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<div class="vector-toc-text"> <span class="vector-toc-numb">2</span> <span>Mechanisms of evolution</span> </div> </a> <ul id="toc-Mechanisms_of_evolution-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Selection_pressure" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Selection_pressure"> <div class="vector-toc-text"> <span class="vector-toc-numb">3</span> <span>Selection pressure</span> </div> </a> <ul id="toc-Selection_pressure-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Functional_adaptations" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Functional_adaptations"> <div class="vector-toc-text"> <span class="vector-toc-numb">4</span> <span>Functional adaptations</span> </div> </a> <button aria-controls="toc-Functional_adaptations-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> <span 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class="vector-toc-link" href="#Tracking_bitten_prey"> <div class="vector-toc-text"> <span class="vector-toc-numb">4.3</span> <span>Tracking bitten prey</span> </div> </a> <ul id="toc-Tracking_bitten_prey-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Diet-based_atrophication" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#Diet-based_atrophication"> <div class="vector-toc-text"> <span class="vector-toc-numb">5</span> <span>Diet-based atrophication</span> </div> </a> <ul id="toc-Diet-based_atrophication-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1 vector-toc-list-item-expanded"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> <span class="vector-toc-numb">6</span> <span>References</span> </div> </a> <button aria-controls="toc-References-sublist" class="cdx-button cdx-button--weight-quiet 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Click here for more information." src="//upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/19px-Symbol_support_vote.svg.png" decoding="async" width="19" height="20" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/29px-Symbol_support_vote.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/39px-Symbol_support_vote.svg.png 2x" data-file-width="180" data-file-height="185" /></a></span></div></div> </div> <div id="siteSub" class="noprint">From Wikipedia, the free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></div></div> <div id="mw-content-text" class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Origin and diversification of snake venom through geologic time</div> <p class="mw-empty-elt"> </p> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Pacific_rattlesnake_(Crotalus_Oreganus)_juvenile_eating_(14232058164).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0f/Pacific_rattlesnake_%28Crotalus_Oreganus%29_juvenile_eating_%2814232058164%29.jpg/260px-Pacific_rattlesnake_%28Crotalus_Oreganus%29_juvenile_eating_%2814232058164%29.jpg" decoding="async" width="260" height="211" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0f/Pacific_rattlesnake_%28Crotalus_Oreganus%29_juvenile_eating_%2814232058164%29.jpg/390px-Pacific_rattlesnake_%28Crotalus_Oreganus%29_juvenile_eating_%2814232058164%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/0/0f/Pacific_rattlesnake_%28Crotalus_Oreganus%29_juvenile_eating_%2814232058164%29.jpg 2x" data-file-width="503" data-file-height="409" /></a><figcaption>The <a href="/wiki/Rattlesnake" title="Rattlesnake">rattlesnake</a> <i><a href="/wiki/Crotalus_oreganus" title="Crotalus oreganus">Crotalus oreganus</a></i> eating its prey, which it uses its venom to subdue</figcaption></figure><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline ol,.mw-parser-output .hlist.inline ul,.mw-parser-output .hlist dl dl,.mw-parser-output .hlist dl ol,.mw-parser-output .hlist dl ul,.mw-parser-output .hlist ol dl,.mw-parser-output .hlist ol ol,.mw-parser-output .hlist ol ul,.mw-parser-output .hlist ul dl,.mw-parser-output .hlist ul ol,.mw-parser-output .hlist ul ul{display:inline}.mw-parser-output .hlist .mw-empty-li{display:none}.mw-parser-output .hlist dt::after{content:": "}.mw-parser-output .hlist dd::after,.mw-parser-output .hlist li::after{content:" · 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.sidebar-title-with-pretitle a{color:var(--color-progressive)!important}}@media print{body.ns-0 .mw-parser-output .sidebar{display:none!important}}</style><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><table class="sidebar sidebar-collapse nomobile nowraplinks" style="border:2px solid #90ee90"><tbody><tr><td class="sidebar-pretitle">Part of a series on</td></tr><tr><th class="sidebar-title-with-pretitle" style="background:#90ee90;padding:0.2em;font-size:175%;font-weight:bold"><a href="/wiki/Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></th></tr><tr><td class="sidebar-image"><span class="notpageimage" typeof="mw:File"><a href="/wiki/File:Darwin%27s_finches_by_Gould.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/ae/Darwin%27s_finches_by_Gould.jpg/180px-Darwin%27s_finches_by_Gould.jpg" decoding="async" width="180" height="136" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/ae/Darwin%27s_finches_by_Gould.jpg/270px-Darwin%27s_finches_by_Gould.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/ae/Darwin%27s_finches_by_Gould.jpg/360px-Darwin%27s_finches_by_Gould.jpg 2x" data-file-width="1748" data-file-height="1319" /></a></span><div class="sidebar-caption"><a href="/wiki/Darwin%27s_finches" title="Darwin's finches">Darwin's finches</a> by <a href="/wiki/John_Gould" title="John Gould">John Gould</a></div></td></tr><tr><td class="sidebar-above" style="display:block; margin-bottom:0.55em; background-color: transparent;"> <div class="hlist"> <ul><li><a href="/wiki/Index_of_evolutionary_biology_articles" title="Index of evolutionary biology articles">Index</a></li> <li><a href="/wiki/Introduction_to_evolution" title="Introduction to evolution">Introduction</a></li> <li><a href="/wiki/Evolution" title="Evolution">Main</a></li> <li><a href="/wiki/Outline_of_evolution" title="Outline of evolution">Outline</a></li></ul> <div class="hlist"> </div> <ul><li><a href="/wiki/Glossary_of_evolutionary_biology" class="mw-redirect" title="Glossary of evolutionary biology">Glossary</a></li> <li><a href="/wiki/Evidence_of_common_descent" title="Evidence of common descent">Evidence</a></li> <li><a href="/wiki/History_of_life" title="History of life">History</a></li></ul> </div></td></tr><tr><td class="sidebar-content" style="background:transparent;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">Processes and outcomes</div><div class="sidebar-list-content mw-collapsible-content plainlist"><div class="hlist"> <ul><li><a href="/wiki/Population_genetics" title="Population genetics">Population genetics</a></li> <li><a href="/wiki/Genetic_variation" title="Genetic variation">Variation</a></li> <li><a href="/wiki/Genetic_diversity" title="Genetic diversity">Diversity</a></li> <li><a href="/wiki/Mutation" title="Mutation">Mutation</a></li> <li><a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a></li> <li><a href="/wiki/Adaptation" title="Adaptation">Adaptation</a></li> <li><a href="/wiki/Polymorphism_(biology)" title="Polymorphism (biology)">Polymorphism</a></li> <li><a href="/wiki/Genetic_drift" title="Genetic drift">Genetic drift</a></li> <li><a href="/wiki/Gene_flow" title="Gene flow">Gene flow</a></li> <li><a href="/wiki/Speciation" title="Speciation">Speciation</a></li> <li><a href="/wiki/Adaptive_radiation" title="Adaptive radiation">Adaptive radiation</a></li> <li><a href="/wiki/Cooperation_(evolution)" title="Cooperation (evolution)">Co-operation</a></li> <li><a href="/wiki/Coevolution" title="Coevolution">Coevolution</a></li> <li><a href="/wiki/Coextinction" title="Coextinction">Coextinction</a></li> <li><a href="/wiki/Contingency_(evolutionary_biology)" title="Contingency (evolutionary biology)">Contingency</a></li> <li><a href="/wiki/Divergent_evolution" title="Divergent evolution">Divergence</a></li> <li><a href="/wiki/Convergent_evolution" title="Convergent evolution">Convergence</a></li> <li><a href="/wiki/Parallel_evolution" title="Parallel evolution">Parallel evolution</a></li> <li><a href="/wiki/Extinction" title="Extinction">Extinction</a></li></ul> </div></div></div></td> </tr><tr><td class="sidebar-content" style="background:transparent;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">Natural history</div><div class="sidebar-list-content mw-collapsible-content plainlist"><div class="hlist"> <ul><li><a href="/wiki/Abiogenesis" title="Abiogenesis">Origin of life</a></li> <li><a href="/wiki/Common_descent" title="Common descent">Common descent</a></li> <li><a href="/wiki/History_of_life" title="History of life">History of life</a></li> <li><a href="/wiki/Timeline_of_the_evolutionary_history_of_life" title="Timeline of the evolutionary history of life">Timeline of evolution</a></li> <li><a href="/wiki/Human_evolution" title="Human evolution">Human evolution</a> <ul><li><a href="/wiki/Recent_human_evolution" title="Recent human evolution">Recent human evolution</a></li></ul></li> <li><a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">Phylogeny</a></li> <li><a href="/wiki/Biodiversity" title="Biodiversity">Biodiversity</a></li> <li><a href="/wiki/Biogeography" title="Biogeography">Biogeography</a></li> <li><a href="/wiki/Taxonomy_(biology)" title="Taxonomy (biology)">Classification</a></li> <li><a href="/wiki/Evolutionary_taxonomy" title="Evolutionary taxonomy">Evolutionary taxonomy</a></li> <li><a href="/wiki/Cladistics" title="Cladistics">Cladistics</a></li> <li><a href="/wiki/Transitional_fossil" title="Transitional fossil">Transitional fossil</a></li> <li><a href="/wiki/Extinction_event" title="Extinction event">Extinction event</a></li></ul> </div></div></div></td> </tr><tr><td class="sidebar-content" style="background:transparent;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">History of evolutionary theory</div><div class="sidebar-list-content mw-collapsible-content plainlist"><div class="hlist"> <ul><li><a href="/wiki/History_of_evolutionary_thought" title="History of evolutionary thought">Overview</a></li> <li><a href="/wiki/Evolutionary_ideas_of_the_Renaissance_and_Enlightenment" title="Evolutionary ideas of the Renaissance and Enlightenment">Renaissance</a></li> <li><a href="/wiki/Transmutation_of_species" title="Transmutation of species">Before Darwin</a></li> <li><a href="/wiki/Charles_Darwin" title="Charles Darwin">Darwin</a></li> <li><i><a href="/wiki/On_the_Origin_of_Species" title="On the Origin of Species">Origin of Species</a></i></li> <li><a href="/wiki/The_eclipse_of_Darwinism" title="The eclipse of Darwinism">Before synthesis</a></li> <li><a href="/wiki/Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis</a></li> <li><a href="/wiki/History_of_molecular_evolution" title="History of molecular evolution">Molecular evolution</a></li> <li><a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evo-devo</a></li> <li><a href="/wiki/Current_research_in_evolutionary_biology" class="mw-redirect" title="Current research in evolutionary biology">Current research</a></li> <li><a href="/wiki/History_of_speciation" title="History of speciation">History of speciation</a></li> <li><a href="/wiki/History_of_paleontology" title="History of paleontology">History of paleontology</a> (<a href="/wiki/Timeline_of_paleontology" title="Timeline of paleontology">timeline</a>)</li></ul> </div></div></div></td> </tr><tr><td class="sidebar-content" style="background:transparent;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">Fields and applications</div><div class="sidebar-list-content mw-collapsible-content plainlist"> <ul><li><a href="/wiki/Applications_of_evolution" title="Applications of evolution">Applications of evolution</a></li> <li><a href="/wiki/Biosocial_criminology" title="Biosocial criminology">Biosocial criminology</a></li> <li><a href="/wiki/Ecological_genetics" title="Ecological genetics">Ecological genetics</a></li> <li><a href="/wiki/Evolutionary_aesthetics" title="Evolutionary aesthetics">Evolutionary aesthetics</a></li> <li><a href="/wiki/Evolutionary_anthropology" title="Evolutionary anthropology">Evolutionary anthropology</a></li> <li><a href="/wiki/Evolutionary_computation" title="Evolutionary computation">Evolutionary computation</a></li> <li><a href="/wiki/Evolutionary_ecology" title="Evolutionary ecology">Evolutionary ecology</a></li> <li><a href="/wiki/Evolutionary_economics" title="Evolutionary economics">Evolutionary economics</a></li> <li><a href="/wiki/Evolutionary_epistemology" title="Evolutionary epistemology">Evolutionary epistemology</a></li> <li><a href="/wiki/Evolutionary_ethics" title="Evolutionary ethics">Evolutionary ethics</a></li> <li><a href="/wiki/Evolutionary_game_theory" title="Evolutionary game theory">Evolutionary game theory</a></li> <li><a href="/wiki/Evolutionary_linguistics" title="Evolutionary linguistics">Evolutionary linguistics</a></li> <li><a href="/wiki/Evolutionary_medicine" title="Evolutionary medicine">Evolutionary medicine</a></li> <li><a href="/wiki/Evolutionary_neuroscience" title="Evolutionary neuroscience">Evolutionary neuroscience</a></li> <li><a href="/wiki/Evolutionary_physiology" title="Evolutionary physiology">Evolutionary physiology</a></li> <li><a href="/wiki/Evolutionary_psychology" title="Evolutionary psychology">Evolutionary psychology</a></li> <li><a href="/wiki/Experimental_evolution" title="Experimental evolution">Experimental evolution</a></li> <li><a href="/wiki/Phylogenetics" title="Phylogenetics">Phylogenetics</a></li> <li><a href="/wiki/Paleontology" title="Paleontology">Paleontology</a></li> <li><a href="/wiki/Selective_breeding" title="Selective breeding">Selective breeding</a></li> <li><a href="/wiki/Laboratory_experiments_of_speciation" title="Laboratory experiments of speciation">Speciation experiments</a></li> <li><a href="/wiki/Sociobiology" title="Sociobiology">Sociobiology</a></li> <li><a href="/wiki/Island_biogeography" class="mw-redirect" title="Island biogeography">Island biogeography</a></li> <li><a href="/wiki/Systematics" title="Systematics">Systematics</a></li> <li><a href="/wiki/Universal_Darwinism" title="Universal Darwinism">Universal Darwinism</a></li></ul></div></div></td> </tr><tr><td class="sidebar-content" style="background:transparent;"> <div class="sidebar-list mw-collapsible mw-collapsed"><div class="sidebar-list-title" style="text-align:center;padding-bottom:0;background:#90ee90;font-size:100%;font-weight:bold;;color: var(--color-base)">Social implications</div><div class="sidebar-list-content mw-collapsible-content plainlist"> <ul><li><a href="/wiki/Eugenics" title="Eugenics">Eugenics</a></li> <li><a href="/wiki/Evolution_as_fact_and_theory" title="Evolution as fact and theory">Evolution as fact and theory</a></li> <li><a href="/wiki/Dysgenics" title="Dysgenics">Dysgenics</a></li> <li><a href="/wiki/Social_effects_of_evolutionary_theory" title="Social effects of evolutionary theory">Social effects</a></li> <li><a href="/wiki/Rejection_of_evolution_by_religious_groups" title="Rejection of evolution by religious groups">Creation–evolution controversy</a></li> <li><a href="/wiki/Theistic_evolution" title="Theistic evolution">Theistic evolution</a></li> <li><a href="/wiki/Objections_to_evolution" title="Objections to evolution">Objections to evolution</a></li> <li><a href="/wiki/Level_of_support_for_evolution" title="Level of support for evolution">Level of support</a></li> <li><a href="/wiki/Nature_versus_nurture" title="Nature versus nurture">Nature-nurture controversy</a></li></ul></div></div></td> </tr><tr><td class="sidebar-below hlist"> <ul><li><span class="nowrap"><span class="nowrap"><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/16px-Tree_of_life.svg.png" decoding="async" width="16" height="13" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/09/Tree_of_life.svg/24px-Tree_of_life.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/09/Tree_of_life.svg/32px-Tree_of_life.svg.png 2x" data-file-width="915" data-file-height="760" /></a></span> </span><a href="/wiki/Portal:Evolutionary_biology" title="Portal:Evolutionary biology">Evolutionary biology portal</a></span></li> <li><span class="nowrap"><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></span></li></ul></td></tr><tr><td class="sidebar-navbar"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><style data-mw-deduplicate="TemplateStyles:r1239400231">.mw-parser-output .navbar{display:inline;font-size:88%;font-weight:normal}.mw-parser-output .navbar-collapse{float:left;text-align:left}.mw-parser-output .navbar-boxtext{word-spacing:0}.mw-parser-output .navbar ul{display:inline-block;white-space:nowrap;line-height:inherit}.mw-parser-output .navbar-brackets::before{margin-right:-0.125em;content:"[ "}.mw-parser-output .navbar-brackets::after{margin-left:-0.125em;content:" ]"}.mw-parser-output .navbar li{word-spacing:-0.125em}.mw-parser-output .navbar a>span,.mw-parser-output .navbar a>abbr{text-decoration:inherit}.mw-parser-output .navbar-mini abbr{font-variant:small-caps;border-bottom:none;text-decoration:none;cursor:inherit}.mw-parser-output .navbar-ct-full{font-size:114%;margin:0 7em}.mw-parser-output .navbar-ct-mini{font-size:114%;margin:0 4em}html.skin-theme-clientpref-night .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Evolutionary_biology" title="Template:Evolutionary biology"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Evolutionary_biology" title="Template talk:Evolutionary biology"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Evolutionary_biology" title="Special:EditPage/Template:Evolutionary biology"><abbr title="Edit this template">e</abbr></a></li></ul></div></td></tr></tbody></table> <p><a href="/wiki/Snake_venom" title="Snake venom">Venom in snakes</a> and some <a href="/wiki/Lizard" title="Lizard">lizards</a> is a form of <a href="/wiki/Saliva" title="Saliva">saliva</a> that has been modified into <a href="/wiki/Venom" title="Venom">venom</a> over its evolutionary history.<sup id="cite_ref-FOOTNOTEHargreaves_et_al._(a)2014_1-0" class="reference"><a href="#cite_note-FOOTNOTEHargreaves_et_al._(a)2014-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> In <a href="/wiki/Snake" title="Snake">snakes</a>, venom has evolved to kill or subdue prey, as well as to perform other diet-related functions.<sup id="cite_ref-FOOTNOTECasewell_et_al.2013218–220_2-0" class="reference"><a href="#cite_note-FOOTNOTECasewell_et_al.2013218–220-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> While snakes occasionally use their venom in self defense, this is not believed to have had a strong effect on venom evolution.<sup id="cite_ref-FOOTNOTEWard-Smith_et_al.2020_3-0" class="reference"><a href="#cite_note-FOOTNOTEWard-Smith_et_al.2020-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> The evolution of venom is thought to be responsible for the enormous expansion of snakes across the globe.<sup id="cite_ref-FOOTNOTEFry_et_al.2012a441–442_4-0" class="reference"><a href="#cite_note-FOOTNOTEFry_et_al.2012a441–442-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTEWuster_et_al.2008_5-0" class="reference"><a href="#cite_note-FOOTNOTEWuster_et_al.2008-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTELomonte_et_al._(a)2014326_6-0" class="reference"><a href="#cite_note-FOOTNOTELomonte_et_al._(a)2014326-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p><p>The evolutionary history of snake venom is a matter of debate. Historically, snake venom was believed to have evolved once, at the base of the <a href="/wiki/Caenophidia" title="Caenophidia">Caenophidia</a>, or derived snakes. Molecular studies published beginning in 2006 suggested that venom originated just once among a putative <a href="/wiki/Clade" title="Clade">clade</a> of reptiles, called <a href="/wiki/Toxicofera" title="Toxicofera">Toxicofera</a>, approximately 170 million years ago.<sup id="cite_ref-FOOTNOTEFry_et_al.2012a434–436_7-0" class="reference"><a href="#cite_note-FOOTNOTEFry_et_al.2012a434–436-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> Under this hypothesis, the original toxicoferan venom was a very simple set of proteins that were assembled in a pair of glands. Subsequently, this set of proteins <a href="/wiki/Adaptive_radiation" title="Adaptive radiation">diversified</a> in the various lineages of toxicoferans, including Serpentes, <a href="/wiki/Anguimorpha" title="Anguimorpha">Anguimorpha</a>, and <a href="/wiki/Iguania" class="mw-redirect" title="Iguania">Iguania</a>: several snake lineages also lost the ability to produce venom.<sup id="cite_ref-FOOTNOTEFry_et_al.2012a424–436_8-0" class="reference"><a href="#cite_note-FOOTNOTEFry_et_al.2012a424–436-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTECasewell_et_al.2013224–227_9-0" class="reference"><a href="#cite_note-FOOTNOTECasewell_et_al.2013224–227-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> The Toxicoferan hypothesis was challenged by studies in the mid-2010s, including a 2015 study which found that venom proteins had <a href="/wiki/Homolog" class="mw-redirect" title="Homolog">homologs</a> in many other tissues in the <a href="/wiki/Burmese_python" title="Burmese python">Burmese python</a>.<sup id="cite_ref-FOOTNOTEReyes-Velasco_et_al.2015_10-0" class="reference"><a href="#cite_note-FOOTNOTEReyes-Velasco_et_al.2015-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTEHargreaves_et_al._(b)2014153–155_11-0" class="reference"><a href="#cite_note-FOOTNOTEHargreaves_et_al._(b)2014153–155-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> The study therefore suggested that venom had evolved independently in different reptile lineages, including once in the Caenophid snakes.<sup id="cite_ref-FOOTNOTEReyes-Velasco_et_al.2015_10-1" class="reference"><a href="#cite_note-FOOTNOTEReyes-Velasco_et_al.2015-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> Venom containing most extant toxin families is believed to have been present in the <a href="/wiki/Last_common_ancestor" class="mw-redirect" title="Last common ancestor">last common ancestor</a> of the Caenophidia: these toxins subsequently underwent tremendous diversification, accompanied by changes in the morphology of venom glands and delivery systems.<sup id="cite_ref-FOOTNOTEXieDashevskyRokytaGhezellou2022_12-0" class="reference"><a href="#cite_note-FOOTNOTEXieDashevskyRokytaGhezellou2022-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p><p>Snake venom evolution is thought to be driven by an <a href="/wiki/Evolutionary_arms_race" title="Evolutionary arms race">evolutionary arms race</a> between venom proteins and prey physiology.<sup id="cite_ref-FOOTNOTECasewellJacksonLaustsenSunagar2020570–581_13-0" class="reference"><a href="#cite_note-FOOTNOTECasewellJacksonLaustsenSunagar2020570–581-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> The common mechanism of evolution is thought to be <a href="/wiki/Gene_duplication" title="Gene duplication">gene duplication</a> followed by <a href="/wiki/Natural_selection" title="Natural selection">natural selection</a> for <a href="/wiki/Adaptation" title="Adaptation">adaptive traits</a>.<sup id="cite_ref-FOOTNOTECasewell_et_al.2013222–223_14-0" class="reference"><a href="#cite_note-FOOTNOTECasewell_et_al.2013222–223-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> The adaptations produced by this process include venom more toxic to specific prey in several lineages,<sup id="cite_ref-FOOTNOTEBarlow_et_al.20092447–2448_15-0" class="reference"><a href="#cite_note-FOOTNOTEBarlow_et_al.20092447–2448-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTECalvete_et_al.20124094–4098_16-0" class="reference"><a href="#cite_note-FOOTNOTECalvete_et_al.20124094–4098-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTELi_et_al.2005_17-0" class="reference"><a href="#cite_note-FOOTNOTELi_et_al.2005-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> proteins that pre-digest prey,<sup id="cite_ref-FOOTNOTEMackessy2010_18-0" class="reference"><a href="#cite_note-FOOTNOTEMackessy2010-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> and a method to track down prey after a bite.<sup id="cite_ref-FOOTNOTESaviola_et_al.2013_19-0" class="reference"><a href="#cite_note-FOOTNOTESaviola_et_al.2013-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> These various adaptations of venom have also led to considerable debate about the definition of venom and venomous snakes.<sup id="cite_ref-FOOTNOTEFry_et_al.2012a443_20-0" class="reference"><a href="#cite_note-FOOTNOTEFry_et_al.2012a443-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> Changes in the diet of a lineage have been linked to atrophication of the venom.<sup id="cite_ref-FOOTNOTEFry_et_al.2012a424–436_8-1" class="reference"><a href="#cite_note-FOOTNOTEFry_et_al.2012a424–436-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTECasewell_et_al.2013224–227_9-1" class="reference"><a href="#cite_note-FOOTNOTECasewell_et_al.2013224–227-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Evolutionary_history">Evolutionary history</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_snake_venom&action=edit&section=1" title="Edit section: Evolutionary history"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The origin of venom is thought to have provided the catalyst for the rapid diversification of snakes in the <a href="/wiki/Cenozoic" title="Cenozoic">Cenozoic</a> period,<sup id="cite_ref-FOOTNOTEFry_et_al.2012a_21-0" class="reference"><a href="#cite_note-FOOTNOTEFry_et_al.2012a-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> particularly to the <a href="/wiki/Colubridae" title="Colubridae">Colubridae</a> and their colonization of the <a href="/wiki/Americas" title="Americas">Americas</a>.<sup id="cite_ref-FOOTNOTEWuster_et_al.2008_5-1" class="reference"><a href="#cite_note-FOOTNOTEWuster_et_al.2008-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> Scholars suggest that the reason for this huge expansion was the shift from a mechanical to a biochemical method of subduing prey.<sup id="cite_ref-FOOTNOTELomonte_et_al._(a)2014326–327_22-0" class="reference"><a href="#cite_note-FOOTNOTELomonte_et_al._(a)2014326–327-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTEMackessy20101464_23-0" class="reference"><a href="#cite_note-FOOTNOTEMackessy20101464-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> Snake venoms attack biological pathways and processes that are also targeted by venoms of other taxa; for instance, <a href="/wiki/Calcium_channel" title="Calcium channel">calcium channel</a> blockers have been found in snakes, spiders, and <a href="/wiki/Cone_snail" title="Cone snail">cone snails</a>, thus suggesting that venom exhibits <a href="/wiki/Convergent_evolution" title="Convergent evolution">convergent evolution</a>.<sup id="cite_ref-FOOTNOTECasewell_et_al.2013225–227_24-0" class="reference"><a href="#cite_note-FOOTNOTECasewell_et_al.2013225–227-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> Venom is common among <a href="/wiki/Primitive_(phylogenetics)" title="Primitive (phylogenetics)">derived</a> snake families.<sup id="cite_ref-FOOTNOTECasewellJacksonLaustsenSunagar2020570–581_13-1" class="reference"><a href="#cite_note-FOOTNOTECasewellJacksonLaustsenSunagar2020570–581-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> Venom containing most extant toxin families is believed to have been present in the <a href="/wiki/Last_common_ancestor" class="mw-redirect" title="Last common ancestor">last common ancestor</a> of the <a href="/wiki/Caenophidia" title="Caenophidia">Caenophidia</a>, also called Colubroidea. These toxins subsequently underwent tremendous diversification, accompanied by changes in the morphology of venom glands and delivery systems. This diversification is linked to the rapid global radiation of the advanced snakes.<sup id="cite_ref-FOOTNOTEXieDashevskyRokytaGhezellou2022_12-1" class="reference"><a href="#cite_note-FOOTNOTEXieDashevskyRokytaGhezellou2022-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> The tubular or grooved <a href="/wiki/Fang" title="Fang">fangs</a> snakes use to deliver their venom to their target have evolved multiple times, and are an example of <a href="/wiki/Convergent_evolution" title="Convergent evolution">convergent evolution</a>. The tubular fangs common to front-fanged snakes are believed to have evolved independently in <a href="/wiki/Viperidae" class="mw-redirect" title="Viperidae">Viperidae</a>, <a href="/wiki/Elapidae" title="Elapidae">Elapidae</a>, and <a href="/wiki/Atractaspidinae" class="mw-redirect" title="Atractaspidinae">Atractaspidinae</a>.<sup id="cite_ref-FOOTNOTEPalciLeBlancPanagiotopoulouCleuren2021_25-0" class="reference"><a href="#cite_note-FOOTNOTEPalciLeBlancPanagiotopoulouCleuren2021-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> </p> <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:85%;line-height:85%"> <tbody><tr> <td class="clade-label first"><b><a href="/wiki/Serpentes" class="mw-redirect" title="Serpentes">Serpentes</a></b> </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/Scolecophidia" title="Scolecophidia">Scolecophidia</a> <span typeof="mw:File"><a href="/wiki/File:Typhlops_vermicularis3_white_background.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c6/Typhlops_vermicularis3_white_background.jpg/70px-Typhlops_vermicularis3_white_background.jpg" decoding="async" width="70" height="18" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c6/Typhlops_vermicularis3_white_background.jpg/105px-Typhlops_vermicularis3_white_background.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c6/Typhlops_vermicularis3_white_background.jpg/140px-Typhlops_vermicularis3_white_background.jpg 2x" data-file-width="876" data-file-height="222" /></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/Booidea" title="Booidea">Booidea</a> inc. <a href="/wiki/Pythonidae" title="Pythonidae">Pythonidae</a> <span typeof="mw:File"><a href="/wiki/File:Python_natalensis_Smith_1840_white_background.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8f/Python_natalensis_Smith_1840_white_background.jpg/80px-Python_natalensis_Smith_1840_white_background.jpg" decoding="async" width="80" height="23" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8f/Python_natalensis_Smith_1840_white_background.jpg/120px-Python_natalensis_Smith_1840_white_background.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8f/Python_natalensis_Smith_1840_white_background.jpg/160px-Python_natalensis_Smith_1840_white_background.jpg 2x" data-file-width="1554" data-file-height="442" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"><a href="/wiki/Caenophidia" title="Caenophidia">Caenophidia</a> </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/Acrochordidae" class="mw-redirect" title="Acrochordidae">Acrochordidae</a> </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/Xenodermatidae" class="mw-redirect" title="Xenodermatidae">Xenodermatidae</a> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"><a href="/wiki/Colubroidea" title="Colubroidea">Colubroidea</a> </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/Pareatidae" class="mw-redirect" title="Pareatidae">Pareatidae</a> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"><b>[A]</b> </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"><b>[B]</b> </td> <td rowspan="2" class="clade-leaf"> <p><a href="/wiki/Viperidae" class="mw-redirect" title="Viperidae">Viperidae</a> <span typeof="mw:File"><a href="/wiki/File:Illustrations_of_the_zoology_of_South_Africa_(Bitis_caudalis).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b2/Illustrations_of_the_zoology_of_South_Africa_%28Bitis_caudalis%29.jpg/70px-Illustrations_of_the_zoology_of_South_Africa_%28Bitis_caudalis%29.jpg" decoding="async" width="70" height="19" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b2/Illustrations_of_the_zoology_of_South_Africa_%28Bitis_caudalis%29.jpg/105px-Illustrations_of_the_zoology_of_South_Africa_%28Bitis_caudalis%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b2/Illustrations_of_the_zoology_of_South_Africa_%28Bitis_caudalis%29.jpg/140px-Illustrations_of_the_zoology_of_South_Africa_%28Bitis_caudalis%29.jpg 2x" data-file-width="1366" data-file-height="368" /></a></span> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"><b>[C]</b> </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/Homalopsidae" title="Homalopsidae">Homalopsidae</a> </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/Colubridae" title="Colubridae">Colubridae</a> <span typeof="mw:File"><a href="/wiki/File:Xenochrophis_piscator_1_Hardwicke_white_background.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/95/Xenochrophis_piscator_1_Hardwicke_white_background.jpg/80px-Xenochrophis_piscator_1_Hardwicke_white_background.jpg" decoding="async" width="80" height="29" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/95/Xenochrophis_piscator_1_Hardwicke_white_background.jpg/120px-Xenochrophis_piscator_1_Hardwicke_white_background.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/95/Xenochrophis_piscator_1_Hardwicke_white_background.jpg/160px-Xenochrophis_piscator_1_Hardwicke_white_background.jpg 2x" data-file-width="761" data-file-height="279" /></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/Lamprophiidae" title="Lamprophiidae">Lamprophiidae</a> </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/Elapidae" title="Elapidae">Elapidae</a> <span typeof="mw:File"><a href="/wiki/File:Bilder-Atlas_zur_wissenschaftlich-popul%C3%A4ren_Naturgeschichte_der_Wirbelthiere_(Naja_naja).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/82/Bilder-Atlas_zur_wissenschaftlich-popul%C3%A4ren_Naturgeschichte_der_Wirbelthiere_%28Naja_naja%29.jpg/70px-Bilder-Atlas_zur_wissenschaftlich-popul%C3%A4ren_Naturgeschichte_der_Wirbelthiere_%28Naja_naja%29.jpg" decoding="async" width="70" height="50" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/82/Bilder-Atlas_zur_wissenschaftlich-popul%C3%A4ren_Naturgeschichte_der_Wirbelthiere_%28Naja_naja%29.jpg/105px-Bilder-Atlas_zur_wissenschaftlich-popul%C3%A4ren_Naturgeschichte_der_Wirbelthiere_%28Naja_naja%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/82/Bilder-Atlas_zur_wissenschaftlich-popul%C3%A4ren_Naturgeschichte_der_Wirbelthiere_%28Naja_naja%29.jpg/140px-Bilder-Atlas_zur_wissenschaftlich-popul%C3%A4ren_Naturgeschichte_der_Wirbelthiere_%28Naja_naja%29.jpg 2x" data-file-width="708" data-file-height="505" /></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 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> <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><small> A cladogram adapted from Fry et al. (2012) showing a subset of suggested protein recruitment events. [A]: 13 toxin families, including <a href="/wiki/Calciseptine#Three-fingered_toxin_family" title="Calciseptine">3FTx</a> and <a href="/wiki/Metalloprotease" class="mw-redirect" title="Metalloprotease">metalloprotease</a>. [B]: 2 toxin families, including PLA2 Type IIA and the P-II class of snake venom metalloproteases [C]: 2 toxin families, including PLA2 Type IB.<sup id="cite_ref-FOOTNOTEFry_et_al.2012a435_26-0" class="reference"><a href="#cite_note-FOOTNOTEFry_et_al.2012a435-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTEAlmeidaVialaNachtigallBroe2021_27-0" class="reference"><a href="#cite_note-FOOTNOTEAlmeidaVialaNachtigallBroe2021-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup></small></p></blockquote> <p>Until the use of <a href="/wiki/DNA_sequencing" title="DNA sequencing">gene sequencing</a> to create <a href="/wiki/Phylogenetic_tree" title="Phylogenetic tree">phylogenetic trees</a> became practical, phylogenies were created on the basis of <a href="/wiki/Morphology_(biology)" title="Morphology (biology)">morphology</a>. Such traditional phylogenies suggested that venom originated along multiple branches among <a href="/wiki/Squamata" title="Squamata">Squamata</a> approximately 100 million years ago: in the <a href="/wiki/Caenophidia" title="Caenophidia">Caenophidia</a>, or derived snakes, and in the lizard genus <i><a href="/wiki/Heloderma" title="Heloderma">Heloderma</a></i>.<sup id="cite_ref-FOOTNOTEFry_et_al.2012a434–436_7-1" class="reference"><a href="#cite_note-FOOTNOTEFry_et_al.2012a434–436-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTEReyes-Velasco_et_al.2015_10-2" class="reference"><a href="#cite_note-FOOTNOTEReyes-Velasco_et_al.2015-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTESunagarAbraham2021_28-0" class="reference"><a href="#cite_note-FOOTNOTESunagarAbraham2021-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> Studies using <a href="/wiki/Nuclear_gene" title="Nuclear gene">nuclear gene</a> sequences in the mid-2000s and early 2010s found the presence of venom proteins in the lizard <a href="/wiki/Clade" title="Clade">clades</a> <a href="/wiki/Anguimorpha" title="Anguimorpha">Anguimorpha</a> and <a href="/wiki/Iguania" class="mw-redirect" title="Iguania">Iguania</a> similar to those of snakes, and suggested that together with Serpentes, these formed a <a href="/wiki/Clade" title="Clade">clade</a>, which they named "<a href="/wiki/Toxicofera" title="Toxicofera">Toxicofera</a>".<sup id="cite_ref-FOOTNOTEFry_et_al.2012a434–436_7-2" class="reference"><a href="#cite_note-FOOTNOTEFry_et_al.2012a434–436-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTESunagarAbraham2021_28-1" class="reference"><a href="#cite_note-FOOTNOTESunagarAbraham2021-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> This led to the theory that venom originated only once within the entire lineage approximately 170 million years ago. This ancestral venom was described as consisting of a very simple set of proteins, assembled in a pair of glands.<sup id="cite_ref-FOOTNOTEFry_et_al.2012a434–436_7-3" class="reference"><a href="#cite_note-FOOTNOTEFry_et_al.2012a434–436-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> The venoms of the different lineages then diversified and evolved independently, along with their means of injecting venom into prey.<sup id="cite_ref-FOOTNOTEFry_et_al.2012a434–436_7-4" class="reference"><a href="#cite_note-FOOTNOTEFry_et_al.2012a434–436-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> This diversification included the independent evolution of front-fanged venom delivery from the ancestral rear-fanged venom delivery system.<sup id="cite_ref-FOOTNOTEFryWuster2004870_29-0" class="reference"><a href="#cite_note-FOOTNOTEFryWuster2004870-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> The single origin hypothesis also suggests that venom systems subsequently atrophied, or were completely lost, independently in a number of lineages.<sup id="cite_ref-FOOTNOTEFry_et_al.2012a443_20-1" class="reference"><a href="#cite_note-FOOTNOTEFry_et_al.2012a443-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> The phylogenetic position of Iguania within Toxicofera is supported by most molecular studies, but not by morphological ones.<sup id="cite_ref-FOOTNOTEMountBrown2022973–985_30-0" class="reference"><a href="#cite_note-FOOTNOTEMountBrown2022973–985-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> </p><p>The "Toxicoferan hypothesis" was subsequently challenged. A study performed in 2014 found that homologs of 16 venom proteins, which had been used to support the single origin hypothesis, were all expressed at high levels in a number of body tissues.<sup id="cite_ref-FOOTNOTEHargreaves_et_al._(b)2014153–155_11-1" class="reference"><a href="#cite_note-FOOTNOTEHargreaves_et_al._(b)2014153–155-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> The authors therefore suggested that previous research, which had found venom proteins to be conserved across the supposed Toxicoferan lineage, might have misinterpreted the presence of more generic "housekeeping" genes across this lineage, as a result of only sampling certain tissues within the reptiles' bodies. Therefore, the authors suggested that instead of evolving just once in an ancestral reptile, venom evolved independently in multiple lineages, including once prior to the radiation of the "advanced" snakes.<sup id="cite_ref-FOOTNOTEHargreaves_et_al._(b)2014153–155_11-2" class="reference"><a href="#cite_note-FOOTNOTEHargreaves_et_al._(b)2014153–155-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> A 2015 study found that homologs of the so-called "toxic" genes were present in numerous tissues of a non-venomous snake, the <a href="/wiki/Burmese_python" title="Burmese python">Burmese python</a>. One of the authors stated that they had found homologs to the venom genes in many tissues outside the oral glands, where venom genes might be expected. This demonstrated the weaknesses of only analyzing <a href="/wiki/Transcriptome" title="Transcriptome">transcriptomes</a> (the total <a href="/wiki/Messenger_RNA" title="Messenger RNA">messenger RNA</a> in a cell).<sup id="cite_ref-FOOTNOTEReyes-Velasco_et_al.2015_10-3" class="reference"><a href="#cite_note-FOOTNOTEReyes-Velasco_et_al.2015-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> The team suggested that pythons represented a period in snake evolution before major venom development. The researchers also found that the expansion of venom gene families occurred mostly in highly venomous <a href="/wiki/Colubroidea" title="Colubroidea">caenophidian snakes</a> (also referred to as "colubrid snakes"), thus suggesting that most venom evolution took place after this lineage diverged from other snakes.<sup id="cite_ref-FOOTNOTEReyes-Velasco_et_al.2015_10-4" class="reference"><a href="#cite_note-FOOTNOTEReyes-Velasco_et_al.2015-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> The debate over the Toxicoferan hypothesis is driven in part by disagreement over the definition of a venom.<sup id="cite_ref-FOOTNOTEReyes-Velasco_et_al.2015_10-5" class="reference"><a href="#cite_note-FOOTNOTEReyes-Velasco_et_al.2015-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTEHargreaves_et_al._(b)2014_31-0" class="reference"><a href="#cite_note-FOOTNOTEHargreaves_et_al._(b)2014-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> As of 2022, the Toxicoferan hypothesis remains a prevalent view.<sup id="cite_ref-FOOTNOTERaoKalogeropoulosAllentoftGopalakrishnan2022_32-0" class="reference"><a href="#cite_note-FOOTNOTERaoKalogeropoulosAllentoftGopalakrishnan2022-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Mechanisms_of_evolution">Mechanisms of evolution</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_snake_venom&action=edit&section=2" title="Edit section: Mechanisms of evolution"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:1poc.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/28/1poc.png/220px-1poc.png" decoding="async" width="220" height="248" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/28/1poc.png/330px-1poc.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/28/1poc.png/440px-1poc.png 2x" data-file-width="522" data-file-height="589" /></a><figcaption><a href="/wiki/Phospholipase_A2" title="Phospholipase A2">Phospholipase A2</a>, an <a href="/wiki/Enzyme" title="Enzyme">enzyme</a> found in normal tissue that has been adapted in certain snake venoms. The example pictured here is found in <a href="/wiki/Bee_sting" title="Bee sting">bee stings</a>.</figcaption></figure> <p>The primary mechanism for the diversification of venom is thought to be the <a href="/wiki/Gene_duplication" title="Gene duplication">duplication of gene</a> coding for other tissues, followed by their expression in the venom glands. The proteins then evolved into various venom proteins through natural selection. This process, known as the birth-and-death model, is responsible for several of the protein recruitment events in snake venom.<sup id="cite_ref-FOOTNOTECasewell_et_al.2013223_33-0" class="reference"><a href="#cite_note-FOOTNOTECasewell_et_al.2013223-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTECasewellJacksonLaustsenSunagar2020570–581_13-2" class="reference"><a href="#cite_note-FOOTNOTECasewellJacksonLaustsenSunagar2020570–581-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> These duplications occurred in a variety of tissue types with a number of ancestral functions. Notable examples include <a href="/wiki/Three-finger_toxin" title="Three-finger toxin">3FTx</a>, ancestrally a <a href="/wiki/Neurotransmitter" title="Neurotransmitter">neurotransmitter</a> found in the brain, which has adapted into a neurotoxin that binds and blocks <a href="/wiki/Acetylcholine_receptor" title="Acetylcholine receptor">acetylcholine receptors</a>. Another example is <a href="/wiki/Phospholipase_A2" title="Phospholipase A2">phospholipase A2</a> (PLA2) type IIA, ancestrally involved with <a href="/wiki/Inflammation" title="Inflammation">inflammatory processes</a> in normal tissue, which has evolved into venom capable of triggering <a href="/wiki/Lipase" title="Lipase">lipase</a> activity and tissue destruction.<sup id="cite_ref-FOOTNOTEFry_et_al.2012a_21-1" class="reference"><a href="#cite_note-FOOTNOTEFry_et_al.2012a-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> The change in function of PLA2, in particular, has been well documented; there is evidence of several separate gene duplication events, often associated with the origin of new snake species.<sup id="cite_ref-FOOTNOTELynch2007_34-0" class="reference"><a href="#cite_note-FOOTNOTELynch2007-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Non-allelic_homologous_recombination" title="Non-allelic homologous recombination">Non-allelic homologous recombination</a> induced by transposon invasion (or recombination between DNA sequences that are similar, but not <a href="/wiki/Allele" title="Allele">alleles</a>) has been proposed as the mechanism of duplication of PLA2 genes in rattlesnakes, as an explanation for its rapid evolution.<sup id="cite_ref-FOOTNOTEDowell_et_al.2016_35-0" class="reference"><a href="#cite_note-FOOTNOTEDowell_et_al.2016-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> These venom proteins have also occasionally been recruited back into tissue genes.<sup id="cite_ref-FOOTNOTECasewell_et_al.2013223–224_36-0" class="reference"><a href="#cite_note-FOOTNOTECasewell_et_al.2013223–224-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> </p><p>Gene duplication is not the only way that venom has become more diverse. There have been instances of new venom proteins generated by <a href="/wiki/Alternative_splicing" title="Alternative splicing">alternative splicing</a>. The <a href="/wiki/Elapidae" title="Elapidae">Elapid</a> snake <i><a href="/wiki/Bungarus_fasciatus" class="mw-redirect" title="Bungarus fasciatus">Bungarus fasciatus</a></i>, for example, possesses a gene that is alternatively spliced to yield both a venom component and a physiological protein.<sup id="cite_ref-FOOTNOTECasewell_et_al.2013223–224_36-1" class="reference"><a href="#cite_note-FOOTNOTECasewell_et_al.2013223–224-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> Further diversification may have occurred by gene loss of specific venom components. For instance, the rattlesnake ancestor is believed to have had the PLA2 genes for a heterodimeric neurotoxin now found in Crotalus scutulatus, but those genes are absent in modern non-neurotoxic <i>Crotalus</i> species; the PLA2 genes for the Lys49-myotoxin supposedly existing in the common ancestor of rattlesnakes were also lost several times on recent lineages to extant species <sup id="cite_ref-FOOTNOTEDowell_et_al.2016_35-1" class="reference"><a href="#cite_note-FOOTNOTEDowell_et_al.2016-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> Domain loss has also been implicated in venom neofunctionalization. Investigation of the evolutionary history of viperid SVMP venom genes revealed repeated occasions of domain loss, coupled with significant positive selection in most of the phylogenetic branches where domain loss was thought to have occurred.<sup id="cite_ref-FOOTNOTECasewell_et_al.2011_37-0" class="reference"><a href="#cite_note-FOOTNOTECasewell_et_al.2011-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup> Venom toxins have also evolved via the gene "hijacking" or "co-opting", or the change in function of unrelated genes. A 2021 study suggested that co-opting explained the evolution of most types of toxins, but not that of the toxins that are most abundant in snake venom.<sup id="cite_ref-FOOTNOTEAlmeidaVialaNachtigallBroe2021_27-1" class="reference"><a href="#cite_note-FOOTNOTEAlmeidaVialaNachtigallBroe2021-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> </p><p>Protein recruitment events have occurred at different points in the evolutionary history of snakes. For example, the 3FTX protein family is absent in the viperid lineage, suggesting that it was recruited into snake venom after the viperid snakes branched off from the remaining colubroidae.<sup id="cite_ref-FOOTNOTEFryWuster2004871_38-0" class="reference"><a href="#cite_note-FOOTNOTEFryWuster2004871-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> PLA2 is thought to have been recruited at least two separate times into snake venom, once in elapids and once in viperids, displaying convergent evolution of this protein into a toxin.<sup id="cite_ref-FOOTNOTEFry_et_al.2012b_39-0" class="reference"><a href="#cite_note-FOOTNOTEFry_et_al.2012b-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTEMikheyevBarua2019_40-0" class="reference"><a href="#cite_note-FOOTNOTEMikheyevBarua2019-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> A 2019 study suggested that gene duplication could have allowed different toxins to evolve independently, allowing snakes to experiment with their venom profiles and explore new and effective venom formulations.<sup id="cite_ref-FOOTNOTEMikheyevBarua2019_40-1" class="reference"><a href="#cite_note-FOOTNOTEMikheyevBarua2019-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> This was proposed as one of the ways snakes have diversified their venom formulations through millions of years of evolution.<sup id="cite_ref-FOOTNOTEMikheyevBarua2019_40-2" class="reference"><a href="#cite_note-FOOTNOTEMikheyevBarua2019-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup> The various recruitment events had resulted in snake venom evolving into a very complex mixture of proteins. The venom of rattlesnakes, for example, includes nearly 40 different proteins from different protein families,<sup id="cite_ref-FOOTNOTEGibbsMackessy2009_41-0" class="reference"><a href="#cite_note-FOOTNOTEGibbsMackessy2009-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> and other snake venoms have been found to contain more than 100 distinct proteins.<sup id="cite_ref-FOOTNOTELomonte_et_al._(a)2014326–327_22-1" class="reference"><a href="#cite_note-FOOTNOTELomonte_et_al._(a)2014326–327-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> The composition of this mixture has been shown to vary geographically, and to be related to the prey species available in a certain region.<sup id="cite_ref-FOOTNOTELi_et_al.2005_17-1" class="reference"><a href="#cite_note-FOOTNOTELi_et_al.2005-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> Snake venom has generally evolved very quickly, with changes occurring faster in the venom than in the rest of the organism.<sup id="cite_ref-FOOTNOTELomonte_et_al._(a)2014334_42-0" class="reference"><a href="#cite_note-FOOTNOTELomonte_et_al._(a)2014334-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Selection_pressure">Selection pressure</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_snake_venom&action=edit&section=3" title="Edit section: Selection pressure"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Long-standing hypotheses of snake venom evolution have assumed that most snakes inject far more venom into their prey than is required to kill them; thus, venom composition would not be subject to natural selection. This is known as the "overkill" hypothesis.<sup id="cite_ref-FOOTNOTEBarlow_et_al.20092443_43-0" class="reference"><a href="#cite_note-FOOTNOTEBarlow_et_al.20092443-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> However, recent studies of the molecular history of snake venom have contradicted this, instead finding evidence of rapid adaptive evolution in many different clades, including the carpet vipers, <i><a href="/wiki/Echis" title="Echis">Echis</a></i>,<sup id="cite_ref-FOOTNOTEBarlow_et_al.20092447_44-0" class="reference"><a href="#cite_note-FOOTNOTEBarlow_et_al.20092447-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> the ground rattlesnakes, <i><a href="/wiki/Sistrurus" title="Sistrurus">Sistrurus</a></i>,<sup id="cite_ref-FOOTNOTEGibbsMackessy2009_41-1" class="reference"><a href="#cite_note-FOOTNOTEGibbsMackessy2009-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> and the <a href="/wiki/Malayan_pit_viper" class="mw-redirect" title="Malayan pit viper">Malayan pit viper</a>,<sup id="cite_ref-FOOTNOTECasewell_et_al.2013220_45-0" class="reference"><a href="#cite_note-FOOTNOTECasewell_et_al.2013220-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> as well as generally in the diversification of PLA2 proteins.<sup id="cite_ref-FOOTNOTELynch2007_34-1" class="reference"><a href="#cite_note-FOOTNOTELynch2007-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> There is phylogenetic evidence of <a href="/wiki/Positive_selection" class="mw-redirect" title="Positive selection">positive selection</a> and rapid rates of gene gain and loss in venom genes of <i>Sistrurus</i> taxa feeding on different prey.<sup id="cite_ref-FOOTNOTEGibbsRossiter2008_46-0" class="reference"><a href="#cite_note-FOOTNOTEGibbsRossiter2008-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> As of 2019, evidence existed both of "overkill" occurring in some lineages, and rapid adaptive evolution, and an <a href="/wiki/Evolutionary_arms_race" title="Evolutionary arms race">evolutionary arms race</a> with prey physiology, in many others.<sup id="cite_ref-FOOTNOTEHealyCarboneJackson527–537_47-0" class="reference"><a href="#cite_note-FOOTNOTEHealyCarboneJackson527–537-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> </p><p>The genes that code for venom proteins in some snake genera have a proportion of <a href="/wiki/Synonymous_mutations" class="mw-redirect" title="Synonymous mutations">synonymous mutations</a> that is lower than would be expected if venom were evolving through <a href="/wiki/Neutral_theory_of_molecular_evolution" title="Neutral theory of molecular evolution">neutral evolutionary processes</a>; the non-synonymous mutation rate, however, was found higher in many cases, indicating directional selection. <sup id="cite_ref-FOOTNOTELynch2007_34-2" class="reference"><a href="#cite_note-FOOTNOTELynch2007-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> In addition, snake venom is metabolically costly for a snake to produce, which scientists have suggested as further evidence that a selection pressure exists on snake venom (in this case, to minimize the volume of venom required).<sup id="cite_ref-FOOTNOTEBarlow_et_al.20092443_43-1" class="reference"><a href="#cite_note-FOOTNOTEBarlow_et_al.20092443-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> The use of <a href="/wiki/Model_organism" title="Model organism">model organisms</a>, rather than snakes' natural prey, to study prey toxicity, has been suggested as a reason why the "overkill" hypothesis may have been overemphasized.<sup id="cite_ref-FOOTNOTECasewell_et_al.2013220–221_48-0" class="reference"><a href="#cite_note-FOOTNOTECasewell_et_al.2013220–221-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> However, the pitviper genus <i><a href="/wiki/Agkistrodon" title="Agkistrodon">Agkistrodon</a></i> has been found to be an exception to this; the composition of venom in <i>Agkistrodon</i> has been found to be related to the position of the species within the phylogeny, suggesting that those venoms have evolved mostly through neutral processes (<a href="/wiki/Mutation" title="Mutation">mutation</a> and <a href="/wiki/Genetic_drift" title="Genetic drift">genetic drift</a>), and that there may be significant variation in the selection pressure upon various snake venoms.<sup id="cite_ref-FOOTNOTELomonte_et_al._(b)2014112–114_49-0" class="reference"><a href="#cite_note-FOOTNOTELomonte_et_al._(b)2014112–114-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> </p><p>Several studies have found evidence that venom and resistance to venom in prey species have evolved in a <a href="/wiki/Evolutionary_arms_race" title="Evolutionary arms race">coevolutionary arms race</a>.<sup id="cite_ref-FOOTNOTEBarlow_et_al.20092443_43-2" class="reference"><a href="#cite_note-FOOTNOTEBarlow_et_al.20092443-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup> For example, wood rats of the genus <i><a href="/wiki/Neotoma" class="mw-redirect" title="Neotoma">Neotoma</a></i> have a high degree of resistance to the venom of rattlesnakes, suggesting that the rats have evolved in response to the snake venom, thus renewing selection pressure upon the snakes.<sup id="cite_ref-FOOTNOTEGibbsMackessy2009_41-2" class="reference"><a href="#cite_note-FOOTNOTEGibbsMackessy2009-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> Resistance to venoms of sympatric predatory snake species has been found in eels,<sup id="cite_ref-FOOTNOTEHeatwolePoran1995_50-0" class="reference"><a href="#cite_note-FOOTNOTEHeatwolePoran1995-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup> ground squirrels,<sup id="cite_ref-FOOTNOTEBiardiChienCoss2005_51-0" class="reference"><a href="#cite_note-FOOTNOTEBiardiChienCoss2005-51"><span class="cite-bracket">[</span>51<span class="cite-bracket">]</span></a></sup> rock squirrels,<sup id="cite_ref-FOOTNOTEBiardiCoss2011_52-0" class="reference"><a href="#cite_note-FOOTNOTEBiardiCoss2011-52"><span class="cite-bracket">[</span>52<span class="cite-bracket">]</span></a></sup> and Eastern gray squirrels.<sup id="cite_ref-FOOTNOTEPomento_et_al.2016_53-0" class="reference"><a href="#cite_note-FOOTNOTEPomento_et_al.2016-53"><span class="cite-bracket">[</span>53<span class="cite-bracket">]</span></a></sup> All these studies suggested a co-evolutionary arms race between prey and predator, indicating another potential selection pressure on snake venom to increase or innovate toxicity. The selection pressure on snake venom is thought to be selecting for functional diversity within the proteins in venom, both within a given species, and across species.<sup id="cite_ref-FOOTNOTESanz_et_al.20062098–2099_54-0" class="reference"><a href="#cite_note-FOOTNOTESanz_et_al.20062098–2099-54"><span class="cite-bracket">[</span>54<span class="cite-bracket">]</span></a></sup> In addition to prey physiology, evidence exists that snake venom has evolved in response to the physiology of predators.<sup id="cite_ref-FOOTNOTECasewellJacksonLaustsenSunagar2020570–581_13-3" class="reference"><a href="#cite_note-FOOTNOTECasewellJacksonLaustsenSunagar2020570–581-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p><p>Besides diet, there are other possible pressures on snake venom composition. A 2019 study found that larger body mass and smaller ecological habitats were correlated with increased venom yield.<sup id="cite_ref-FOOTNOTEHealyCarboneJackson527–537_47-1" class="reference"><a href="#cite_note-FOOTNOTEHealyCarboneJackson527–537-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> Another study found that weather and temperature had stronger correlations with snake venom than diets or types of prey.<sup id="cite_ref-FOOTNOTEZancolli_et_al.2019_55-0" class="reference"><a href="#cite_note-FOOTNOTEZancolli_et_al.2019-55"><span class="cite-bracket">[</span>55<span class="cite-bracket">]</span></a></sup> While venomous snakes use their venoms in defence (hence the problem of <a href="/wiki/Snakebite" title="Snakebite">snakebite</a> in humans), it is not well known to what extent natural selection for defence has driven venom evolution. The venoms of the Texas coral snake, <i><a href="/wiki/Micrurus_tener" class="mw-redirect" title="Micrurus tener">Micrurus tener</a></i>, and other species of <i><a href="/wiki/Micrurus" title="Micrurus">Micrurus</a></i> have been found to contain toxins with specific pain-inducing activity, suggesting a defensive function.<sup id="cite_ref-FOOTNOTEBohlen2011_56-0" class="reference"><a href="#cite_note-FOOTNOTEBohlen2011-56"><span class="cite-bracket">[</span>56<span class="cite-bracket">]</span></a></sup> However, a questionnaire survey of snakebite patients bitten by a wide variety of venomous species showed that pain after most snakebites is of slow onset, arguing against widespread selection for defence.<sup id="cite_ref-FOOTNOTEWard-Smith_et_al.2020_3-1" class="reference"><a href="#cite_note-FOOTNOTEWard-Smith_et_al.2020-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> The spitting of venom displayed by some species of <a href="/wiki/Spitting_cobra" title="Spitting cobra">spitting cobra</a> is solely a defensive adaptation. A 2021 study showed that the venoms of all three lineages of spitting cobra convergently evolved higher levels of sensory neuron activation (i.e., cause more pain) than the venoms of non-spitting cobras, through the synergistic action of <a href="/wiki/Cytotoxins" class="mw-redirect" title="Cytotoxins">cytotoxins</a> and <a href="/wiki/Phospholipase_A2" title="Phospholipase A2">Phospholipase A2</a> toxins, indicating selection for a defensive function.<sup id="cite_ref-FOOTNOTEKazandjian_et_al.2021_57-0" class="reference"><a href="#cite_note-FOOTNOTEKazandjian_et_al.2021-57"><span class="cite-bracket">[</span>57<span class="cite-bracket">]</span></a></sup> </p><p>In contrast to venom composition and toxicity to specific lineages, venom yield, or the quantity of venom produced by an individual snake, has not been found to vary with the body-mass of prey animals, and instead to vary with the body-mass of snakes producing it. Yield increases with snake body-mass in a consistent with the hypothesis that snakes invest a constant proportion of metabolic output into producing venom, which is metabolically costly.<sup id="cite_ref-FOOTNOTEHealyCarboneJackson527–537_47-2" class="reference"><a href="#cite_note-FOOTNOTEHealyCarboneJackson527–537-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Functional_adaptations">Functional adaptations</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_snake_venom&action=edit&section=4" title="Edit section: Functional adaptations"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Mangrovennachtbaumnatter_(Boiga_dendrophila).JPG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/9e/Mangrovennachtbaumnatter_%28Boiga_dendrophila%29.JPG/220px-Mangrovennachtbaumnatter_%28Boiga_dendrophila%29.JPG" decoding="async" width="220" height="142" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/9e/Mangrovennachtbaumnatter_%28Boiga_dendrophila%29.JPG/330px-Mangrovennachtbaumnatter_%28Boiga_dendrophila%29.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/9e/Mangrovennachtbaumnatter_%28Boiga_dendrophila%29.JPG/440px-Mangrovennachtbaumnatter_%28Boiga_dendrophila%29.JPG 2x" data-file-width="1443" data-file-height="933" /></a><figcaption>The Mangrove snake <i>Boiga dendrophila</i>, whose venom is toxic to birds.</figcaption></figure> <p>Snakes use their venom to kill or subdue prey, as well as for other diet-related functions, such as digestion. Current scientific theory suggests that snake venom is not used for defense or for competition between members of the same species, unlike in other taxa. Thus adaptive evolution in snake venom has resulted in several adaptations with respect to these diet-related functions that increase the fitness of the snakes that carry them.<sup id="cite_ref-FOOTNOTECasewell_et_al.2013219–220_58-0" class="reference"><a href="#cite_note-FOOTNOTECasewell_et_al.2013219–220-58"><span class="cite-bracket">[</span>58<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTEBarlow_et_al.20092443_43-3" class="reference"><a href="#cite_note-FOOTNOTEBarlow_et_al.20092443-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTELi_et_al.2005_17-2" class="reference"><a href="#cite_note-FOOTNOTELi_et_al.2005-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> This is also reflected in variation in venom composition within a species; venom is known to vary geographically, and by age and size, likely reflecting variation in the prey consumed by different groups within a species.<sup id="cite_ref-FOOTNOTECasewellJacksonLaustsenSunagar2020570–581_13-4" class="reference"><a href="#cite_note-FOOTNOTECasewellJacksonLaustsenSunagar2020570–581-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> Geographic variation is also present at the species level; island snakes tend to have less complex venoms, while those living in highly productive habitats have more complex venoms, suggesting a <a href="/wiki/Biogeography" title="Biogeography">biogeographic pattern</a>.<sup id="cite_ref-FOOTNOTESiqueira‐Silva_et_al.20211978–1989_59-0" class="reference"><a href="#cite_note-FOOTNOTESiqueira‐Silva_et_al.20211978–1989-59"><span class="cite-bracket">[</span>59<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Prey-specific_venom_toxicity">Prey-specific venom toxicity</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_snake_venom&action=edit&section=5" title="Edit section: Prey-specific venom toxicity"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Echis_carinatus_sal.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/42/Echis_carinatus_sal.jpg/220px-Echis_carinatus_sal.jpg" decoding="async" width="220" height="180" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/42/Echis_carinatus_sal.jpg/330px-Echis_carinatus_sal.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/42/Echis_carinatus_sal.jpg/440px-Echis_carinatus_sal.jpg 2x" data-file-width="640" data-file-height="524" /></a><figcaption><i><a href="/wiki/Echis_carinatus" title="Echis carinatus">Echis carinatus</a></i>, one of the many species of the widespread genus <i>Echis</i>. The toxicity of <i>Echis</i> venom to scorpions has been found to vary with the proportion of arthropods in the snake's diet.</figcaption></figure> <p>Venom that is toxic only to a certain taxon, or strongly toxic only to a certain taxon, has been found in a number of snakes, suggesting that these venoms have evolved via natural selection to subdue preferred prey species. Examples of this phenomenon have been found in the Mangrove snake <i><a href="/wiki/Boiga_dendrophila" title="Boiga dendrophila">Boiga dendrophila</a></i>, which has a venom specifically toxic to birds,<sup id="cite_ref-FOOTNOTECasewell_et_al.2013220–221_48-1" class="reference"><a href="#cite_note-FOOTNOTECasewell_et_al.2013220–221-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> as well as in the genera <i>Echis</i> and <i>Sistrurus</i>, and in sea snakes. The venom of <a href="/wiki/Pseustes_sulphureus" class="mw-redirect" title="Pseustes sulphureus"><i>Spilotes sulphureus</i></a> which has two components, one of which is toxic to lizards but non-toxic in mammals, while the other is toxic in mammals and non-toxic in lizards.<sup id="cite_ref-FOOTNOTEModahl_et_al.2018_60-0" class="reference"><a href="#cite_note-FOOTNOTEModahl_et_al.2018-60"><span class="cite-bracket">[</span>60<span class="cite-bracket">]</span></a></sup> However, while several snakes possess venom that is highly toxic to their preferred prey species, the reverse correlation is not necessarily true: the venoms of several snakes are toxic to taxa that they do not consume in high proportions. Most snake venom, for instance, is highly toxic to lizards, although the proportion of lizard prey varies among snake species. This has led researchers to suggest that toxicity to a certain taxon is nearly independent of toxicity to another distantly related taxon.<sup id="cite_ref-FOOTNOTEGibbsMackessy2009_41-3" class="reference"><a href="#cite_note-FOOTNOTEGibbsMackessy2009-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> </p> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Hydrophis_-_Brehms.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/10/Hydrophis_-_Brehms.jpg/220px-Hydrophis_-_Brehms.jpg" decoding="async" width="220" height="212" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/10/Hydrophis_-_Brehms.jpg/330px-Hydrophis_-_Brehms.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/1/10/Hydrophis_-_Brehms.jpg 2x" data-file-width="415" data-file-height="400" /></a><figcaption><i><a href="/wiki/Hydrophis_cyanocinctus" title="Hydrophis cyanocinctus">Hydrophis cyanocinctus</a></i>, a member of a clade that has a greatly simplified venom that evolved in response to a diet of fish</figcaption></figure> <p>The natural diets of snakes in the widespread <a href="/wiki/Viperidae" class="mw-redirect" title="Viperidae">viper</a> genus <i><a href="/wiki/Echis" title="Echis">Echis</a></i> are highly varied, and include <a href="/wiki/Arthropods" class="mw-redirect" title="Arthropods">arthropods</a>, such as <a href="/wiki/Scorpion" title="Scorpion">scorpions</a>, as well as vertebrates. Various <i>Echis</i> species consume different quantities of arthropods in their diet.<sup id="cite_ref-FOOTNOTEBarlow_et_al.20092444,_2447_61-0" class="reference"><a href="#cite_note-FOOTNOTEBarlow_et_al.20092444,_2447-61"><span class="cite-bracket">[</span>61<span class="cite-bracket">]</span></a></sup> A 2009 study injected scorpions with the venom of various <i>Echis</i> species, and found a high correlation between the proportion of arthropods that the snakes consumed in their natural habitat, and the toxicity of their venom to scorpions.<sup id="cite_ref-FOOTNOTEBarlow_et_al.20092447_44-1" class="reference"><a href="#cite_note-FOOTNOTEBarlow_et_al.20092447-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> The researchers also found evidence that the evolution of venom more toxic to arthropods was related to an increase in the proportion of arthropods in the snakes' diet, and that diet and venom may have evolved by a process of <a href="/wiki/Coevolution" title="Coevolution">coevolution</a>. A phylogeny of the genus constructed using <a href="/wiki/Mitochondrial_DNA" title="Mitochondrial DNA">mitochondrial DNA</a> showed that one instance of a change in venom composition in the species ancestral to all <i>Echis</i> snakes was correlated with a shift to an arthropod based diet, whereas another shift in a more recent lineage was correlated with a shift to a diet of <a href="/wiki/Vertebrate" title="Vertebrate">vertebrates</a>.<sup id="cite_ref-FOOTNOTEBarlow_et_al.20092446–2448_62-0" class="reference"><a href="#cite_note-FOOTNOTEBarlow_et_al.20092446–2448-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup> Despite the higher toxicity of the venom of arthropod-consuming species, it was not found to incapacitate or kill prey any faster than that of species with fewer arthropods in their diet. Thus, the venom is thought to have evolved to minimize the volume required, as the production of venom carries a significant metabolic cost, thus providing a fitness benefit.<sup id="cite_ref-FOOTNOTEBarlow_et_al.20092446–2448_62-1" class="reference"><a href="#cite_note-FOOTNOTEBarlow_et_al.20092446–2448-62"><span class="cite-bracket">[</span>62<span class="cite-bracket">]</span></a></sup> This pattern is also found in other lineages.<sup id="cite_ref-FOOTNOTECasewell_et_al.2013223–225_63-0" class="reference"><a href="#cite_note-FOOTNOTECasewell_et_al.2013223–225-63"><span class="cite-bracket">[</span>63<span class="cite-bracket">]</span></a></sup> Similar results were obtained by a 2012 study which found that the venom of arthropod-consuming <i>Echis</i> species was more toxic to locusts than that of vertebrate-consuming species.<sup id="cite_ref-FOOTNOTERichards_et_al.2012_64-0" class="reference"><a href="#cite_note-FOOTNOTERichards_et_al.2012-64"><span class="cite-bracket">[</span>64<span class="cite-bracket">]</span></a></sup> </p><p>A 2009 study of the venom of four <i><a href="/wiki/Sistrurus" title="Sistrurus">Sistrurus</a></i> pit viper species found significant variation in the toxicity to mice.<sup id="cite_ref-FOOTNOTEGibbsMackessy2009_41-4" class="reference"><a href="#cite_note-FOOTNOTEGibbsMackessy2009-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> This variation was related to the proportion of small mammals in the diet of those species.<sup id="cite_ref-FOOTNOTEGibbsMackessy2009_41-5" class="reference"><a href="#cite_note-FOOTNOTEGibbsMackessy2009-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> The idea that <i>Sistrurus</i> venom had evolved to accommodate a mammal-based diet was supported by phylogenetic analysis. The researchers suggested that the basis for the difference in toxicity was the difference in muscle physiology in the various prey animals.<sup id="cite_ref-FOOTNOTEGibbsMackessy2009_41-6" class="reference"><a href="#cite_note-FOOTNOTEGibbsMackessy2009-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> Two lineages of <a href="/wiki/Elapid" class="mw-redirect" title="Elapid">elapid</a> snakes, common sea snakes and <i><a href="/wiki/Laticauda" class="mw-redirect" title="Laticauda">Laticauda</a></i> sea kraits, have independently colonized marine environments, and shifted to a very simple diet based on <a href="/wiki/Teleost" title="Teleost">teleosts</a>, or ray-finned fish.<sup id="cite_ref-FOOTNOTELi_et_al.2005_17-3" class="reference"><a href="#cite_note-FOOTNOTELi_et_al.2005-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> A 2005 study found that both these lineages have a much simpler set of venom proteins than their terrestrial relatives on the <a href="/wiki/Australia_(continent)" title="Australia (continent)">Australian continent</a>, which have a more varied and complex diet.<sup id="cite_ref-FOOTNOTELi_et_al.2005_17-4" class="reference"><a href="#cite_note-FOOTNOTELi_et_al.2005-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> These findings were confirmed by a 2012 study, which compared the venoms of <i><a href="/wiki/Toxicocalamus" title="Toxicocalamus">Toxicocalamus</a> longissimus</i>, a terrestrial species, and <i><a href="/wiki/Hydrophis_cyanocinctus" title="Hydrophis cyanocinctus">Hydrophis cyanocinctus</a></i>, a marine species, both within the subfamily Hydrophiinae (which is also within the Elapid family). Despite being closely related to one another, the marine species had a significantly simpler set of venom proteins.<sup id="cite_ref-FOOTNOTECalvete_et_al.20124094–4098_16-1" class="reference"><a href="#cite_note-FOOTNOTECalvete_et_al.20124094–4098-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> The venoms of the sea snakes are nonetheless among the most toxic venoms known. It has been argued that since sea snakes are typically unable to prevent the escape of bitten prey, their venoms have evolved to act very rapidly.<sup id="cite_ref-FOOTNOTECalvete_et_al.20124092–4093_65-0" class="reference"><a href="#cite_note-FOOTNOTECalvete_et_al.20124092–4093-65"><span class="cite-bracket">[</span>65<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Pre-digestion_of_prey">Pre-digestion of prey</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_snake_venom&action=edit&section=6" title="Edit section: Pre-digestion of prey"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1237032888/mw-parser-output/.tmulti">.mw-parser-output .tmulti .multiimageinner{display:flex;flex-direction:column}.mw-parser-output .tmulti .trow{display:flex;flex-direction:row;clear:left;flex-wrap:wrap;width:100%;box-sizing:border-box}.mw-parser-output .tmulti .tsingle{margin:1px;float:left}.mw-parser-output .tmulti .theader{clear:both;font-weight:bold;text-align:center;align-self:center;background-color:transparent;width:100%}.mw-parser-output .tmulti .thumbcaption{background-color:transparent}.mw-parser-output .tmulti .text-align-left{text-align:left}.mw-parser-output .tmulti .text-align-right{text-align:right}.mw-parser-output .tmulti .text-align-center{text-align:center}@media all and (max-width:720px){.mw-parser-output .tmulti .thumbinner{width:100%!important;box-sizing:border-box;max-width:none!important;align-items:center}.mw-parser-output .tmulti .trow{justify-content:center}.mw-parser-output .tmulti .tsingle{float:none!important;max-width:100%!important;box-sizing:border-box;text-align:center}.mw-parser-output .tmulti .tsingle .thumbcaption{text-align:left}.mw-parser-output .tmulti .trow>.thumbcaption{text-align:center}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .tmulti .multiimageinner img{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .tmulti .multiimageinner img{background-color:white}}</style><div class="thumb tmulti tright"><div class="thumbinner multiimageinner" style="width:312px;max-width:312px"><div class="trow"><div class="tsingle" style="width:162px;max-width:162px"><div class="thumbimage"><span typeof="mw:File"><a href="/wiki/File:PrairieRattlesnake.jpg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/6/60/PrairieRattlesnake.jpg/160px-PrairieRattlesnake.jpg" decoding="async" width="160" height="108" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/60/PrairieRattlesnake.jpg/240px-PrairieRattlesnake.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/60/PrairieRattlesnake.jpg/320px-PrairieRattlesnake.jpg 2x" data-file-width="3030" data-file-height="2045" /></a></span></div></div><div class="tsingle" style="width:146px;max-width:146px"><div class="thumbimage"><span typeof="mw:File"><a href="/wiki/File:PDB_1rm8_EBI.jpg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/7/75/PDB_1rm8_EBI.jpg/144px-PDB_1rm8_EBI.jpg" decoding="async" width="144" height="108" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/75/PDB_1rm8_EBI.jpg/216px-PDB_1rm8_EBI.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/75/PDB_1rm8_EBI.jpg/288px-PDB_1rm8_EBI.jpg 2x" data-file-width="800" data-file-height="600" /></a></span></div></div></div><div class="trow" style="display:flex"><div class="thumbcaption">The venom of the prairie rattlesnake, <i><a href="/wiki/Crotalus_viridis" title="Crotalus viridis">Crotalus viridis</a></i> (left) includes <a href="/wiki/Metalloproteinase" title="Metalloproteinase">metalloproteinases</a> (example on the right) which help digest the prey before the snake eats it.</div></div></div></div> <p>The various subspecies of the rattlesnake genus <i><a href="/wiki/Crotalus" title="Crotalus">Crotalus</a></i>, produce venoms that carry out two conflicting functions. The venom immobilizes prey after a bite, and also helps digestion by breaking down tissues before the snake eats its prey.<sup id="cite_ref-FOOTNOTEMackessy2010_18-1" class="reference"><a href="#cite_note-FOOTNOTEMackessy2010-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> As with other members of the family <a href="/wiki/Viperidae" class="mw-redirect" title="Viperidae">Viperidae</a>, the venoms of <i>Crotalus</i> disrupt the <a href="/wiki/Homeostasis" title="Homeostasis">homeostatic processes</a> of prey animals. However, there is a wide variety of venom compositions among the species of <i>Crotalus</i>. A 2010 study found a 100-fold difference in the amount of <a href="/wiki/Metalloproteinase" title="Metalloproteinase">metalloproteinase</a> activity among the various snakes, with <i><a href="/wiki/Crotalus_cerberus" title="Crotalus cerberus">Crotalus cerberus</a></i> having the highest activity and <i><a href="/wiki/Crotalus_oreganus_concolor" class="mw-redirect" title="Crotalus oreganus concolor">Crotalus oreganus concolor</a></i> having the lowest. There was also a 15-fold variation in the amount of protease activity, with <i>C. o. concolor</i> and <i>C. cerberus</i> having the highest and lowest activities, respectively.<sup id="cite_ref-FOOTNOTEMackessy2010_18-2" class="reference"><a href="#cite_note-FOOTNOTEMackessy2010-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> </p><p>Metalloproteinase activity causes <a href="/wiki/Hemorrhage" class="mw-redirect" title="Hemorrhage">hemorrhage</a> and <a href="/wiki/Necrosis" title="Necrosis">necrosis</a> following a snake bite, a process which aids digestion. The activity of proteases, on the other hand, disrupts <a href="/wiki/Platelet" title="Platelet">platelet</a> and muscle function and damages <a href="/wiki/Cell_membrane" title="Cell membrane">cell membranes</a>, and thus contributes to a quick death for the prey animal.<sup id="cite_ref-FOOTNOTEMackessy2010_18-3" class="reference"><a href="#cite_note-FOOTNOTEMackessy2010-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> The study found that the venoms of <i>Crotalus</i> fell into two categories; those that favored metalloproteinases (Type I) and those that favored proteases (Type II). The study stated that these functions were essentially mutually exclusive; venoms had been selected for based on either their toxicity or their tenderizing potential. The researchers also hypothesized that the reason for this dichotomy was that a venom high in <a href="/wiki/Neurotoxin" title="Neurotoxin">neurotoxicity</a>, such as a type II venom, kills an animal quickly, preventing the relatively slower acting metalloproteinase from digesting tissue.<sup id="cite_ref-FOOTNOTEMackessy2010_18-4" class="reference"><a href="#cite_note-FOOTNOTEMackessy2010-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Tracking_bitten_prey">Tracking bitten prey</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_snake_venom&action=edit&section=7" title="Edit section: Tracking bitten prey"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1237032888/mw-parser-output/.tmulti"><div class="thumb tmulti tright"><div class="thumbinner multiimageinner" style="width:306px;max-width:306px"><div class="trow"><div class="tsingle" style="width:160px;max-width:160px"><div class="thumbimage"><span typeof="mw:File"><a href="/wiki/File:Crotalus_atrox_diamantklapperschlange_kopf.jpg" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/9/99/Crotalus_atrox_diamantklapperschlange_kopf.jpg/158px-Crotalus_atrox_diamantklapperschlange_kopf.jpg" decoding="async" width="158" height="105" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/99/Crotalus_atrox_diamantklapperschlange_kopf.jpg/237px-Crotalus_atrox_diamantklapperschlange_kopf.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/99/Crotalus_atrox_diamantklapperschlange_kopf.jpg/316px-Crotalus_atrox_diamantklapperschlange_kopf.jpg 2x" data-file-width="3000" data-file-height="2000" /></a></span></div></div><div class="tsingle" style="width:142px;max-width:142px"><div class="thumbimage"><span typeof="mw:File"><a href="/wiki/File:Desintegrin_heterodimer.png" class="mw-file-description"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/1/11/Desintegrin_heterodimer.png/140px-Desintegrin_heterodimer.png" decoding="async" width="140" height="105" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/11/Desintegrin_heterodimer.png/210px-Desintegrin_heterodimer.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/11/Desintegrin_heterodimer.png/280px-Desintegrin_heterodimer.png 2x" data-file-width="640" data-file-height="480" /></a></span></div></div></div><div class="trow" style="display:flex"><div class="thumbcaption">The western diamondback rattlesnake, <i><a href="/wiki/Crotalus_atrox" class="mw-redirect" title="Crotalus atrox">Crotalus atrox</a></i> (left), whose venom contains <a href="/wiki/Disintegrin" title="Disintegrin">disintegrins</a> (right) which allow it to track bitten prey</div></div></div></div> <p>Another example of an adaptive function other than prey immobilization is the role of <a href="/wiki/Viperidae" class="mw-redirect" title="Viperidae">viperid</a> venom in allowing the snake to track a prey animal it has bitten, a process known as "prey relocalization." This important adaptation allowed rattlesnakes to evolve the strike-and-release bite mechanism, which provided a huge benefit to snakes by minimizing contact with potentially dangerous prey animals.<sup id="cite_ref-FOOTNOTESaviola_et_al.2013_19-1" class="reference"><a href="#cite_note-FOOTNOTESaviola_et_al.2013-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> However, this adaptation then requires the snake to track down the bitten animal in order to eat it, in an environment full of other animals of the same species. A 2013 study found that western diamondback rattlesnakes (<i><a href="/wiki/Crotalus_atrox" class="mw-redirect" title="Crotalus atrox">Crotalus atrox</a></i>) responded more actively to mouse carcasses that had been injected with crude rattlesnake venom. When the various components of the venom were separated out, the snakes responded to mice injected with two kinds of <a href="/wiki/Disintegrin" title="Disintegrin">disintegrins</a>. The study concluded that these disintegrin proteins were responsible for allowing the snakes to track their prey, by changing the odor of the bitten animal.<sup id="cite_ref-FOOTNOTESaviola_et_al.2013_19-2" class="reference"><a href="#cite_note-FOOTNOTESaviola_et_al.2013-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Diet-based_atrophication">Diet-based atrophication</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_snake_venom&action=edit&section=8" title="Edit section: Diet-based atrophication"><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:Eierschlange_frisst_Zwergwachtelei.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/26/Eierschlange_frisst_Zwergwachtelei.jpg/170px-Eierschlange_frisst_Zwergwachtelei.jpg" decoding="async" width="170" height="234" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/26/Eierschlange_frisst_Zwergwachtelei.jpg/255px-Eierschlange_frisst_Zwergwachtelei.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/26/Eierschlange_frisst_Zwergwachtelei.jpg/340px-Eierschlange_frisst_Zwergwachtelei.jpg 2x" data-file-width="1024" data-file-height="1411" /></a><figcaption>A shift to a diet of eggs has resulted in an atrophied venom system in the common egg-eater <i>Dasypeltis scabra</i></figcaption></figure> <p>Venom in a number of lineages of snakes is thought to have atrophied in response to dietary shifts.<sup id="cite_ref-FOOTNOTEFry_et_al.2012a434–436_7-5" class="reference"><a href="#cite_note-FOOTNOTEFry_et_al.2012a434–436-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> A 2005 study in the marbled sea snake, <i><a href="/wiki/Aipysurus_eydouxii" title="Aipysurus eydouxii">Aipysurus eydouxii</a></i> found that the gene for a <a href="/wiki/Calciseptine#Three-fingered_toxin_family" title="Calciseptine">three-fingered protein</a> found in the venom had undergone a deletion of two <a href="/wiki/Nucleobase" class="mw-redirect" title="Nucleobase">nucleotide bases</a> which made the venom 50–100 times less toxic than it had been previously. This change was correlated with a change in diet from fish to a diet consisting almost entirely of fish eggs, suggesting that the adaptation to an egg diet had removed the <a href="/wiki/Selection_pressure" class="mw-redirect" title="Selection pressure">selection pressure</a> needed to maintain a highly toxic venom, allowing the venom genes to accumulate deleterious mutations.<sup id="cite_ref-FOOTNOTELi_et_al.2005_17-5" class="reference"><a href="#cite_note-FOOTNOTELi_et_al.2005-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-FOOTNOTECalvete_et_al.20124097–4098_66-0" class="reference"><a href="#cite_note-FOOTNOTECalvete_et_al.20124097–4098-66"><span class="cite-bracket">[</span>66<span class="cite-bracket">]</span></a></sup> A similar venom degradation following a shift to an egg-based diet has been found in the Common egg-eater <i><a href="/wiki/Dasypeltis_scabra" title="Dasypeltis scabra">Dasypeltis scabra</a></i>, whose diet consists entirely of birds' eggs, meaning that the snake had no use for its venom. This has led biologists to suggest that if venom is not used by a species, it is rapidly lost.<sup id="cite_ref-FOOTNOTEFry_et_al.2008_67-0" class="reference"><a href="#cite_note-FOOTNOTEFry_et_al.2008-67"><span class="cite-bracket">[</span>67<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_snake_venom&action=edit&section=9" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Citations">Citations</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_snake_venom&action=edit&section=10" title="Edit section: Citations"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width" style="column-width: 22em;"> <ol class="references"> <li id="cite_note-FOOTNOTEHargreaves_et_al._(a)2014-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-FOOTNOTEHargreaves_et_al._(a)2014_1-0">^</a></b></span> <span class="reference-text"><a href="#CITEREFHargreaves_et_al._(a)2014">Hargreaves et al. 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class="reference-text"><a href="#CITEREFFry_et_al.2008">Fry et al. 2008</a>.</span> </li> </ol></div> <div class="mw-heading mw-heading3"><h3 id="Cited_sources">Cited sources</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Evolution_of_snake_venom&action=edit&section=11" title="Edit section: Cited sources"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239549316">.mw-parser-output .refbegin{margin-bottom:0.5em}.mw-parser-output .refbegin-hanging-indents>ul{margin-left:0}.mw-parser-output .refbegin-hanging-indents>ul>li{margin-left:0;padding-left:3.2em;text-indent:-3.2em}.mw-parser-output .refbegin-hanging-indents ul,.mw-parser-output .refbegin-hanging-indents ul li{list-style:none}@media(max-width:720px){.mw-parser-output .refbegin-hanging-indents>ul>li{padding-left:1.6em;text-indent:-1.6em}}.mw-parser-output .refbegin-columns{margin-top:0.3em}.mw-parser-output .refbegin-columns ul{margin-top:0}.mw-parser-output .refbegin-columns li{page-break-inside:avoid;break-inside:avoid-column}@media screen{.mw-parser-output .refbegin{font-size:90%}}</style><div class="refbegin refbegin-columns references-column-width" style="column-width: 30em"> <ul><li><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFAlmeidaVialaNachtigallBroe2021" class="citation journal cs1">Almeida, Diego Dantas; Viala, Vincent Louis; Nachtigall, Pedro Gabriel; Broe, Michael; Gibbs, H. 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title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proceedings+of+the+Royal+Society+B%3A+Biological+Sciences&rft.atitle=When+one+phenotype+is+not+enough%3A+divergent+evolutionary+trajectories+govern+venom+variation+in+a+widespread+rattlesnake+species&rft.volume=286&rft.issue=1898&rft.pages=20182735&rft.date=2019-03-13&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC6458317%23id-name%3DPMC&rft_id=info%3Apmid%2F30862287&rft_id=info%3Adoi%2F10.1098%2Frspb.2018.2735&rft.aulast=Zancolli&rft.aufirst=Giulia&rft.au=Calvete%2C+Juan+J.&rft.au=Cardwell%2C+Michael+D.&rft.au=Greene%2C+Harry+W.&rft.au=Hayes%2C+William+K.&rft.au=Hegarty%2C+Matthew+J.&rft.au=Herrmann%2C+Hans-Werner&rft.au=Holycross%2C+Andrew+T.&rft.au=Lannutti%2C+Dominic+I.&rft.au=Mulley%2C+John+F.&rft.au=Sanz%2C+Libia&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC6458317&rfr_id=info%3Asid%2Fen.wikipedia.org%3AEvolution+of+snake+venom" 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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" 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evolution">Parallel evolution</a></li> <li><a href="/wiki/Signalling_theory" title="Signalling theory">Signalling theory</a> <ul><li><a href="/wiki/Handicap_principle" title="Handicap principle">Handicap principle</a></li></ul></li> <li><a href="/wiki/Speciation" title="Speciation">Speciation</a> <ul><li><a href="/wiki/Species" title="Species">Species</a></li> <li><a href="/wiki/Species_complex" title="Species complex">Species complex</a></li></ul></li> <li><a href="/wiki/Evolutionary_taxonomy" title="Evolutionary taxonomy">Taxonomy</a></li> <li><a href="/wiki/Unit_of_selection" title="Unit of selection">Unit of selection</a> <ul><li><a href="/wiki/Gene-centered_view_of_evolution" title="Gene-centered view of evolution">Gene-centered view of evolution</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Population_genetics" title="Population genetics">Population<br />genetics</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Selective_breeding" title="Selective breeding">Artificial selection</a></li> <li><a href="/wiki/Biodiversity" title="Biodiversity">Biodiversity</a></li> <li><a href="/wiki/Evolutionarily_stable_strategy" title="Evolutionarily stable strategy">Evolutionarily stable strategy</a></li> <li><a href="/wiki/Fisher%27s_principle" title="Fisher's principle">Fisher's principle</a></li> <li><a href="/wiki/Fitness_(biology)" title="Fitness (biology)">Fitness</a> <ul><li><a href="/wiki/Inclusive_fitness" title="Inclusive fitness">Inclusive</a></li></ul></li> <li><a href="/wiki/Gene_flow" title="Gene flow">Gene flow</a></li> <li><a href="/wiki/Genetic_drift" title="Genetic drift">Genetic drift</a></li> <li><a href="/wiki/Kin_selection" title="Kin selection">Kin selection</a> <ul><li><a href="/wiki/Parental_investment" title="Parental investment">Parental investment</a></li> <li><a href="/wiki/Parent%E2%80%93offspring_conflict" title="Parent–offspring conflict">Parent–offspring conflict</a></li></ul></li> <li><a href="/wiki/Mutation" title="Mutation">Mutation</a></li> <li><a href="/wiki/Population" title="Population">Population</a></li> <li><a href="/wiki/Natural_selection" title="Natural selection">Natural selection</a></li> <li><a href="/wiki/Sexual_dimorphism" title="Sexual dimorphism">Sexual dimorphism</a></li> <li><a href="/wiki/Sexual_selection" title="Sexual selection">Sexual selection</a> <ul><li><a href="/wiki/Sexual_selection_in_flowering_plants" title="Sexual selection in flowering plants">Flowering plants</a></li> <li><a href="/wiki/Sexual_selection_in_fungi" title="Sexual selection in fungi">Fungi</a></li> <li><a href="/wiki/Mate_choice" title="Mate choice">Mate choice</a></li></ul></li> <li><a href="/wiki/Social_selection" title="Social selection">Social selection</a></li> <li><a href="/wiki/Trivers%E2%80%93Willard_hypothesis" title="Trivers–Willard hypothesis">Trivers–Willard hypothesis</a></li> <li><a href="/wiki/Genetic_variation" title="Genetic variation">Variation</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Development</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Canalisation_(genetics)" title="Canalisation (genetics)">Canalisation</a></li> <li><a href="/wiki/Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evolutionary developmental biology</a></li> <li><a href="/wiki/Genetic_assimilation" title="Genetic assimilation">Genetic assimilation</a></li> <li><a href="/wiki/Inversion_(evolutionary_biology)" title="Inversion (evolutionary biology)">Inversion</a></li> <li><a href="/wiki/Modularity_(biology)" title="Modularity (biology)">Modularity</a></li> <li><a href="/wiki/Phenotypic_plasticity" title="Phenotypic plasticity">Phenotypic plasticity</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Taxon" title="Taxon">taxa</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_bacteria" title="Evolution of bacteria">Bacteria</a></li> <li><a href="/wiki/Evolution_of_birds" title="Evolution of birds">Birds</a> <ul><li><a href="/wiki/Origin_of_birds" title="Origin of birds">origin</a></li></ul></li> <li><a href="/wiki/Evolution_of_brachiopods" title="Evolution of brachiopods">Brachiopods</a></li> <li><a href="/wiki/Evolution_of_molluscs" title="Evolution of molluscs">Molluscs</a> <ul><li><a href="/wiki/Evolution_of_cephalopods" title="Evolution of cephalopods">Cephalopods</a></li></ul></li> <li><a href="/wiki/Dinosaur#Evolutionary_history" title="Dinosaur">Dinosaurs</a></li> <li><a href="/wiki/Evolution_of_fish" title="Evolution of fish">Fish</a></li> <li><a href="/wiki/Evolution_of_fungi" title="Evolution of fungi">Fungi</a></li> <li><a href="/wiki/Evolution_of_insects" title="Evolution of insects">Insects</a> <ul><li><a href="/wiki/Evolution_of_butterflies" title="Evolution of butterflies">butterflies</a></li></ul></li> <li><a href="/wiki/History_of_life" title="History of life">Life</a></li> <li><a href="/wiki/Evolution_of_mammals" title="Evolution of mammals">Mammals</a> <ul><li><a href="/wiki/Cat_gap" title="Cat gap">cats</a></li> <li><a href="/wiki/Canidae#Evolution" title="Canidae">canids</a> <ul><li><a href="/wiki/Evolution_of_the_wolf" title="Evolution of the wolf">wolves</a></li> <li><a href="/wiki/Domestication_of_the_dog" title="Domestication of the dog">dogs</a></li></ul></li> <li><a href="/wiki/Hyena#Evolution" title="Hyena">hyenas</a></li> <li><a href="/wiki/Evolution_of_cetaceans" title="Evolution of cetaceans">dolphins and whales</a></li> <li><a href="/wiki/Evolution_of_the_horse" title="Evolution of the horse">horses</a></li> <li><a href="/wiki/Evolution_of_Macropodidae" title="Evolution of Macropodidae">Kangaroos</a></li> <li><a href="/wiki/Evolution_of_primates" title="Evolution of primates">primates</a> <ul><li><a href="/wiki/Human_evolution" title="Human evolution">humans</a></li> <li><a href="/wiki/Evolution_of_lemurs" title="Evolution of lemurs">lemurs</a></li></ul></li> <li><a href="/wiki/Evolution_of_sirenians" title="Evolution of sirenians">sea cows</a></li></ul></li> <li><a href="/wiki/Evolutionary_history_of_plants" title="Evolutionary history of plants">Plants</a> <ul><li><a href="/wiki/Pollinator-mediated_selection" title="Pollinator-mediated selection">pollinator-mediated</a></li></ul></li> <li><a href="/wiki/Evolution_of_reptiles" title="Evolution of reptiles">Reptiles</a></li> <li><a href="/wiki/Evolution_of_spiders" title="Evolution of spiders">Spiders</a></li> <li><a href="/wiki/Evolution_of_tetrapods" title="Evolution of tetrapods">Tetrapods</a></li> <li><a href="/wiki/Viral_evolution" title="Viral evolution">Viruses</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Organ_(anatomy)" class="mw-redirect" title="Organ (anatomy)">organs</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_cells" title="Evolution of cells">Cell</a></li> <li><a href="/wiki/Models_of_DNA_evolution" title="Models of DNA evolution">DNA</a></li> <li><a href="/wiki/Evolution_of_flagella" title="Evolution of flagella">Flagella</a></li> <li><a href="/wiki/Eukaryote#Origin_of_eukaryotes" title="Eukaryote">Eukaryotes</a> <ul><li><a href="/wiki/Symbiogenesis" title="Symbiogenesis">symbiogenesis</a></li> <li><a href="/wiki/Chromosome#Eukaryotes" title="Chromosome">chromosome</a></li> <li><a href="/wiki/Endomembrane_system#Evolution" title="Endomembrane system">endomembrane system</a></li> <li><a href="/wiki/Mitochondrion#Origin_and_evolution" title="Mitochondrion">mitochondria</a></li> <li><a href="/wiki/Cell_nucleus#Evolution" title="Cell nucleus">nucleus</a></li> <li><a href="/wiki/Plastid#Origin" title="Plastid">plastids</a></li></ul></li> <li>In animals <ul><li><a href="/wiki/Evolution_of_the_eye" title="Evolution of the eye">eye</a></li> <li><a href="/wiki/Hair#Evolution" title="Hair">hair</a></li> <li><a href="/wiki/Evolution_of_mammalian_auditory_ossicles" title="Evolution of mammalian auditory ossicles">auditory ossicle</a></li> <li><a href="/wiki/Evolution_of_nervous_systems" title="Evolution of nervous systems">nervous system</a></li> <li><a href="/wiki/Evolution_of_the_brain" title="Evolution of the brain">brain</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Of <a href="/wiki/Biological_process" title="Biological process">processes</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Evolution_of_ageing" title="Evolution of ageing">Aging</a> <ul><li><a href="/wiki/Death#Evolution_of_aging_and_mortality" title="Death">Death</a></li> <li><a href="/wiki/Programmed_cell_death#Evolutionary_origin_of_mitochondrial_apoptosis" title="Programmed cell death">Programmed cell death</a></li></ul></li> <li><a href="/wiki/Origin_of_avian_flight" title="Origin of avian flight">Avian flight</a></li> <li><a href="/wiki/Evolution_of_biological_complexity" title="Evolution of biological complexity">Biological complexity</a></li> <li><a href="/wiki/Cooperation_(evolution)" title="Cooperation (evolution)">Cooperation</a></li> <li><a href="/wiki/Evolution_of_color_vision" title="Evolution of color vision">Color vision</a> <ul><li><a href="/wiki/Evolution_of_color_vision_in_primates" title="Evolution of color vision in primates">in primates</a></li></ul></li> <li><a href="/wiki/Evolution_of_emotion" title="Evolution of emotion">Emotion</a></li> <li><a href="/wiki/Empathy#Evolution_across_species" title="Empathy">Empathy</a></li> <li><a href="/wiki/Evolutionary_ethics" title="Evolutionary ethics">Ethics</a></li> <li><a href="/wiki/Evolution_of_eusociality" title="Evolution of eusociality">Eusociality</a></li> <li><a href="/wiki/Immune_system#Evolution_and_other_mechanisms" title="Immune system">Immune system</a></li> <li><a href="/wiki/Metabolism#Evolution" title="Metabolism">Metabolism</a></li> <li><a href="/wiki/Monogamy_in_animals" title="Monogamy in animals">Monogamy</a></li> <li><a href="/wiki/Evolution_of_morality" title="Evolution of morality">Morality</a></li> <li><a href="/wiki/Mosaic_evolution" title="Mosaic evolution">Mosaic evolution</a></li> <li><a href="/wiki/Multicellular_organism#Evolutionary_history" title="Multicellular organism">Multicellularity</a></li> <li><a href="/wiki/Evolution_of_sexual_reproduction" title="Evolution of sexual reproduction">Sexual reproduction</a> <ul><li><a href="/wiki/Anisogamy#Evolution" title="Anisogamy">Gamete differentiation/sexes</a></li> <li><a href="/wiki/Biological_life_cycle" title="Biological life cycle">Life cycles/nuclear phases</a></li> <li><a href="/wiki/Mating_type" title="Mating type">Mating types</a></li> <li><a href="/wiki/Origin_and_function_of_meiosis" title="Origin and function of meiosis">Meiosis</a></li> <li><a href="/wiki/Sex-determination_system#Evolution" title="Sex-determination system">Sex-determination</a></li></ul></li> <li><a class="mw-selflink selflink">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="" 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