monophyly" title="Monophyly">Monophyly</a></li> <li><a href="/wiki/Paraphyly" title="Paraphyly">Paraphyly</a></li> <li><a href="/wiki/Polyphyly" title="Polyphyly">Polyphyly</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Nomenclature</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/Phylogenetic_nomenclature" title="Phylogenetic nomenclature">Phylogenetic nomenclature</a></li> <li><a href="/wiki/Crown_group" title="Crown group">Crown group</a></li> <li><a href="/wiki/Sister_group" title="Sister group">Sister group</a></li> <li><a href="/wiki/Basal_(phylogenetics)" title="Basal (phylogenetics)">Basal</a></li> <li><a href="/wiki/Supertree" title="Supertree">Supertree</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="3"><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:Phylogenetics" title="Category:Phylogenetics">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:Phylogenetics" class="extiw" title="commons:Category:Phylogenetics">Commons</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="Gene_expression" 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:Gene_expression" title="Template:Gene expression"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Gene_expression" title="Template talk:Gene expression"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Gene_expression" title="Special:EditPage/Template:Gene expression"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Gene_expression" style="font-size:114%;margin:0 4em"><a href="/wiki/Gene_expression" title="Gene expression">Gene expression</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Introduction_to_genetics" title="Introduction to genetics">Introduction<br />to genetics</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/Genetic_code" title="Genetic code">Genetic code</a></li> <li><a href="/wiki/Central_dogma_of_molecular_biology" title="Central dogma of molecular biology">Central dogma</a> <ul><li><a href="/wiki/DNA" title="DNA">DNA</a>&#160;→ <a href="/wiki/Messenger_RNA" title="Messenger RNA">RNA</a>&#160;→ <a href="/wiki/Protein" title="Protein">Protein</a></li></ul></li> <li><a href="/wiki/Central_dogma_of_molecular_biology#Special_transfers_of_biological_sequential_information" title="Central dogma of molecular biology">Special transfers</a> <ul><li><a href="/wiki/RNA-dependent_RNA_polymerase" title="RNA-dependent RNA polymerase">RNA→RNA</a></li> <li><a href="/wiki/Reverse_transcription" class="mw-redirect" title="Reverse transcription">RNA→DNA</a></li> <li><a href="/wiki/Prion" title="Prion">Protein→Protein</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Transcription_(biology)" title="Transcription (biology)">Transcription</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight: normal;">Types</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bacterial_transcription" title="Bacterial transcription">Bacterial</a></li> <li><a href="/wiki/Archaeal_transcription" title="Archaeal transcription">Archaeal</a></li> <li><a href="/wiki/Eukaryotic_transcription" title="Eukaryotic transcription">Eukaryotic</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight: normal;">Key elements</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Transcription_factor" title="Transcription factor">Transcription factor</a></li> <li><a href="/wiki/RNA_polymerase" title="RNA polymerase">RNA polymerase</a></li> <li><a href="/wiki/Promoter_(genetics)" title="Promoter (genetics)">Promoter</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight: normal;"><a href="/wiki/Post-transcriptional_modification" title="Post-transcriptional modification">Post-transcription</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Primary_transcript" title="Primary transcript">Precursor mRNA (pre-mRNA&#160;/&#32;hnRNA)</a></li> <li><a href="/wiki/Five-prime_cap" title="Five-prime cap">5' capping</a></li> <li><a href="/wiki/RNA_splicing" title="RNA splicing">Splicing</a></li> <li><a href="/wiki/Polyadenylation" title="Polyadenylation">Polyadenylation</a></li> <li><a href="/wiki/Histone_acetylation_and_deacetylation" title="Histone acetylation and deacetylation">Histone acetylation and deacetylation</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Translation_(biology)" title="Translation (biology)">Translation</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight: normal;">Types</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bacterial_translation" title="Bacterial translation">Bacterial</a></li> <li><a href="/wiki/Archaeal_translation" title="Archaeal translation">Archaeal</a></li> <li><a href="/wiki/Eukaryotic_translation" title="Eukaryotic translation">Eukaryotic</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:8.0em;font-weight: normal;">Key elements</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0;white-space:nowrap;"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Ribosome" title="Ribosome">Ribosome</a></li> <li><a href="/wiki/Transfer_RNA" title="Transfer RNA">Transfer RNA (tRNA)</a></li> <li><a href="/wiki/Ribosome-nascent_chain_complex" title="Ribosome-nascent chain complex">Ribosome-nascent chain complex (RNC)</a></li> <li><a href="/wiki/Post-translational_modification" title="Post-translational modification">Post-translational modification</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Regulation_of_gene_expression" title="Regulation of gene expression">Regulation</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/Epigenetics" title="Epigenetics">Epigenetic</a> <ul><li><a href="/wiki/Genomic_imprinting" title="Genomic imprinting">imprinting</a></li></ul></li> <li><a href="/wiki/Transcriptional_regulation" title="Transcriptional regulation">Transcriptional</a> <ul><li><a href="/wiki/Gene_regulatory_network" title="Gene regulatory network">Gene regulatory network</a></li> <li><a href="/wiki/Cis-regulatory_element" title="Cis-regulatory element">cis-regulatory element</a></li></ul></li> <li><a href="/wiki/Lac_operon" title="Lac operon">lac operon</a></li> <li><a href="/wiki/Post-transcriptional_regulation" title="Post-transcriptional regulation">Post-transcriptional</a> <ul><li><a href="/wiki/P-bodies" title="P-bodies">sequestration (P-bodies)</a></li> <li><a href="/wiki/Alternative_splicing" title="Alternative splicing">alternative splicing</a></li> <li><a href="/wiki/MicroRNA" title="MicroRNA">microRNA</a></li></ul></li> <li><a href="/wiki/Translational_regulation" title="Translational regulation">Translational</a></li> <li><a href="/wiki/Post-translational_regulation" title="Post-translational regulation">Post-translational</a> <ul><li><a href="/wiki/Phosphorylation" title="Phosphorylation">reversible</a></li> <li><a href="/wiki/Proteolysis" title="Proteolysis">irreversible</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Influential people</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/Fran%C3%A7ois_Jacob" title="François Jacob">François Jacob</a></li> <li><a href="/wiki/Jacques_Monod" title="Jacques Monod">Jacques Monod</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Genetics" 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:Genetics" title="Template:Genetics"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Genetics" title="Template talk:Genetics"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Genetics" title="Special:EditPage/Template:Genetics"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Genetics" style="font-size:114%;margin:0 4em"><a href="/wiki/Genetics" title="Genetics">Genetics</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><a href="/wiki/Introduction_to_genetics" title="Introduction to genetics">Introduction</a></li> <li><a href="/wiki/Outline_of_genetics" title="Outline of genetics">Outline</a></li> <li><a href="/wiki/History_of_genetics" title="History of genetics">History</a></li> <li><a href="/wiki/Timeline_of_the_history_of_genetics" title="Timeline of the history of genetics">Timeline</a></li> <li><a href="/wiki/Index_of_genetics_articles" title="Index of genetics articles">Index</a></li> <li><a href="/wiki/Glossary_of_genetics" class="mw-redirect" title="Glossary of genetics">Glossary</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Key components</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/Chromosome" title="Chromosome">Chromosome</a></li> <li><a href="/wiki/DNA" title="DNA">DNA</a></li> <li><a href="/wiki/RNA" title="RNA">RNA</a></li> <li><a href="/wiki/Genome" title="Genome">Genome</a></li> <li><a href="/wiki/Heredity" title="Heredity">Heredity</a></li> <li><a href="/wiki/Nucleotide" title="Nucleotide">Nucleotide</a></li> <li><a href="/wiki/Mutation" title="Mutation">Mutation</a></li> <li><a href="/wiki/Genetic_variation" title="Genetic variation">Genetic variation</a></li> <li><a href="/wiki/Allele" title="Allele">Allele</a></li> <li><a href="/wiki/Amino_acid" title="Amino acid">Amino acid</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Fields</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/Classical_genetics" title="Classical genetics">Classical</a></li> <li><a href="/wiki/Conservation_genetics" title="Conservation genetics">Conservation</a></li> <li><a href="/wiki/Cytogenetics" title="Cytogenetics">Cytogenetics</a></li> <li><a href="/wiki/Ecological_genetics" title="Ecological genetics">Ecological</a></li> <li><a href="/wiki/Immunogenetics" title="Immunogenetics">Immunogenetics</a></li> <li><a href="/wiki/Microbial_genetics" title="Microbial genetics">Microbial</a></li> <li><a href="/wiki/Molecular_genetics" title="Molecular genetics">Molecular</a></li> <li><a href="/wiki/Population_genetics" title="Population genetics">Population</a></li> <li><a href="/wiki/Quantitative_genetics" title="Quantitative genetics">Quantitative</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Archaeogenetics" title="Archaeogenetics">Archaeogenetics</a> of</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/Genetic_history_of_Africa" title="Genetic history of Africa">Africa</a></li> <li><a href="/wiki/Genetic_history_of_indigenous_peoples_of_the_Americas" class="mw-redirect" title="Genetic history of indigenous peoples of the Americas">the Americas</a></li> <li><a href="/wiki/Genetic_history_of_the_British_Isles" title="Genetic history of the British Isles">the British Isles</a></li> <li><a href="/wiki/Genetic_history_of_Europe" title="Genetic history of Europe">Europe</a></li> <li><a href="/wiki/Genetic_history_of_Italy" title="Genetic history of Italy">Italy</a></li> <li><a href="/wiki/Genetic_history_of_the_Middle_East" title="Genetic history of the Middle East">the Middle East</a></li> <li><a href="/wiki/Genetics_and_archaeogenetics_of_South_Asia" title="Genetics and archaeogenetics of South Asia">South Asia</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related topics</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/Behavioural_genetics" title="Behavioural genetics">Behavioural genetics</a></li> <li><a href="/wiki/Epigenetics" title="Epigenetics">Epigenetics</a></li> <li><a href="/wiki/Geneticist" title="Geneticist">Geneticist</a></li> <li><a href="/wiki/Genome_editing" title="Genome editing">Genome editing</a></li> <li><a href="/wiki/Genomics" title="Genomics">Genomics</a></li> <li><a href="/wiki/Genetic_code" title="Genetic code">Genetic code</a></li> <li><a href="/wiki/Genetic_engineering" title="Genetic engineering">Genetic engineering</a></li> <li><a href="/wiki/Genetic_diversity" title="Genetic diversity">Genetic diversity</a></li> <li><a href="/wiki/Genetic_monitoring" title="Genetic monitoring">Genetic monitoring</a></li> <li><a href="/wiki/Genetic_genealogy" title="Genetic genealogy">Genetic genealogy</a></li> <li><a href="/wiki/Heredity" title="Heredity">Heredity</a></li> <li><a href="/wiki/He_Jiankui_affair" title="He Jiankui affair">He Jiankui affair</a></li> <li><a href="/wiki/Medical_genetics" title="Medical genetics">Medical genetics</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/Plant_genetics" title="Plant genetics">Plant genetics</a></li> <li><a href="/wiki/Population_genomics" title="Population genomics">Population genomics</a></li> <li><a href="/wiki/Reverse_genetics" title="Reverse genetics">Reverse genetics</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Lists</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/List_of_genetic_codes" title="List of genetic codes">List of genetic codes</a></li> <li><a href="/wiki/List_of_genetics_research_organizations" title="List of genetics research organizations">List of genetics research organizations</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> <a href="/wiki/Category:Genetics" title="Category:Genetics">Category</a></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Evolutionary_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&#39;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 class="mw-selflink selflink">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" typeof="mw:File"><span title="Category"><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/16px-Symbol_category_class.svg.png" decoding="async" width="16" height="16" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/23px-Symbol_category_class.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/96/Symbol_category_class.svg/31px-Symbol_category_class.svg.png 2x" data-file-width="180" data-file-height="185" /></span></span> <a href="/wiki/Category:Evolutionary_biology" title="Category:Evolutionary biology">Category</a></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> <a href="/wiki/Portal:Evolutionary_biology" title="Portal:Evolutionary biology">Portal</a></li></ul> </div></td></tr></tbody></table></div> <!-- NewPP limit report Parsed by mw‐web.codfw.main‐f69cdc8f6‐ckrcl Cached time: 20241124053517 Cache expiry: 2592000 Reduced expiry: false Complications: [vary‐revision‐sha1, show‐toc] CPU time usage: 3.111 seconds Real time usage: 3.456 seconds Preprocessor visited node count: 25216/1000000 Post‐expand include size: 910382/2097152 bytes Template argument size: 26738/2097152 bytes Highest expansion depth: 26/100 Expensive parser function count: 16/500 Unstrip recursion depth: 1/20 Unstrip post‐expand size: 952739/5000000 bytes Lua time usage: 1.907/10.000 seconds Lua memory usage: 9180859/52428800 bytes Lua Profile: ? 260 ms 14.0% MediaWiki\Extension\Scribunto\Engines\LuaSandbox\LuaSandboxCallback::match 220 ms 11.8% MediaWiki\Extension\Scribunto\Engines\LuaSandbox\LuaSandboxCallback::callParserFunction 200 ms 10.8% <mw.lua:694> 160 ms 8.6% 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