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<title>Bat wing development</title>
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<span id="openzim-page-title" class="mw-page-title-main"><span class="mw-page-title-main">Bat wing development</span></span>
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<p>The order <a href="Chiroptera" class="mw-redirect" title="Chiroptera">Chiroptera</a>, comprising all bats, has evolved the unique mammalian adaptation of <a href="Bat_flight" title="Bat flight">flight</a>. Bat wings are modified <a href="Tetrapod" title="Tetrapod">tetrapod</a> forelimbs. Because bats are <a href="Mammals" class="mw-redirect" title="Mammals">mammals</a>, the <a href="Skeletal" class="mw-redirect" title="Skeletal">skeletal</a> structures in their wings are morphologically <a href="Homology_(biology)" title="Homology (biology)">homologous</a> to the skeletal components found in other tetrapod forelimbs. Through adaptive evolution these structures in bats have undergone many morphological changes, such as webbed digits, elongation of the forelimb, and reduction in bone thickness.<sup id="cite_ref-review_1-0" class="reference"><a href="#cite_note-review-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Recently, there have been comparative studies of mouse and bat forelimb development to understand the genetic basis of morphological evolution. Consequently, the bat wing is a valuable <a href="Evolutionary_developmental_biology" title="Evolutionary developmental biology">evo-devo</a> model for studying the evolution of vertebrate limb diversity.
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<div class="mw-heading mw-heading2"><h2 id="Comparisons_to_mouse_limb_development">Comparisons to mouse limb development</h2></div>
<p>Tetrapod <a href="Limb_development" title="Limb development">limb development</a> involves many signaling molecules such as <a href="Fibroblast_growth_factor" title="Fibroblast growth factor">FGF</a>, <a href="Bone_morphogenetic_protein" title="Bone morphogenetic protein">BMP</a>, <a href="Sonic_hedgehog" class="mw-redirect" title="Sonic hedgehog">SHH</a> and <a href="Wnt_signaling_pathway" title="Wnt signaling pathway">WNT</a>. The <a href="Apical_ectodermal_ridge" title="Apical ectodermal ridge">apical ectodermal ridge</a> is a structure found at the distal most tip which becomes a key signaling center for the developing limb.<sup id="cite_ref-aer_2-0" class="reference"><a href="#cite_note-aer-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Surprisingly many of the same signaling pathways known to play a role in tetrapod limb development have been found to play a role in bat forelimb development but the timing, intensity, and spatial <a href="Gene_expression" title="Gene expression">gene expression</a> of some <a href="Ortholog" class="mw-redirect" title="Ortholog">orthologous</a> genes have changed. Since mice are also mammals, it is convenient to compare morphology and development of forelimbs between mice and bats; these comparisons may elucidate the genetic basis of adaptive bat wing development.
</p><p>Although many of the molecular mechanisms involved in limb development are conserved between mouse and bat, there are a number of differences primarily seen in gene expression patterns. Surprisingly, the coding regions of many of these genes with different expression domains are highly conserved between mouse and bat. Thus, it is likely that this major morphological transition was a consequence of <a href="Cis-acting" class="mw-redirect" title="Cis-acting">cis</a>-<a href="Gene_regulation" class="mw-redirect" title="Gene regulation">regulatory</a> changes. Researchers can study the genetic basis of bat wing development by using comparative <a href="In_situ_hybridization" title="In situ hybridization">in situ hybridization</a> to examine gene expression domains and using experimental embryology in mice and bats.
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<div class="mw-heading mw-heading2"><h2 id="Presence_of_webbed_digits">Presence of webbed digits</h2></div>
<p>Formation of the bat wing membrane (the <a href="Patagium" title="Patagium">patagium</a>) allowed a greater surface area of the wing necessary for flight. All vertebrate limb formation initially has tissue between the digits after which <a href="Apoptosis" title="Apoptosis">apoptosis</a> occurs to separate the digits. BMP signals are most likely responsible for the interdigit apoptosis as they are expressed in the interdigit tissue and blocking BMP signaling will prevent interdigital <a href="Apoptosis" title="Apoptosis">apoptosis</a>.<sup id="cite_ref-bmp_3-0" class="reference"><a href="#cite_note-bmp-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> However, in bats, BMP genes are still expressed in the interdigits and yet interdigit apoptosis is repressed. FGF signaling has been associated with blocking cell death.<sup id="cite_ref-fibroblast_4-0" class="reference"><a href="#cite_note-fibroblast-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> <i><a href="Fibroblast_growth_factor" title="Fibroblast growth factor">Fgf8</a></i> is expressed in bat interdigit tissue during a time when apoptosis occurs which does not occur in mice. Thus, FGFs may play a role in blocking the apoptotic effects of BMPs in the bat wing interdigit. Finally, applying ectopic BMPs and FGF antagonists to developing bat wings results in apoptosis of the patagium.<sup id="cite_ref-webbing_5-0" class="reference"><a href="#cite_note-webbing-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup>
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<div class="mw-heading mw-heading2"><h2 id="Elongation_of_forelimb">Elongation of forelimb</h2></div>
<p>One major difference in bat forearms is that their skeletal limb structures are elongated. This elongation of the forelimb skeleton is required to support the wing membrane. Comparative in situ hybridization studies have revealed that the expression domain of <i>fgf8</i> in bat forelimb AER are expanded in comparison to the mouse forelimb, suggesting that expanded expression of <i>fgf8</i> may contribute to the larger size of the bat forelimb. Because the mouse and bat orthologs are conserved, there is likely to be a regulatory change in <i>fgf8</i>.<sup id="cite_ref-fgf_6-0" class="reference"><a href="#cite_note-fgf-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> In mice, one gene known to regulate limb growth is <i><a href="PRX_(gene)" title="PRX (gene)">prx1</a></i>, which encodes a transcription factor.<sup id="cite_ref-HOX_7-0" class="reference"><a href="#cite_note-HOX-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> The expression patterns of <i>prx1</i> in bats differs from mice in that <i>prx1</i> has an expanded expression domain and is upregulated.
Researchers found that the coding region of <i>prx1</i> in bats is nearly identical to mice but found a bat-specific <i>prx1</i> <a href="Enhancer_(genetics)" title="Enhancer (genetics)">enhancer</a>.<sup id="cite_ref-length_8-0" class="reference"><a href="#cite_note-length-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> When they replaced the bat <i>prx1</i> enhancer with the endogenous enhancer found in mice, these transgenic mice had slightly increased forelimbs. Comparative studies have established that bat digits undergo a more rapid rate of <a href="Chondrocyte" title="Chondrocyte">chondrocyte</a> proliferation.<sup id="cite_ref-flight_9-0" class="reference"><a href="#cite_note-flight-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> In addition to interdigit apoptosis, BMPs have been shown to affect chondrocyte proliferation and digit length in mice.<sup id="cite_ref-plate_10-0" class="reference"><a href="#cite_note-plate-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> <i><a href="Bone_morphogenetic_protein_2" title="Bone morphogenetic protein 2">Bmp-2</a></i> shows increased expression in the digits of bats compared with mice, suggesting that a change in the BMP pathway has occurred to give rise to longer bat digits.<sup id="cite_ref-flight_9-1" class="reference"><a href="#cite_note-flight-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup>
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<div class="mw-heading mw-heading2"><h2 id="Reduction_in_bone_thickness">Reduction in bone thickness</h2></div>
<p>Another major difference in bat forelimbs is in the density of their skeletal limbs. The bones found in their forelimbs are reduced to achieve a light body weight required for flight. In particular, their <a href="Ulna" title="Ulna">ulna</a> is reduced in width and fused to the other zeugopod element, the radius.<sup id="cite_ref-review_1-1" class="reference"><a href="#cite_note-review-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> One of the possible molecular pathways involved in reduction of bat skeletal forelimb thickness is differences in <a href="Sonic_hedgehog" class="mw-redirect" title="Sonic hedgehog">SHH</a> expression. Mice with <i>shh</i> null mutations lose their ulna structure.<sup id="cite_ref-shh_11-0" class="reference"><a href="#cite_note-shh-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Another good candidate for bat bone reduction is <i><a href="HOXD13" title="HOXD13">Hox-d13</a></i>, a gene belonging to the <a href="Hox_gene" title="Hox gene">Hox gene</a> family. In situ hybridization studies have found that the <i>Hoxd13</i> expression domain in bat limbs has been shifted posteriorly in comparison to mouse.<sup id="cite_ref-hoxy_12-0" class="reference"><a href="#cite_note-hoxy-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> This observed difference in the expression pattern of <i>Hoxd13</i> may also explain reduced size and density of the ulna found in bats. Overall, these studies suggest that the molecular changes responsible for the evolution of wings in bats is due to genetic regulatory changes.
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<div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2></div>
<ul><li><a href="Evolution_of_mammals" title="Evolution of mammals">Evolution of mammals</a></li></ul>
<div class="mw-heading mw-heading2"><h2 id="References">References</h2></div>
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<li id="cite_note-length-8"><span class="mw-cite-backlink"><b><a href="#cite_ref-length_8-0">^</a></b></span> <span class="reference-text"><cite id="CITEREFCretekosWangGreen2008" class="citation journal cs1">Cretekos CJ, Wang Y, Green ED, et&nbsp;al. (2008). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2192750">"Regulatory divergence modifies limb length between mammals"</a>. <i>Genes Dev</i>. <b>22</b> (2): <span class="nowrap">144–</span>151. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1101%2Fgad.1620408">10.1101/gad.1620408</a>. <a href="PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a>&nbsp;<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2192750">2192750</a></span>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a>&nbsp;<a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/18198333">18198333</a>.</cite></span>
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<li id="cite_note-flight-9"><span class="mw-cite-backlink">^ <a href="#cite_ref-flight_9-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-flight_9-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><cite id="CITEREFSearsBehringerRasweilerNiswander2006" class="citation journal cs1">Sears KE, Behringer RR, Rasweiler JJ, Niswander LA (2006). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1458926">"Development of bat flight: morphologic and molecular evolution of bat wing digits"</a>. <i>Proceedings of the National Academy of Sciences of the United States of America</i>. <b>103</b> (17): <span class="nowrap">6581–</span>6. <a href="Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2006PNAS..103.6581S">2006PNAS..103.6581S</a>. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.0509716103">10.1073/pnas.0509716103</a></span>. <a href="PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a>&nbsp;<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1458926">1458926</a></span>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a>&nbsp;<a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/16618938">16618938</a>.</cite></span>
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<li id="cite_note-plate-10"><span class="mw-cite-backlink"><b><a href="#cite_ref-plate_10-0">^</a></b></span> <span class="reference-text"><cite id="CITEREFDe_LucaBarnesUyedaDe-Levi2001" class="citation journal cs1">De Luca F, Barnes KM, Uyeda JA, De-Levi S, Abad V, Palese T, Mericq V, Baron J (2001). <a rel="nofollow" class="external text" href="https://doi.org/10.1210%2Fendo.142.1.7901">"Regulation of growth plate chondrogenesis by bone morphogenetic protein-2"</a>. <i>Endocrinology</i>. <b>142</b> (1): <span class="nowrap">430–</span>436. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1210%2Fendo.142.1.7901">10.1210/endo.142.1.7901</a></span>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a>&nbsp;<a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/11145607">11145607</a>.</cite></span>
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<li id="cite_note-shh-11"><span class="mw-cite-backlink"><b><a href="#cite_ref-shh_11-0">^</a></b></span> <span class="reference-text"><cite id="CITEREFChiangLitingtungHarrisSimandl2001" class="citation journal cs1">Chiang C, Litingtung Y, Harris MP, Simandl BK, Li Y, Beachy PA, Fallon JF (2001). <a rel="nofollow" class="external text" href="https://doi.org/10.1006%2Fdbio.2001.0346">"Manifestation of the limb prepattern: limb development in the absence of sonic hedgehog pattern"</a>. <i>Dev. Biol</i>. <b>236</b> (2): <span class="nowrap">421–</span>435. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1006%2Fdbio.2001.0346">10.1006/dbio.2001.0346</a></span>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a>&nbsp;<a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/11476582">11476582</a>.</cite></span>
</li>
<li id="cite_note-hoxy-12"><span class="mw-cite-backlink"><b><a href="#cite_ref-hoxy_12-0">^</a></b></span> <span class="reference-text"><cite id="CITEREFChenCretekosRasweilerBehringer2005" class="citation journal cs1">Chen CH, Cretekos CJ, Rasweiler JJ, Behringer RR (2005). "Hoxd13 expression in the developing limbs of the short-tailed fruit bat, Carollia perspicillata". <i>Evolution &amp; Development</i>. <b>7</b> (2): <span class="nowrap">130–</span>141. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fj.1525-142X.2005.05015.x">10.1111/j.1525-142X.2005.05015.x</a>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a>&nbsp;<a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/15733311">15733311</a>. <a href="S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a>&nbsp;<a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:12204277">12204277</a>.</cite></span>
</li>
</ol></div>
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</style><div id="Evolutionary_biology753" style="font-size:114%;margin:0 4em"><a href="Evolutionary_biology" title="Evolutionary biology">Evolutionary biology</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div>
<ul><li><a href="Introduction_to_evolution" title="Introduction to evolution">Introduction</a></li>
<li><a href="Outline_of_evolution" title="Outline of evolution">Outline</a></li>
<li><a href="Timeline_of_the_evolutionary_history_of_life" title="Timeline of the evolutionary history of life">Timeline of evolution</a></li>
<li><a href="History_of_life" title="History of life">History of life</a></li>
<li><a href="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="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="Abiogenesis" title="Abiogenesis">Abiogenesis</a></li>
<li><a href="Adaptation" title="Adaptation">Adaptation</a></li>
<li><a href="Adaptive_radiation" title="Adaptive radiation">Adaptive radiation</a></li>
<li><a href="Altruism_(biology)" title="Altruism (biology)">Altruism</a>
<ul><li><a href="Cheating_(biology)" title="Cheating (biology)">Cheating</a></li>
<li><a href="Reciprocal_altruism" title="Reciprocal altruism">Reciprocal</a></li></ul></li>
<li><a href="Baldwin_effect" title="Baldwin effect">Baldwin effect</a></li>
<li><a href="Cladistics" title="Cladistics">Cladistics</a></li>
<li><a href="Coevolution" title="Coevolution">Coevolution</a>
<ul><li><a href="Mutualism_(biology)" title="Mutualism (biology)">Mutualism</a></li></ul></li>
<li><a href="Common_descent" title="Common descent">Common descent</a></li>
<li><a href="Convergent_evolution" title="Convergent evolution">Convergence</a></li>
<li><a href="Divergent_evolution" title="Divergent evolution">Divergence</a></li>
<li><a href="Earliest_known_life_forms" title="Earliest known life forms">Earliest known life forms</a></li>
<li><a href="Evidence_of_common_descent" title="Evidence of common descent">Evidence of evolution</a></li>
<li><a href="Evolutionary_arms_race" title="Evolutionary arms race">Evolutionary arms race</a></li>
<li><a href="Evolutionary_pressure" title="Evolutionary pressure">Evolutionary pressure</a></li>
<li><a href="Exaptation" title="Exaptation">Exaptation</a></li>
<li><a href="Extinction" title="Extinction">Extinction</a>
<ul><li><a href="Extinction_event" title="Extinction event">Event</a></li></ul></li>
<li><a href="Homology_(biology)" title="Homology (biology)">Homology</a></li>
<li><a href="Last_universal_common_ancestor" title="Last universal common ancestor">Last universal common ancestor</a></li>
<li><a href="Macroevolution" title="Macroevolution">Macroevolution</a></li>
<li><a href="Microevolution" title="Microevolution">Microevolution</a></li>
<li><a href="Evolutionary_mismatch" title="Evolutionary mismatch">Mismatch</a></li>
<li><a href="Nonadaptive_radiation" title="Nonadaptive radiation">Non-adaptive radiation</a></li>
<li><a href="Abiogenesis" title="Abiogenesis">Origin of life</a></li>
<li><a href="Panspermia" title="Panspermia">Panspermia</a></li>
<li><a href="Parallel_evolution" title="Parallel evolution">Parallel evolution</a></li>
<li><a href="Signalling_theory" title="Signalling theory">Signalling theory</a>
<ul><li><a href="Handicap_principle" title="Handicap principle">Handicap principle</a></li></ul></li>
<li><a href="Speciation" title="Speciation">Speciation</a>
<ul><li><a href="Species" title="Species">Species</a></li>
<li><a href="Species_complex" title="Species complex">Species complex</a></li></ul></li>
<li><a href="Evolutionary_taxonomy" title="Evolutionary taxonomy">Taxonomy</a></li>
<li><a href="Unit_of_selection" title="Unit of selection">Unit of selection</a>
<ul><li><a href="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="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="Selective_breeding" title="Selective breeding">Artificial selection</a></li>
<li><a href="Biodiversity" title="Biodiversity">Biodiversity</a></li>
<li><a href="Evolutionarily_stable_strategy" title="Evolutionarily stable strategy">Evolutionarily stable strategy</a></li>
<li><a href="Fisher's_principle" title="Fisher's principle">Fisher's principle</a></li>
<li><a href="Fitness_(biology)" title="Fitness (biology)">Fitness</a>
<ul><li><a href="Inclusive_fitness" title="Inclusive fitness">Inclusive</a></li></ul></li>
<li><a href="Gene_flow" title="Gene flow">Gene flow</a></li>
<li><a href="Genetic_drift" title="Genetic drift">Genetic drift</a></li>
<li><a href="Kin_selection" title="Kin selection">Kin selection</a>
<ul><li><a href="Inbreeding_avoidance" title="Inbreeding avoidance">Inbreeding avoidance</a></li>
<li><a href="Kin_recognition" title="Kin recognition">Kin recognition</a></li>
<li><a href="Parental_investment" title="Parental investment">Parental investment</a></li>
<li><a href="Parent%E2%80%93offspring_conflict" title="Parent–offspring conflict">Parent–offspring conflict</a></li></ul></li>
<li><a href="Mutation" title="Mutation">Mutation</a></li>
<li><a href="Population" title="Population">Population</a></li>
<li><a href="Natural_selection" title="Natural selection">Natural selection</a></li>
<li><a href="Sexual_dimorphism" title="Sexual dimorphism">Sexual dimorphism</a></li>
<li><a href="Sexual_selection" title="Sexual selection">Sexual selection</a>
<ul><li><a href="Sexual_selection_in_flowering_plants" title="Sexual selection in flowering plants">Flowering plants</a></li>
<li><a href="Sexual_selection_in_fungi" title="Sexual selection in fungi">Fungi</a></li>
<li><a href="Mate_choice" title="Mate choice">Mate choice</a></li></ul></li>
<li><a href="Social_selection" title="Social selection">Social selection</a></li>
<li><a href="Trivers%E2%80%93Willard_hypothesis" title="Trivers–Willard hypothesis">Trivers–Willard hypothesis</a></li>
<li><a href="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="Canalisation_(genetics)" title="Canalisation (genetics)">Canalisation</a></li>
<li><a href="Evolutionary_developmental_biology" title="Evolutionary developmental biology">Evolutionary developmental biology</a></li>
<li><a href="Genetic_assimilation" title="Genetic assimilation">Genetic assimilation</a></li>
<li><a href="Inversion_(evolutionary_biology)" title="Inversion (evolutionary biology)">Inversion</a></li>
<li><a href="Modularity_(biology)" title="Modularity (biology)">Modularity</a></li>
<li><a href="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="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="Evolution_of_bacteria" title="Evolution of bacteria">Bacteria</a></li>
<li><a href="Evolution_of_birds" title="Evolution of birds">Birds</a>
<ul><li><a href="Origin_of_birds" title="Origin of birds">origin</a></li></ul></li>
<li><a href="Evolution_of_brachiopods" title="Evolution of brachiopods">Brachiopods</a></li>
<li><a href="Evolution_of_molluscs" title="Evolution of molluscs">Molluscs</a>
<ul><li><a href="Evolution_of_cephalopods" title="Evolution of cephalopods">Cephalopods</a></li></ul></li>
<li><a href="Dinosaur#Evolutionary_history" title="Dinosaur">Dinosaurs</a></li>
<li><a href="Evolution_of_fish" title="Evolution of fish">Fish</a></li>
<li><a href="Evolution_of_fungi" title="Evolution of fungi">Fungi</a></li>
<li><a href="Evolution_of_insects" title="Evolution of insects">Insects</a>
<ul><li><a href="Evolution_of_butterflies" title="Evolution of butterflies">butterflies</a></li></ul></li>
<li><a href="History_of_life" title="History of life">Life</a></li>
<li><a href="Evolution_of_mammals" title="Evolution of mammals">Mammals</a>
<ul><li><a href="Cat_gap" title="Cat gap">cats</a></li>
<li><a href="Canidae#Evolution" title="Canidae">canids</a>
<ul><li><a href="Evolution_of_the_wolf" title="Evolution of the wolf">wolves</a></li>
<li><a href="Domestication_of_the_dog" title="Domestication of the dog">dogs</a></li></ul></li>
<li><a href="Hyena#Evolution" title="Hyena">hyenas</a></li>
<li><a href="Evolution_of_cetaceans" title="Evolution of cetaceans">dolphins and whales</a></li>
<li><a href="Evolution_of_the_horse" title="Evolution of the horse">horses</a></li>
<li><a href="Evolution_of_Macropodidae" title="Evolution of Macropodidae">Kangaroos</a></li>
<li><a href="Evolution_of_primates" title="Evolution of primates">primates</a>
<ul><li><a href="Human_evolution" title="Human evolution">humans</a></li>
<li><a href="Evolution_of_lemurs" title="Evolution of lemurs">lemurs</a></li></ul></li>
<li><a href="Evolution_of_sirenians" title="Evolution of sirenians">sea cows</a></li></ul></li>
<li><a href="Evolutionary_history_of_plants" title="Evolutionary history of plants">Plants</a>
<ul><li><a href="Pollinator-mediated_selection" title="Pollinator-mediated selection">pollinator-mediated</a></li></ul></li>
<li><a href="Evolution_of_reptiles" title="Evolution of reptiles">Reptiles</a></li>
<li><a href="Evolution_of_spiders" title="Evolution of spiders">Spiders</a></li>
<li><a href="Evolution_of_tetrapods" title="Evolution of tetrapods">Tetrapods</a></li>
<li><a href="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="Organ_(biology)" title="Organ (biology)">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="Evolution_of_cells" title="Evolution of cells">Cell</a></li>
<li><a href="Models_of_DNA_evolution" title="Models of DNA evolution">DNA</a></li>
<li><a href="Evolution_of_flagella" title="Evolution of flagella">Flagella</a></li>
<li><a href="Eukaryote#Origin_of_eukaryotes" title="Eukaryote">Eukaryotes</a>
<ul><li><a href="Symbiogenesis" title="Symbiogenesis">symbiogenesis</a></li>
<li><a href="Chromosome#Eukaryotes" title="Chromosome">chromosome</a></li>
<li><a href="Endomembrane_system#Evolution" title="Endomembrane system">endomembrane system</a></li>
<li><a href="Mitochondrion#Origin_and_evolution" title="Mitochondrion">mitochondria</a></li>
<li><a href="Cell_nucleus#Evolution" title="Cell nucleus">nucleus</a></li>
<li><a href="Plastid#Origin" title="Plastid">plastids</a></li></ul></li>
<li>In animals
<ul><li><a href="Evolution_of_the_eye" title="Evolution of the eye">eye</a></li>
<li><a href="Hair#Evolution" title="Hair">hair</a></li>
<li><a href="Evolution_of_mammalian_auditory_ossicles" title="Evolution of mammalian auditory ossicles">auditory ossicle</a></li>
<li><a href="Evolution_of_nervous_systems" title="Evolution of nervous systems">nervous system</a></li>
<li><a href="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="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="Evolution_of_ageing" title="Evolution of ageing">Aging</a>
<ul><li><a href="Death#Evolution_of_aging_and_mortality" title="Death">Death</a></li>
<li><a href="Programmed_cell_death#Evolutionary_origin_of_mitochondrial_apoptosis" title="Programmed cell death">Programmed cell death</a></li></ul></li>
<li><a href="Origin_of_avian_flight" title="Origin of avian flight">Avian flight</a></li>
<li><a href="Evolution_of_biological_complexity" title="Evolution of biological complexity">Biological complexity</a></li>
<li><a href="Cooperation_(evolution)" title="Cooperation (evolution)">Cooperation</a></li>
<li><a href="Evolution_of_color_vision" title="Evolution of color vision">Color vision</a>
<ul><li><a href="Evolution_of_color_vision_in_primates" title="Evolution of color vision in primates">in primates</a></li></ul></li>
<li><a href="Evolution_of_emotion" title="Evolution of emotion">Emotion</a></li>
<li><a href="Empathy#Evolution_across_species" title="Empathy">Empathy</a></li>
<li><a href="Evolutionary_ethics" title="Evolutionary ethics">Ethics</a></li>
<li><a href="Evolution_of_eusociality" title="Evolution of eusociality">Eusociality</a></li>
<li><a href="Immune_system#Evolution_and_other_mechanisms" title="Immune system">Immune system</a></li>
<li><a href="Metabolism#Evolution" title="Metabolism">Metabolism</a></li>
<li><a href="Monogamy_in_animals" title="Monogamy in animals">Monogamy</a></li>
<li><a href="Evolution_of_morality" title="Evolution of morality">Morality</a></li>
<li><a href="Mosaic_evolution" title="Mosaic evolution">Mosaic evolution</a></li>
<li><a href="Multicellular_organism#Evolutionary_history" title="Multicellular organism">Multicellularity</a></li>
<li><a href="Evolution_of_sexual_reproduction" title="Evolution of sexual reproduction">Sexual reproduction</a>
<ul><li><a href="Anisogamy#Evolution" title="Anisogamy">Gamete differentiation/sexes</a></li>
<li><a href="Biological_life_cycle" title="Biological life cycle">Life cycles/nuclear phases</a></li>
<li><a href="Mating_type" title="Mating type">Mating types</a></li>
<li><a href="Origin_and_function_of_meiosis" title="Origin and function of meiosis">Meiosis</a></li>
<li><a href="Sex-determination_system#Evolution" title="Sex-determination system">Sex-determination</a></li></ul></li>
<li><a href="Evolution_of_snake_venom" title="Evolution of snake venom">Snake venom</a></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="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="Phyletic_gradualism" title="Phyletic gradualism">Gradualism</a>/<a href="Punctuated_equilibrium" title="Punctuated equilibrium">Punctuated equilibrium</a>/<a href="Saltation_(biology)" title="Saltation (biology)">Saltationism</a></li>
<li><a href="Point_mutation" title="Point mutation">Micromutation</a>/<a href="Mutationism#Later_mutationist_theories" title="Mutationism">Macromutation</a></li>
<li><a href="Uniformitarianism" title="Uniformitarianism">Uniformitarianism</a>/<a href="Catastrophism" title="Catastrophism">Catastrophism</a></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="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="Allopatric_speciation" title="Allopatric speciation">Allopatric</a></li>
<li><a href="Anagenesis" title="Anagenesis">Anagenesis</a></li>
<li><a href="Catagenesis_(biology)" title="Catagenesis (biology)">Catagenesis</a></li>
<li><a href="Cladogenesis" title="Cladogenesis">Cladogenesis</a></li>
<li><a href="Cospeciation" title="Cospeciation">Cospeciation</a></li>
<li><a href="Ecological_speciation" title="Ecological speciation">Ecological</a></li>
<li><a href="Hybrid_speciation" title="Hybrid speciation">Hybrid</a></li>
<li><a href="Nonecological_speciation" title="Nonecological speciation">Non-ecological</a></li>
<li><a href="Parapatric_speciation" title="Parapatric speciation">Parapatric</a></li>
<li><a href="Peripatric_speciation" title="Peripatric speciation">Peripatric</a></li>
<li><a href="Reinforcement_(speciation)" title="Reinforcement (speciation)">Reinforcement</a></li>
<li><a href="Sympatric_speciation" title="Sympatric speciation">Sympatric</a></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="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="Evolutionary_ideas_of_the_Renaissance_and_Enlightenment" title="Evolutionary ideas of the Renaissance and Enlightenment">Renaissance and Enlightenment</a></li>
<li><a href="Transmutation_of_species" title="Transmutation of species">Transmutation of species</a></li>
<li><a href="David_Hume" title="David Hume">David Hume</a>
<ul><li><i><a href="Dialogues_Concerning_Natural_Religion" title="Dialogues Concerning Natural Religion">Dialogues Concerning Natural Religion</a></i></li></ul></li>
<li><a href="Charles_Darwin" title="Charles Darwin">Charles Darwin</a>
<ul><li><i><a href="On_the_Origin_of_Species" title="On the Origin of Species">On the Origin of Species</a></i></li></ul></li>
<li><a href="History_of_paleontology" title="History of paleontology">History of paleontology</a></li>
<li><a href="Transitional_fossil" title="Transitional fossil">Transitional fossil</a></li>
<li><a href="Blending_inheritance" title="Blending inheritance">Blending inheritance</a></li>
<li><a href="Mendelian_inheritance" title="Mendelian inheritance">Mendelian inheritance</a></li>
<li><a href="The_eclipse_of_Darwinism" title="The eclipse of Darwinism">The eclipse of Darwinism</a></li>
<li><a href="Neo-Darwinism" title="Neo-Darwinism">Neo-Darwinism</a></li>
<li><a href="Modern_synthesis_(20th_century)" title="Modern synthesis (20th century)">Modern synthesis</a></li>
<li><a href="History_of_molecular_evolution" title="History of molecular evolution">History of molecular evolution</a></li>
<li><a href="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="Darwinism" title="Darwinism">Darwinism</a></li>
<li><a href="Alternatives_to_Darwinian_evolution" title="Alternatives to Darwinian evolution">Alternatives</a>
<ul><li><a href="Catastrophism" title="Catastrophism">Catastrophism</a></li>
<li><a href="Lamarckism" title="Lamarckism">Lamarckism</a></li>
<li><a href="Orthogenesis" title="Orthogenesis">Orthogenesis</a></li>
<li><a href="Mutationism" title="Mutationism">Mutationism</a></li>
<li><a href="Saltation_(biology)" title="Saltation (biology)">Saltationism</a></li>
<li><a href="Structuralism_(biology)" title="Structuralism (biology)">Structuralism</a>
<ul><li><a href="Spandrel_(biology)" title="Spandrel (biology)">Spandrel</a></li></ul></li>
<li><a href="Theistic_evolution" title="Theistic evolution">Theistic</a></li>
<li><a href="Vitalism" title="Vitalism">Vitalism</a></li></ul></li>
<li><a href="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="Biogeography" title="Biogeography">Biogeography</a></li>
<li><a href="Ecological_genetics" title="Ecological genetics">Ecological genetics</a></li>
<li><a href="Evolutionary_medicine" title="Evolutionary medicine">Evolutionary medicine</a></li>
<li><a href="Group_selection" title="Group selection">Group selection</a>
<ul><li><a href="Cultural_evolution" title="Cultural evolution">Cultural evolution</a></li>
<li><a href="Cultural_group_selection" title="Cultural group selection">Cultural group selection</a></li>
<li><a href="Dual_inheritance_theory" title="Dual inheritance theory">Dual inheritance theory</a></li></ul></li>
<li><a href="Hologenome_theory_of_evolution" title="Hologenome theory of evolution">Hologenome theory of evolution</a></li>
<li><a href="Missing_heritability_problem" title="Missing heritability problem">Missing heritability problem</a></li>
<li><a href="Molecular_evolution" title="Molecular evolution">Molecular evolution</a></li>
<li><a href="Astrobiology" title="Astrobiology">Astrobiology</a></li>
<li><a href="Phylogenetics" title="Phylogenetics">Phylogenetics</a>
<ul><li><a href="Phylogenetic_tree" title="Phylogenetic tree">Tree</a></li></ul></li>
<li><a href="Polymorphism_(biology)" title="Polymorphism (biology)">Polymorphism</a></li>
<li><a href="Protocell" title="Protocell">Protocell</a></li>
<li><a href="Systematics" title="Systematics">Systematics</a></li>
<li><a href="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"></span></span> Category</li>
<li><span class="noviewer" typeof="mw:File"></span> <a href="Portal%3AEvolutionary_biology" title="Portal:Evolutionary biology">Portal</a></li></ul>
</div></td></tr></tbody></table></div>
<div class="navbox-styles"></div><div role="navigation" class="navbox" aria-labelledby="Fins,_limbs_and_wings105" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="3"><div id="Fins,_limbs_and_wings105" style="font-size:114%;margin:0 4em"><a href="Fin" title="Fin">Fins</a>, <a href="Limb_(anatomy)" title="Limb (anatomy)">limbs</a> and <a href="Wing" title="Wing">wings</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Fin" title="Fin">Fins</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="Aquatic_locomotion" title="Aquatic locomotion">Aquatic locomotion</a></li>
<li><a href="Cephalopod_fin" title="Cephalopod fin">Cephalopod fin</a></li>
<li><a href="Fish_locomotion" title="Fish locomotion">Fish locomotion</a></li>
<li><a href="Fin_and_flipper_locomotion" title="Fin and flipper locomotion">Fin and flipper locomotion</a></li>
<li><a href="Caudal_fin" class="mw-redirect" title="Caudal fin">Caudal fin</a></li>
<li><a href="Dorsal_fin" title="Dorsal fin">Dorsal fin</a></li>
<li><a href="Fish_fin" title="Fish fin">Fish fin</a></li>
<li><a href="Flipper_(anatomy)" title="Flipper (anatomy)">Flipper</a></li>
<li><a href="Sarcopterygii" title="Sarcopterygii">Lobe-finned fish</a></li>
<li><a href="Actinopterygii" title="Actinopterygii">Ray-finned fish</a></li>
<li><a href="Pectoral_fins" class="mw-redirect" title="Pectoral fins">Pectoral fins</a></li>
<li><a href="Pelvic_fin" title="Pelvic fin">Pelvic fin</a></li></ul>
</div></td><td class="noviewer navbox-image" rowspan="5" style="width:1px;padding:0 0 0 2px"><div><span typeof="mw:File"></span></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Limb_(anatomy)" title="Limb (anatomy)">Limbs</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="Limb_development" title="Limb development">Limb development</a></li>
<li><a href="Comparative_foot_morphology" title="Comparative foot morphology">Limb morphology</a>
<ul><li><a href="Digitigrade" title="Digitigrade">digitigrade</a></li>
<li><a href="Plantigrade" title="Plantigrade">plantigrade</a></li>
<li><a href="Unguligrade" class="mw-redirect" title="Unguligrade">unguligrade</a></li>
<li><a href="Uniped" class="mw-redirect" title="Uniped">uniped</a></li>
<li><a href="Bipedalism" title="Bipedalism">biped</a></li>
<li><a href="Facultative_biped" class="mw-redirect" title="Facultative biped">facultative biped</a></li>
<li><a href="Tripedal" class="mw-redirect" title="Tripedal">triped</a></li>
<li><a href="Quadrupedalism" title="Quadrupedalism">quadruped</a></li></ul></li>
<li><a href="Arthropod_leg" title="Arthropod leg">Arthropod</a></li>
<li><a href="Cephalopod_limb" title="Cephalopod limb">Cephalopod</a></li>
<li><a href="Tetrapod#Limbs" title="Tetrapod">Tetrapod</a>
<ul><li><a href="Dactyly" title="Dactyly">dactyly</a></li></ul></li>
<li><a href="Digit_(anatomy)" title="Digit (anatomy)">Digit</a></li>
<li><a href="Webbed_foot" title="Webbed foot">Webbed foot</a></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Wing" title="Wing">Wings</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="Flying_and_gliding_animals" title="Flying and gliding animals">Flying and gliding animals</a></li>

<li><a href="Bird_wing" title="Bird wing">Bird wing</a>
<ul><li><a href="Keel_(bird_anatomy)" title="Keel (bird anatomy)">keel</a></li>
<li><a href="Bird_anatomy#Skeletal_system" title="Bird anatomy">skeleton</a></li>
<li><a href="Flight_feather" title="Flight feather">feathers</a></li></ul></li>
<li><a href="Insect_wing" title="Insect wing">Insect wing</a></li>
<li><a href="Pterosaur#Wings" title="Pterosaur">Pterosaur wing</a></li>
<li><a href="Wingspan" title="Wingspan">Wingspan</a></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Evolution</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="Evolution_of_fish" title="Evolution of fish">Evolution of fish</a></li>
<li><a href="Evolution_of_tetrapods" title="Evolution of tetrapods">Evolution of tetrapods</a></li>
<li><a href="Evolution_of_birds" title="Evolution of birds">Evolution of birds</a></li>
<li><a href="Origin_of_birds" title="Origin of birds">Origin of birds</a></li>
<li><a href="Origin_of_avian_flight" title="Origin of avian flight">Origin of avian flight</a></li>
<li><a href="Evolution_of_cetaceans" title="Evolution of cetaceans">Evolution of cetaceans</a></li>
<li><a href="Comparative_anatomy" title="Comparative anatomy">Comparative anatomy</a></li>
<li><a href="Convergent_evolution" title="Convergent evolution">Convergent evolution</a></li>
<li><a href="Analogy_(biology)" class="mw-redirect" title="Analogy (biology)">Analogous structures</a></li>
<li><a href="Homology_(biology)" title="Homology (biology)">Homologous structures</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="Animal_locomotion" title="Animal locomotion">Animal locomotion</a></li>
<li><a href="Gait" title="Gait">Gait</a></li>
<li><a href="Robot_locomotion" title="Robot locomotion">Robot locomotion</a></li>
<li><a href="Samara_(fruit)" title="Samara (fruit)">Samara</a></li>
<li><a href="Terrestrial_locomotion" title="Terrestrial locomotion">Terrestrial locomotion</a></li>
<li><a href="Tradeoffs_for_locomotion_in_air_and_water" title="Tradeoffs for locomotion in air and water">Tradeoffs for locomotion in air and water</a></li>
<li><a href="Rotating_locomotion_in_living_systems" title="Rotating locomotion in living systems">Rotating locomotion</a></li>
<li><a href="Undulatory_locomotion" title="Undulatory locomotion">Undulatory locomotion</a></li></ul>
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