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Convergent evolution
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<h2 id="overview">Overview</h2> <p class="note">Further information: <a href="/facts/List_of_examples_of_convergent_evolution/GLd7WlMu">List of examples of convergent evolution</a></p> <p>In morphology, analogous traits arise when different species live in similar ways and/or a similar environment, and so face the same environmental factors. When occupying similar <a href="/facts/Ecological_niche/YnhNkL2e">ecological niches</a> (that is, a distinctive way of life) similar problems can lead to similar solutions.<a class="footnote-ref" id="fnref:1" href="#fn:1"><sup>1</sup></a><a class="footnote-ref" id="fnref:2" href="#fn:2"><sup>2</sup></a><a class="footnote-ref" id="fnref:3" href="#fn:3"><sup>3</sup></a> The British anatomist <a href="/facts/Richard_Owen/M8H3td41">Richard Owen</a> was the first to identify the fundamental difference between analogies and <a href="/facts/Homology_(biology)/muctKlyT">homologies</a>.<a class="footnote-ref" id="fnref:4" href="#fn:4"><sup>4</sup></a> </p><p>In biochemistry, physical and chemical constraints on <a href="/facts/Enzyme_mechanism/4a283vgL">mechanisms</a> have caused some <a href="/facts/Active_site/u9ua7rGu">active site</a> arrangements such as the <a href="/facts/Catalytic_triad/WskaH1zM">catalytic triad</a> to evolve independently in separate <a href="/facts/Enzyme_superfamilies/FJTXCha0">enzyme superfamilies</a>.<a class="footnote-ref" id="fnref:5" href="#fn:5"><sup>5</sup></a> </p><p>In his 1989 book <i><a href="/facts/Wonderful_Life_(book)/d975HBhr">Wonderful Life</a></i>, <a href="/facts/Stephen_Jay_Gould/FtAcabjg">Stephen Jay Gould</a> argued that if one could "rewind the tape of life [and] the same conditions were encountered again, evolution could take a very different course."<a class="footnote-ref" id="fnref:6" href="#fn:6"><sup>6</sup></a> <a href="/facts/Simon_Conway_Morris/c4kHVblJ">Simon Conway Morris</a> disputes this conclusion, arguing that convergence is a dominant force in evolution, and given that the same environmental and physical constraints are at work, life will inevitably evolve toward an "optimum" body plan, and at some point, evolution is bound to stumble upon <a href="/facts/Animal_intelligence/LpP3H18I">intelligence</a>, a trait presently identified with at least <a href="/facts/Primates/Hl8RNE2X">primates</a>, <a href="/facts/Corvids/PAr1u8fF">corvids</a>, and <a href="/facts/Cetaceans/rtC33h63">cetaceans</a>.<a class="footnote-ref" id="fnref:7" href="#fn:7"><sup>7</sup></a> </p> <h2 id="distinctions">Distinctions</h2> <h3>Cladistics</h3> <p class="note">Main article: <a href="/facts/Cladistics/JasKVABk">Cladistics</a></p> <p>In cladistics, a homoplasy is a trait shared by two or more <a href="/facts/Taxon/caQC1e87">taxa</a> for any reason other than that they share a common ancestry. Taxa which do share ancestry are part of the same <a href="/facts/Clade/XefyYmWn">clade</a>; cladistics seeks to arrange them according to their degree of relatedness to describe their <a href="/facts/Phylogeny/f5vVEpeY">phylogeny</a>. Homoplastic traits caused by convergence are therefore, from the point of view of cladistics, confounding factors which could lead to an incorrect analysis.<a class="footnote-ref" id="fnref:8" href="#fn:8"><sup>8</sup></a><a class="footnote-ref" id="fnref:9" href="#fn:9"><sup>9</sup></a><a class="footnote-ref" id="fnref:10" href="#fn:10"><sup>10</sup></a><a class="footnote-ref" id="fnref:11" href="#fn:11"><sup>11</sup></a> </p> <h3>Atavism</h3> <p class="note">Main article: <a href="/facts/Atavism/GDGHCs3R">Atavism</a></p> <p>In some cases, it is difficult to tell whether a trait has been lost and then re-evolved convergently, or whether a gene has simply been switched off and then re-enabled later. Such a re-emerged trait is called an <a href="/facts/Atavism/GDGHCs3R">atavism</a>. From a mathematical standpoint, an unused gene (<a href="/facts/Genetic_drift/lvoEEctX">selectively neutral</a>) has a steadily decreasing <a href="/facts/Probability/fgYvMhwb">probability</a> of retaining potential functionality over time. The time scale of this process varies greatly in different phylogenies; in mammals and birds, there is a reasonable probability of remaining in the genome in a potentially functional state for around 6 million years.<a class="footnote-ref" id="fnref:12" href="#fn:12"><sup>12</sup></a> </p> <h3>Parallel vs. convergent evolution</h3> <p>When two species are similar in a particular character, evolution is defined as parallel if the ancestors were also similar, and convergent if they were not.<a class="footnote-ref" id="fnref:13" href="#fn:13"><sup>13</sup></a> Some scientists have argued that there is a continuum between parallel and convergent evolution,<a class="footnote-ref" id="fnref:14" href="#fn:14"><sup>14</sup></a><a class="footnote-ref" id="fnref:15" href="#fn:15"><sup>15</sup></a><a class="footnote-ref" id="fnref:16" href="#fn:16"><sup>16</sup></a><a class="footnote-ref" id="fnref:17" href="#fn:17"><sup>17</sup></a> while others maintain that despite some overlap, there are still important distinctions between the two.<a class="footnote-ref" id="fnref:18" href="#fn:18"><sup>18</sup></a><a class="footnote-ref" id="fnref:19" href="#fn:19"><sup>19</sup></a> </p><p>When the ancestral forms are unspecified or unknown, or the range of traits considered is not clearly specified, the distinction between parallel and convergent evolution becomes more subjective. For instance, the striking example of similar placental and marsupial forms is described by <a href="/facts/Richard_Dawkins/x8Q24PJG">Richard Dawkins</a> in <i><a href="/facts/The_Blind_Watchmaker/HuzXYmae">The Blind Watchmaker</a></i> as a case of convergent evolution, because mammals on each continent had a long evolutionary history prior to the extinction of the dinosaurs under which to accumulate relevant differences.<a class="footnote-ref" id="fnref:20" href="#fn:20"><sup>20</sup></a> </p> <h2 id="at-molecular-level">At molecular level</h2> <h3>Proteins</h3> <h4>Tertiary structures</h4> <p>Many proteins share analogous <a href="/facts/Protein_structure/BptnTs9X">structural elements</a> that arose independently across different genomes. There are several examples of convergent protein motifs sharing similar arrangements of structural elements.<a class="footnote-ref" id="fnref:21" href="#fn:21"><sup>21</sup></a> Whole protein structures too have arisen through convergent evolution.<a class="footnote-ref" id="fnref:22" href="#fn:22"><sup>22</sup></a> </p> <h4>Protease active sites</h4> <p>The <a href="/facts/Enzymology/DSI1Fh7y">enzymology</a> of <a href="/facts/Proteases/G1ASdVat">proteases</a> provides some of the clearest examples of convergent evolution. These examples reflect the intrinsic chemical constraints on enzymes, leading evolution to converge on equivalent solutions independently and repeatedly.<a class="footnote-ref" id="fnref:23" href="#fn:23"><sup>23</sup></a><a class="footnote-ref" id="fnref:24" href="#fn:24"><sup>24</sup></a> </p><p>Serine and cysteine proteases use different amino acid functional groups (alcohol or thiol) as a <a href="/facts/Nucleophile/xYccoaJQ">nucleophile</a>. To activate that nucleophile, they orient an acidic and a basic residue in a <a href="/facts/Catalytic_triad/WskaH1zM">catalytic triad</a>. The chemical and physical constraints on <a href="/facts/Enzyme_catalysis/4a283vgL">enzyme catalysis</a> have caused identical triad arrangements to evolve independently more than 20 times in different <a href="/facts/Enzyme_superfamilies/FJTXCha0">enzyme superfamilies</a>.<a class="footnote-ref" id="fnref:25" href="#fn:25"><sup>25</sup></a> </p><p><a href="/facts/Threonine_protease/HIQQrmvJ">Threonine proteases</a> use the amino acid threonine as their catalytic <a href="/facts/Nucleophile/xYccoaJQ">nucleophile</a>. Unlike cysteine and serine, threonine is a <a href="/facts/Secondary_alcohol/Y0SCTDkh">secondary alcohol</a> (i.e. has a methyl group). The methyl group of threonine greatly restricts the possible orientations of triad and substrate, as the methyl clashes with either the enzyme backbone or the histidine base. Consequently, most threonine proteases use an N-terminal threonine in order to avoid such <a href="/facts/Steric_clash/TnX5c8HG">steric clashes</a>. Several evolutionarily independent <a href="/facts/Enzyme_superfamilies/FJTXCha0">enzyme superfamilies</a> with different <a href="/facts/Protein_fold/FJTXCha0">protein folds</a> use the N-terminal residue as a nucleophile. This commonality of <a href="/facts/Active_site/u9ua7rGu">active site</a> but difference of protein fold indicates that the active site evolved convergently in those families.<a class="footnote-ref" id="fnref:26" href="#fn:26"><sup>26</sup></a><a class="footnote-ref" id="fnref:27" href="#fn:27"><sup>27</sup></a> </p> <h4>Cone snail and fish insulin</h4> <p><i><a href="/facts/Conus_geographus/QPxcjSqz">Conus geographus</a></i> produces a distinct form of <a href="/facts/Insulin/oXG0myt1">insulin</a> that is more similar to fish insulin protein sequences than to insulin from more closely related molluscs, suggesting convergent evolution,<a class="footnote-ref" id="fnref:28" href="#fn:28"><sup>28</sup></a> though with the possibility of <a href="/facts/Horizontal_gene_transfer/FPyUIsf2">horizontal gene transfer</a>.<a class="footnote-ref" id="fnref:29" href="#fn:29"><sup>29</sup></a> </p> <h4>Ferrous iron uptake via protein transporters in land plants and chlorophytes</h4> <p>Distant homologues of the metal ion transporters <a href="/facts/Zinc_transporter_protein/sBkh3UXA">ZIP</a> in <a href="/facts/Land_plants/8pzYKGvv">land plants</a> and <a href="/facts/Chlorophytes/hRL4moDT">chlorophytes</a> have converged in structure, likely to take up Fe2+ efficiently. The IRT1 proteins from <i><a href="/facts/Arabidopsis_thaliana/wt2IMJsC">Arabidopsis thaliana</a></i> and <a href="/facts/Rice/wvL1hBCz">rice</a> have extremely different amino acid sequences from <i><a href="/facts/Chlamydomonas/Skokunvd">Chlamydomonas</a></i>'s IRT1, but their three-dimensional structures are similar, suggesting convergent evolution.<a class="footnote-ref" id="fnref:30" href="#fn:30"><sup>30</sup></a> </p> <h4>Na+,K+-ATPase and Insect resistance to cardiotonic steroids</h4> <p>Many examples of convergent evolution exist in insects in terms of developing resistance at a molecular level to toxins. One well-characterized example is the evolution of resistance to cardiotonic steroids (CTSs) via amino acid substitutions at well-defined positions of the α-subunit of <a href="/facts/Na%2B/K%2B-ATPase/SuA4d4Na">Na+,K+-ATPase</a> (ATPalpha). Variation in ATPalpha has been surveyed in various CTS-adapted species spanning six insect orders.<a class="footnote-ref" id="fnref:31" href="#fn:31"><sup>31</sup></a><a class="footnote-ref" id="fnref:32" href="#fn:32"><sup>32</sup></a><a class="footnote-ref" id="fnref:33" href="#fn:33"><sup>33</sup></a> Among 21 CTS-adapted species, 58 (76%) of 76 amino acid substitutions at sites implicated in CTS resistance occur in parallel in at least two lineages.<a class="footnote-ref" id="fnref:34" href="#fn:34"><sup>34</sup></a> 30 of these substitutions (40%) occur at just two sites in the protein (positions 111 and 122). CTS-adapted species have also recurrently evolved <a href="/facts/Neofunctionalization/BCfPYEhz">neo-functionalized</a> duplications of ATPalpha, with convergent tissue-specific expression patterns.<a class="footnote-ref" id="fnref:35" href="#fn:35"><sup>35</sup></a><a class="footnote-ref" id="fnref:36" href="#fn:36"><sup>36</sup></a> </p> <h3>Nucleic acids</h3> <p>Convergence occurs at the level of <a href="/facts/DNA/nB89Iauz">DNA</a> and the <a href="/facts/Amino_acid/WDxYwBny">amino acid</a> sequences produced by <a href="/facts/Translation_(biology)/JcPC1rth">translating</a> <a href="/facts/Structural_gene/dhTA2nkS">structural genes</a> into <a href="/facts/Protein/Jxmagu3E">proteins</a>. Studies have found convergence in amino acid sequences in echolocating bats and the dolphin;<a class="footnote-ref" id="fnref:37" href="#fn:37"><sup>37</sup></a> among marine mammals;<a class="footnote-ref" id="fnref:38" href="#fn:38"><sup>38</sup></a> between giant and red pandas;<a class="footnote-ref" id="fnref:39" href="#fn:39"><sup>39</sup></a> and between the thylacine and canids.<a class="footnote-ref" id="fnref:40" href="#fn:40"><sup>40</sup></a> Convergence has also been detected in a type of <a href="/facts/Non-coding_DNA/zyTU8q8M">non-coding DNA</a>, <a href="/facts/Cis-regulatory_element/q6u6VSBs">cis-regulatory elements</a>, such as in their rates of evolution; this could indicate either <a href="/facts/Positive_selection/gHPnrflX">positive selection</a> or relaxed <a href="/facts/Negative_selection_(natural_selection)/DCkRLqb4">purifying selection</a>.<a class="footnote-ref" id="fnref:41" href="#fn:41"><sup>41</sup></a><a class="footnote-ref" id="fnref:42" href="#fn:42"><sup>42</sup></a> </p> <h2 id="in-animal-morphology">In animal morphology</h2> <h3>Bodyplans</h3> <p>Swimming animals including <a href="/facts/Fish/fln4gH9V">fish</a> such as <a href="/facts/Herring/QH12ADpV">herrings</a>, <a href="/facts/Marine_mammals/WCoC2ye8">marine mammals</a> such as <a href="/facts/Dolphins/NUxUP7Ph">dolphins</a>, and <a href="/facts/Ichthyosaur/fBNIIJaJ">ichthyosaurs</a> (<a href="/facts/Mesozoic_era/yYsvXGPa">of the Mesozoic</a>) all converged on the same streamlined shape.<a class="footnote-ref" id="fnref:43" href="#fn:43"><sup>43</sup></a><a class="footnote-ref" id="fnref:44" href="#fn:44"><sup>44</sup></a> A similar shape and swimming adaptations are even present in molluscs, such as <i><a href="/facts/Phylliroe/emRmE5bH">Phylliroe</a></i>.<a class="footnote-ref" id="fnref:45" href="#fn:45"><sup>45</sup></a> The fusiform bodyshape (a tube tapered at both ends) adopted by many aquatic animals is an adaptation to enable them to <a href="/facts/Animal_locomotion/trhhYTVR">travel at high speed</a> in a high <a href="/facts/Drag_(physics)/vdl4jF7I">drag</a> environment.<a class="footnote-ref" id="fnref:46" href="#fn:46"><sup>46</sup></a> Similar body shapes are found in the <a href="/facts/Earless_seal/7fFosRas">earless seals</a> and the <a href="/facts/Eared_seals/7lhN2Klw">eared seals</a>: they still have four legs, but these are strongly modified for swimming.<a class="footnote-ref" id="fnref:47" href="#fn:47"><sup>47</sup></a> </p><p>The marsupial fauna of Australia and the placental mammals of the Old World have several strikingly similar forms, developed in two clades, isolated from each other.<a class="footnote-ref" id="fnref:48" href="#fn:48"><sup>48</sup></a> The body, and especially the skull shape, of the <a href="/facts/Thylacine/maFXXmmJ">thylacine</a> (Tasmanian tiger or Tasmanian wolf) converged with those of <a href="/facts/Canidae/5Fj02eeS">Canidae</a> such as the red fox, <i><a href="/facts/Vulpes_vulpes/sJCRHyt8">Vulpes vulpes</a></i>.<a class="footnote-ref" id="fnref:49" href="#fn:49"><sup>49</sup></a> </p> <h3>Echolocation</h3> <p>As a sensory adaptation, <a href="/facts/Animal_echolocation/2hggutJV">echolocation</a> has evolved separately in <a href="/facts/Cetaceans/rtC33h63">cetaceans</a> (dolphins and whales) and bats, but from the same genetic mutations.<a class="footnote-ref" id="fnref:50" href="#fn:50"><sup>50</sup></a> </p> <h3>Electric fishes</h3> <p>The <a href="/facts/Gymnotiformes/qxAk3qqJ">Gymnotiformes</a> of South America and the <a href="/facts/Mormyridae/yKbL7X3n">Mormyridae</a> of Africa independently evolved <a href="/facts/Electroreception_and_electrogenesis/W19Jo0ay">passive electroreception</a> (around 119 and 110 million years ago, respectively). Around 20 million years after acquiring that ability, both groups evolved active <a href="/facts/Electric_fish/nUoAsvqm">electrogenesis</a>, producing weak electric fields to help them detect prey.<a class="footnote-ref" id="fnref:51" href="#fn:51"><sup>51</sup></a> </p> <h3>Eyes</h3> <p class="note">Main article: <a href="/facts/Eye_evolution/gxtqI2QM">Eye evolution</a></p> <p>One of the best-known examples of convergent evolution is the camera eye of <a href="/facts/Cephalopod_eye/FaK6em8v">cephalopods</a> (such as squid and octopus), <a href="/facts/Vertebrate/tT54FSTT">vertebrates</a> (including mammals) and <a href="/facts/Cnidaria/d0HRc6w0">cnidarians</a> (such as jellyfish).<a class="footnote-ref" id="fnref:52" href="#fn:52"><sup>52</sup></a> Their last common ancestor had at most a simple photoreceptive spot, but a range of processes led to the <a href="/facts/Evolution_of_the_eye/gxtqI2QM">progressive refinement of camera eyes</a>—with one sharp difference: the cephalopod eye is "wired" in the opposite direction, with blood and nerve vessels entering from the back of the retina, rather than the front as in vertebrates. As a result, vertebrates have a <a href="/facts/Blind_spot_(vision)/XKEkbp8e">blind spot</a>.<a class="footnote-ref" id="fnref:53" href="#fn:53"><sup>53</sup></a> </p> <h3>Sex organs</h3> <p>Hydrostatic <a href="/facts/Penis/s9zpC49k">penises</a> have convergently evolved at least six times in male <a href="/facts/Amniote/TeLtDTrJ">amniotes</a>. In these species, males <a href="/facts/Copulation_(zoology)/rYcheMjL">copulate</a> with females and <a href="/facts/Internal_fertilization/CJtuSyeG">internally fertilize</a> their eggs. Similar <a href="/facts/Intromittent_organ/0bhhQ2hY">intromittent organs</a> have evolved in invertebrates such as <a href="/facts/Octopus/z5uokKFF">octopuses</a> and <a href="/facts/Gastropod/rYPqm3PT">gastropods</a>.<a class="footnote-ref" id="fnref:54" href="#fn:54"><sup>54</sup></a> </p> <h3>Flight</h3> <p class="note">Further information: <a href="/facts/Flying_and_gliding_animals/dzTJgy86">Flying and gliding animals § Evolution and ecology of aerial locomotion</a></p> <p><a href="/facts/Birds/RJF6zwLP">Birds</a> and <a href="/facts/Bat/CJudzNt1">bats</a> have <a href="/facts/Homology_(biology)/muctKlyT">homologous</a> limbs because they are both ultimately derived from terrestrial <a href="/facts/Tetrapod/vWpAxzAW">tetrapods</a>, but their flight mechanisms are only analogous, so their wings are examples of functional convergence. The two groups have independently evolved their own means of powered flight. Their wings differ substantially in construction. The bat wing is a membrane stretched across four extremely elongated fingers and the legs. The airfoil of the bird wing is made of <a href="/facts/Feather/sxbf96nF">feathers</a>, strongly attached to the forearm (the ulna) and the highly fused bones of the wrist and hand (the <a href="/facts/Carpometacarpus/rIQfXG0M">carpometacarpus</a>), with only tiny remnants of two fingers remaining, each anchoring a single feather. So, while the wings of bats and birds are functionally convergent, they are not anatomically convergent.<a class="footnote-ref" id="fnref:55" href="#fn:55"><sup>55</sup></a><a class="footnote-ref" id="fnref:56" href="#fn:56"><sup>56</sup></a> Birds and bats also share a high concentration of <a href="/facts/Cerebroside/qYjLH6IP">cerebrosides</a> in the skin of their wings. This improves skin flexibility, a trait useful for flying animals; other mammals have a far lower concentration.<a class="footnote-ref" id="fnref:57" href="#fn:57"><sup>57</sup></a> The extinct <a href="/facts/Pterosaur/YNQwa1ru">pterosaurs</a> independently evolved wings from their fore- and hindlimbs, while <a href="/facts/Insect/IgDNvnUL">insects</a> have <a href="/facts/Insect_wing/e9v05xWY">wings</a> that evolved separately from different organs.<a class="footnote-ref" id="fnref:58" href="#fn:58"><sup>58</sup></a> </p><p><a href="/facts/Flying_squirrel/yTFwGwgc">Flying squirrels</a> and <a href="/facts/Sugar_glider/4YLNQ8Nb">sugar gliders</a> are much alike in their mammalian body plans, with gliding wings stretched between their limbs, but flying squirrels are placentals while sugar gliders are marsupials, widely separated within the mammal lineage from the placentals.<a class="footnote-ref" id="fnref:59" href="#fn:59"><sup>59</sup></a> </p><p><a href="/facts/Hummingbird_hawk-moth/6UAlBLMm">Hummingbird hawk-moths</a> and <a href="/facts/Hummingbird/LfcJhMzm">hummingbirds</a> have evolved similar flight and feeding patterns.<a class="footnote-ref" id="fnref:60" href="#fn:60"><sup>60</sup></a> </p> <h3>Insect mouthparts</h3> <p>Insect mouthparts show many examples of convergent evolution. The mouthparts of different insect groups consist of a set of <a href="/facts/Homology_(biology)/muctKlyT">homologous</a> organs, specialised for the dietary intake of that insect group. Convergent evolution of many groups of insects led from original biting-chewing mouthparts to different, more specialised, derived function types. These include, for example, the <a href="/facts/Proboscis/Ym7nscSW">proboscis</a> of flower-visiting insects such as <a href="/facts/Bee/HglXpXty">bees</a> and <a href="/facts/Flower_beetle/KC5w5A2C">flower beetles</a>,<a class="footnote-ref" id="fnref:61" href="#fn:61"><sup>61</sup></a><a class="footnote-ref" id="fnref:62" href="#fn:62"><sup>62</sup></a><a class="footnote-ref" id="fnref:63" href="#fn:63"><sup>63</sup></a> or the biting-sucking mouthparts of blood-sucking insects such as <a href="/facts/Flea/hFLUFK24">fleas</a> and <a href="/facts/Mosquito/u8hTMTfc">mosquitos</a>. </p> <h3>Opposable thumbs</h3> <p><a href="/facts/Opposable_thumb/vx1ycnbA">Opposable thumbs</a> allowing the grasping of objects are most often associated with <a href="/facts/Primates/Hl8RNE2X">primates</a>, like humans and other apes, monkeys, and lemurs. Opposable thumbs also evolved in <a href="/facts/Giant_pandas/BMbbMGg7">giant pandas</a>, but these are completely different in structure, having six fingers including the thumb, which develops from a wrist bone entirely separately from other fingers.<a class="footnote-ref" id="fnref:64" href="#fn:64"><sup>64</sup></a> </p> <h3>Primates</h3> <p class="note">Further information: <a href="/facts/Human_skin_color/nkMestc6">Human skin color § Genetics of skin color variation</a></p> <table><tbody><tr><td></td><td></td><td></td></tr><tr><td colspan="3"> Despite the similar lightening of <a href="/facts/Human_skin_color/nkMestc6">skin colour</a> after moving <a href="/facts/Out_of_Africa_hypothesis/SJXAFKMA">out of Africa</a>, different genes were involved in European (left) and East Asian (right) lineages.</td></tr></tbody></table> <p>Convergent evolution in humans includes blue eye colour and light skin colour.<a class="footnote-ref" id="fnref:65" href="#fn:65"><sup>65</sup></a> When humans migrated <a href="/facts/Out_of_Africa_hypothesis/SJXAFKMA">out of Africa</a>, they moved to more northern latitudes with less intense sunlight.<a class="footnote-ref" id="fnref:66" href="#fn:66"><sup>66</sup></a> It was beneficial to them to reduce their <a href="/facts/Human_skin_color/nkMestc6">skin pigmentation</a>.<a class="footnote-ref" id="fnref:67" href="#fn:67"><sup>67</sup></a> It appears certain that there was some lightening of skin colour <i>before</i> European and East Asian lineages diverged, as there are some skin-lightening genetic differences that are common to both groups.<a class="footnote-ref" id="fnref:68" href="#fn:68"><sup>68</sup></a> However, after the lineages diverged and became genetically isolated, the skin of both groups lightened more, and that additional lightening was due to <i>different</i> genetic changes.<a class="footnote-ref" id="fnref:69" href="#fn:69"><sup>69</sup></a> </p> <table><tbody><tr><th colspan="2">Humans</th><th colspan="2">Lemurs</th></tr><tr><td></td><td></td><td></td><td></td></tr><tr><td colspan="4">Despite the similarity of appearance, the genetic basis of blue eyes is different in humans and <a href="/facts/Lemur/lgLEmFnr">lemurs</a>.</td></tr></tbody></table> <p><a href="/facts/Lemurs/lgLEmFnr">Lemurs</a> and <a href="/facts/Humans/rlcTENsr">humans</a> are both primates. Ancestral primates had brown eyes, as most primates do today. The genetic basis of blue eyes in humans has been studied in detail and much is known about it. It is not the case that one <a href="/facts/Locus_(genetics)/JcIT8wLm">gene locus</a> is responsible, say with brown dominant to blue <a href="/facts/Eye_colour/50sYzD22">eye colour</a>. However, a single locus is responsible for about 80% of the variation. In lemurs, the differences between blue and brown eyes are not completely known, but the same gene locus is not involved.<a class="footnote-ref" id="fnref:70" href="#fn:70"><sup>70</sup></a></p> <h2 id="in-plants">In plants</h2> <h3>The annual life-cycle</h3> <p>While most plant species are <a href="/facts/Perennial_plant/MQ3NvAUW">perennial</a>, about 6% follow an <a href="/facts/Annual_plant/SkPv72DA">annual</a> life cycle, living for only one growing season.<a class="footnote-ref" id="fnref:71" href="#fn:71"><sup>71</sup></a> The annual life cycle independently emerged in over 120 plant families of angiosperms.<a class="footnote-ref" id="fnref:72" href="#fn:72"><sup>72</sup></a><a class="footnote-ref" id="fnref:73" href="#fn:73"><sup>73</sup></a> The prevalence of annual species increases under hot-dry summer conditions in the four species-rich families of annuals (<a href="/facts/Asteraceae/bouc6v8C">Asteraceae</a>, <a href="/facts/Brassicaceae/xMjNIBzU">Brassicaceae</a>, <a href="/facts/Fabaceae/uxuHNdAr">Fabaceae</a>, and <a href="/facts/Poaceae/q2CRK8q0">Poaceae</a>), indicating that the annual life cycle is adaptive.<a class="footnote-ref" id="fnref:74" href="#fn:74"><sup>74</sup></a><a class="footnote-ref" id="fnref:75" href="#fn:75"><sup>75</sup></a> </p> <h3>Carbon fixation</h3> <p><a href="/facts/C4_photosynthesis/2dLTxsae">C4 photosynthesis</a>, one of the three major carbon-fixing biochemical processes, has <a href="/facts/Evolutionary_history_of_plants/1HfU0VLs">arisen independently up to 40 times</a>.<a class="footnote-ref" id="fnref:76" href="#fn:76"><sup>76</sup></a><a class="footnote-ref" id="fnref:77" href="#fn:77"><sup>77</sup></a> About 7,600 plant species of <a href="/facts/Angiosperm/E5eGYIvY">angiosperms</a> use C4 carbon fixation, with many <a href="/facts/Monocot/mXCuuCYo">monocots</a> including 46% of grasses such as <a href="/facts/Zea_mays/lJ6c5xMH">maize</a> and <a href="/facts/Sugar_cane/925qszH4">sugar cane</a>,<a class="footnote-ref" id="fnref:78" href="#fn:78"><sup>78</sup></a><a class="footnote-ref" id="fnref:79" href="#fn:79"><sup>79</sup></a> and <a href="/facts/Dicot/yat51Ncq">dicots</a> including several species in the <a href="/facts/Chenopodiaceae/8tkMs8jo">Chenopodiaceae</a> and the <a href="/facts/Amaranthaceae/8tkMs8jo">Amaranthaceae</a>.<a class="footnote-ref" id="fnref:80" href="#fn:80"><sup>80</sup></a><a class="footnote-ref" id="fnref:81" href="#fn:81"><sup>81</sup></a> </p> <h3>Fruits</h3> <p><a href="/facts/Fruit/PBpuzk60">Fruits</a> with a wide variety of structural origins have converged to become edible. <a href="/facts/Apple/SF1WND0v">Apples</a> are <a href="/facts/Pome/X5BLQAAp">pomes</a> with five <a href="/facts/Carpel/P5qFQMk1">carpels</a>; their accessory tissues form the apple's core, surrounded by structures from outside the botanical fruit, the <a href="/facts/Receptacle_(botany)/sUVrrdwS">receptacle</a> or <a href="/facts/Hypanthium/mxS9G82W">hypanthium</a>. Other edible fruits include other plant tissues;<a class="footnote-ref" id="fnref:82" href="#fn:82"><sup>82</sup></a> the fleshy part of a <a href="/facts/Tomato/3d5Xsqw7">tomato</a> is the walls of the <a href="/facts/Pericarp/VDLWzDkf">pericarp</a>.<a class="footnote-ref" id="fnref:83" href="#fn:83"><sup>83</sup></a> This implies convergent evolution under selective pressure, in this case the competition for <a href="/facts/Seed_dispersal/yNofEeyA">seed dispersal</a> by animals through consumption of fleshy fruits.<a class="footnote-ref" id="fnref:84" href="#fn:84"><sup>84</sup></a> </p><p>Seed dispersal by ants (<a href="/facts/Myrmecochory/3rcqJHwv">myrmecochory</a>) has evolved independently more than 100 times, and is present in more than 11,000 plant species. It is one of the most dramatic examples of convergent evolution in biology.<a class="footnote-ref" id="fnref:85" href="#fn:85"><sup>85</sup></a> </p> <h3>Carnivory</h3> <p><a href="/facts/Carnivorous_plant/GWTld9Ke">Carnivory</a> has evolved multiple times independently in plants in widely separated groups. In three species studied, <i><a href="/facts/Cephalotus/HBbckSwz">Cephalotus follicularis</a></i>, <i><a href="/facts/Nepenthes_alata/qduuBak1">Nepenthes alata</a></i> and <i><a href="/facts/Sarracenia_purpurea/sSpMUwX8">Sarracenia purpurea</a></i>, there has been convergence at the molecular level. Carnivorous plants secrete <a href="/facts/Enzymes/DSI1Fh7y">enzymes</a> into the digestive fluid they produce. By studying <a href="/facts/Purple_acid_phosphatases/eN1WBQPq">phosphatase</a>, <a href="/facts/Glycoside_hydrolase_family_19/n6VUTmud">glycoside hydrolase</a>, <a href="/facts/Glucanase/YlteKvBn">glucanase</a>, <a href="/facts/RNASET2/QLxKHsND">RNAse</a> and <a href="/facts/Chitinase/tAmj0pB5">chitinase</a> <a href="/facts/Enzyme/DSI1Fh7y">enzymes</a> as well as a <a href="/facts/Pathogenesis-related_protein/vCT1VSvk">pathogenesis-related protein</a> and a <a href="/facts/Thaumatin/ytpkDdUs">thaumatin</a>-related protein, the authors found many convergent <a href="/facts/Amino_acid/WDxYwBny">amino acid</a> substitutions. These changes were not at the enzymes' catalytic sites, but rather on the exposed surfaces of the proteins, where they might interact with other components of the cell or the digestive fluid. The authors also found that <a href="/facts/Homologous_gene/mCIVStrA">homologous genes</a> in the non-carnivorous plant <i><a href="/facts/Arabidopsis_thaliana/wt2IMJsC">Arabidopsis thaliana</a></i> tend to have their expression increased when the plant is stressed, leading the authors to suggest that stress-responsive proteins have often been co-opted<a class="footnote-ref" id="fnref:86" href="#fn:86"><sup>86</sup></a> in the repeated evolution of carnivory.<a class="footnote-ref" id="fnref:87" href="#fn:87"><sup>87</sup></a> </p> <h2 id="methods-of-inference">Methods of inference</h2> <p>Phylogenetic reconstruction and <a href="/facts/Ancestral_reconstruction/a7m8yX7C">ancestral state reconstruction</a> proceed by assuming that evolution has occurred without convergence. Convergent patterns may, however, appear at higher levels in a phylogenetic reconstruction, and are sometimes explicitly sought by investigators. The methods applied to infer convergent evolution depend on whether pattern-based or process-based convergence is expected. Pattern-based convergence is the broader term, for when two or more lineages independently evolve patterns of similar traits. Process-based convergence is when the convergence is due to similar forces of <a href="/facts/Natural_selection/JXTte6en">natural selection</a>.<a class="footnote-ref" id="fnref:88" href="#fn:88"><sup>88</sup></a> </p> <h3>Pattern-based measures</h3> <p>Earlier methods for measuring convergence incorporate ratios of phenotypic and <a href="/facts/Phylogenetic/bw4rsovt">phylogenetic</a> distance by simulating evolution with a <a href="/facts/Brownian_motion/Th0g0wkr">Brownian motion</a> model of trait evolution along a phylogeny.<a class="footnote-ref" id="fnref:89" href="#fn:89"><sup>89</sup></a><a class="footnote-ref" id="fnref:90" href="#fn:90"><sup>90</sup></a> More recent methods also quantify the strength of convergence.<a class="footnote-ref" id="fnref:91" href="#fn:91"><sup>91</sup></a> One drawback to keep in mind is that these methods can confuse long-term stasis with convergence due to phenotypic similarities. Stasis occurs when there is little evolutionary change among taxa.<a class="footnote-ref" id="fnref:92" href="#fn:92"><sup>92</sup></a> </p><p>Distance-based measures assess the degree of similarity between lineages over time. Frequency-based measures assess the number of lineages that have evolved in a particular trait space.<a class="footnote-ref" id="fnref:93" href="#fn:93"><sup>93</sup></a> </p> <h3>Process-based measures</h3> <p>Methods to infer process-based convergence fit models of selection to a phylogeny and continuous trait data to determine whether the same selective forces have acted upon lineages. This uses the <a href="/facts/Ornstein%E2%80%93Uhlenbeck_process/oFaDcovm">Ornstein–Uhlenbeck process</a> to test different scenarios of selection. Other methods rely on an <i><a href="/facts/A_priori_knowledge/C567SkcV">a priori</a></i> specification of where shifts in selection have occurred.<a class="footnote-ref" id="fnref:94" href="#fn:94"><sup>94</sup></a> </p> <h2 id="see-also">See also</h2> <ul><li><a href="/facts/Incomplete_lineage_sorting/Nh3x93Xy">Incomplete lineage sorting</a> – Characteristic of phylogenetic analysis: the presence of multiple alleles in ancestral populations might lead to the impression that convergent evolution has occurred.</li> <li><a href="/facts/Carcinisation/nrr183QG">Carcinisation</a> – Evolution of crustaceans into crab-like forms</li> <li><a href="/facts/Morphology_(biology)/Nl3B0myy">Morphology (biology)</a> – Study of external forms and structures of organisms</li> <li><a href="/facts/Iterative_evolution/vgHRND7m">Iterative evolution</a> – The repeated evolution of a specific trait or body plan from the same ancestral lineage at different points in time.</li> <li><a href="/facts/Elvis_taxon/jCctHR0S">Elvis taxon</a> – Misidentification of later taxon superficially resembling earlier extinct taxon</li> <li><a href="/facts/Breeding_back/d0QT01n0">Breeding back</a> – A form of selective breeding to recreate the traits of an extinct species, but the genome will differ from the original species.</li> <li><a href="/facts/Orthogenesis/TKjwUPmA">Orthogenesis</a> (contrastable with convergent evolution; involves teleology)</li> <li><a href="/facts/Contingency_(evolutionary_biology)/3S118ARo">Contingency (evolutionary biology)</a> – effect of evolutionary history on outcomes</li></ul> <h2 id="notes">Notes</h2> <h2 id="further-reading">Further reading</h2> <ul><li>Losos, Jonathan B. (2017). <i>Improbable Destinies: Fate, Chance, and the Future of Evolution</i>. Riverhead Books. <a href="/facts/ISBN_(identifier)/15AdSPa9">ISBN</a> 978-0399184925.</li></ul> <h2 id="external-links">External links</h2> <ul><li> Media related to Convergent evolution at Wikimedia Commons</li></ul> <h2 id="references">References</h2> <ol> <li id="fn:1"><p>Kirk, John Thomas Osmond (2007). Science & Certainty. Csiro Publishing. p. 79. ISBN 978-0-643-09391-1. Archived from the original on 15 February 2017. Retrieved 23 January 2017. evolutionary convergence, which, quoting .. Simon Conway Morris .. is the 'recurring tendency of biological organization to arrive at the same "solution" to a particular "need". .. the 'Tasmanian tiger' .. looked and behaved like a wolf and occupied a similar ecological niche, but was in fact a marsupial not a placental mammal. <a href="978-0-643-09391-1" target="_blank">978-0-643-09391-1</a> <a href="#fnref:1" class="footnote-back-ref">↩</a></p></li> <li id="fn:2"><p>Reece, J.; Meyers, N.; Urry, L.; Cain, M.; Wasserman, S.; Minorsky, P.; Jackson, R.; Cooke, B. (5 September 2011). Cambell Biology, 9th Edition. Pearson. p. 586. ISBN 978-1-4425-3176-5. <a href="978-1-4425-3176-5" target="_blank">978-1-4425-3176-5</a> <a href="#fnref:2" class="footnote-back-ref">↩</a></p></li> <li id="fn:3"><p>"Homologies and analogies". University of California Berkeley. Archived from the original on 19 November 2016. Retrieved 10 January 2017. <a href="http://evolution.berkeley.edu/evolibrary/article/evo_09" target="_blank">http://evolution.berkeley.edu/evolibrary/article/evo_09</a> <a href="#fnref:3" class="footnote-back-ref">↩</a></p></li> <li id="fn:4"><p>Thunstad, Erik (2009). Darwins teori, evolusjon gjennom 400 år (in Norwegian). Oslo, Norway: Humanist forlag. p. 404. ISBN 978-82-92622-53-7. <a href="978-82-92622-53-7" target="_blank">978-82-92622-53-7</a> <a href="#fnref:4" class="footnote-back-ref">↩</a></p></li> <li id="fn:5"><p>Buller, A. R.; Townsend, C. A. (19 February 2013). "Intrinsic evolutionary constraints on protease structure, enzyme acylation, and the identity of the catalytic triad". Proceedings of the National Academy of Sciences of the United States of America. 110 (8): E653–61. Bibcode:2013PNAS..110E.653B. doi:10.1073/pnas.1221050110. PMC 3581919. PMID 23382230. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3581919" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3581919</a> <a href="#fnref:5" class="footnote-back-ref">↩</a></p></li> <li id="fn:6"><p>Gould, S.J. (1989). Wonderful Life: The Burgess Shale and the Nature of History. W.W. Norton. pp. 282–285. ISBN 978-0-09-174271-3. <a href="978-0-09-174271-3" target="_blank">978-0-09-174271-3</a> <a href="#fnref:6" class="footnote-back-ref">↩</a></p></li> <li id="fn:7"><p>Conway Morris, Simon (2005). Life's solution: inevitable humans in a lonely universe. Cambridge University Press. pp. 164, 167, 170 and 235. ISBN 978-0-521-60325-6. OCLC 156902715. <a href="978-0-521-60325-6" target="_blank">978-0-521-60325-6</a> <a href="#fnref:7" class="footnote-back-ref">↩</a></p></li> <li id="fn:8"><p>Chirat, R.; Moulton, D. E.; Goriely, A. (2013). "Mechanical basis of morphogenesis and convergent evolution of spiny seashells". Proceedings of the National Academy of Sciences. 110 (15): 6015–6020. Bibcode:2013PNAS..110.6015C. doi:10.1073/pnas.1220443110. PMC 3625336. PMID 23530223. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3625336" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3625336</a> <a href="#fnref:8" class="footnote-back-ref">↩</a></p></li> <li id="fn:9"><p>Lomolino, M; Riddle, B; Whittaker, R; Brown, J (2010). Biogeography, Fourth Edition. Sinauer Associates. p. 426. ISBN 978-0-87893-494-2. <a href="978-0-87893-494-2" target="_blank">978-0-87893-494-2</a> <a href="#fnref:9" class="footnote-back-ref">↩</a></p></li> <li id="fn:10"><p>West-Eberhard, Mary Jane (2003). Developmental Plasticity and Evolution. Oxford University Press. pp. 353–376. ISBN 978-0-19-512235-0. <a href="978-0-19-512235-0" target="_blank">978-0-19-512235-0</a> <a href="#fnref:10" class="footnote-back-ref">↩</a></p></li> <li id="fn:11"><p>Sanderson, Michael J.; Hufford, Larry (1996). Homoplasy: The Recurrence of Similarity in Evolution. Academic Press. pp. 330, and passim. ISBN 978-0-08-053411-4. Archived from the original on 14 February 2017. Retrieved 21 January 2017. <a href="978-0-08-053411-4" target="_blank">978-0-08-053411-4</a> <a href="#fnref:11" class="footnote-back-ref">↩</a></p></li> <li id="fn:12"><p>Collin, R.; Cipriani, R. (2003). "Dollo's law and the re-evolution of shell coiling". Proceedings of the Royal Society B. 270 (1533): 2551–2555. doi:10.1098/rspb.2003.2517. PMC 1691546. 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