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<h1 id="firstHeading" class="firstHeading" lang="en">Fungus</h1>
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<div id="siteSub" class="noprint">From Wikipedia, the free encyclopedia</div>
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<div role="note" class="hatnote navigation-not-searchable">For the US radio station, see <a href="/wiki/Fungus_(XM)" title="Fungus (XM)">Fungus (XM)</a>.</div>
<div role="note" class="hatnote navigation-not-searchable">"Fungi" redirects here. For other uses, see <a href="/wiki/Fungi_(disambiguation)" class="mw-disambig" title="Fungi (disambiguation)">Fungi (disambiguation)</a>.</div>
<p class="mw-empty-elt"></p>
<div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Any member of the eukaryotic kingdom that includes organisms such as yeasts, molds and mushrooms</div>
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<table class="infobox biota" style="text-align: left; width: 200px; font-size: 100%">
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<th colspan="2" style="text-align: center; background-color: rgb(145,250,250)">
Fungi<br />
<small>
Temporal range: Early <a href="/wiki/Devonian" title="Devonian">Devonian</a>–<a href="/wiki/Holocene" title="Holocene">Present</a> (but <a href="#Evolution">see text</a>) <span style="display:inline-block;"></span><span style="display:inline-block;">410–0 <a href="/wiki/Megaannum" class="mw-redirect" title="Megaannum">Ma</a></span> <span style="display:inline-block;"></span>
<div id="Timeline-row" style="margin: 4px auto 0; clear:both; width:220px; padding:0px; height:18px; overflow:visible; border:1px #666; border-style:solid none; position:relative; z-index:0; font-size:13px;">
<div style="position:absolute; height:100%; left:0px; width:208.07692307692px; padding-left:5px; text-align:left; background-color:rgb(254,214,123); background-image: -moz-linear-gradient(left, rgba(255,255,255,1), rgba(254,217,106,1) 15%, rgba(254,217,106,1)); background-image: -o-linear-gradient(left, rgba(255,255,255,1), rgba(254,217,106,1) 15%, rgba(254,217,106,1)); background-image: -webkit-linear-gradient(left, rgba(255,255,255,1), rgba(254,217,106,1) 15%, rgba(254,217,106,1)); background-image: linear-gradient(to right, rgba(255,255,255,1), rgba(254,217,106,1) 15%, rgba(254,217,106,1));"><a href="/wiki/Precambrian" title="Precambrian">PreЄ</a></div>
<div style="position:absolute; height:100%; text-align:center; background-color:rgb(129,170,114); left:36.892307692308px; width:18.818461538462px;"><a href="/wiki/Cambrian" title="Cambrian">Є</a></div>
<div style="position:absolute; height:100%; text-align:center; background-color:rgb(0,169,138); left:55.710769230769px; width:14.08px;"><a href="/wiki/Ordovician" title="Ordovician">O</a></div>
<div style="position:absolute; height:100%; text-align:center; background-color:rgb(166,223,197); left:69.790769230769px; width:8.3261538461539px;"><a href="/wiki/Silurian" title="Silurian">S</a></div>
<div style="position:absolute; height:100%; text-align:center; background-color:rgb(221,150,81); left:78.116923076923px; width:20.409230769231px;"><a href="/wiki/Devonian" title="Devonian">D</a></div>
<div style="position:absolute; height:100%; text-align:center; background-color:rgb(63,174,173); left:98.526153846154px; width:20.307692307692px;"><a href="/wiki/Carboniferous" title="Carboniferous">C</a></div>
<div style="position:absolute; height:100%; text-align:center; background-color:rgb(247,88,60); left:118.83384615385px; width:15.907015384615px;"><a href="/wiki/Permian" title="Permian">P</a></div>
<div style="position:absolute; height:100%; text-align:center; background-color:rgb(153,78,150); left:134.74086153846px; width:17.126830769231px;"><a href="/wiki/Triassic" title="Triassic">T</a></div>
<div style="position:absolute; height:100%; text-align:center; background-color:rgb(0,187,231); left:151.86769230769px; width:19.055384615385px;"><a href="/wiki/Jurassic" title="Jurassic">J</a></div>
<div style="position:absolute; height:100%; text-align:center; background-color:rgb(111,200,107); left:170.92307692308px; width:26.738461538462px;"><a href="/wiki/Cretaceous" title="Cretaceous">K</a></div>
<div style="position:absolute; height:100%; text-align:center; background-color:rgb(254,161,99); left:197.66153846154px; width:14.543692307692px;"><a href="/wiki/Paleogene" title="Paleogene"><small>Pg</small></a></div>
<div style="position:absolute; height:100%; text-align:center; background-color:rgb(254,221,45); left:212.20523076923px; width:6.9215384615385px;"><a href="/wiki/Neogene" title="Neogene"><small>N</small></a></div>
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</div>
</small>
</th>
</tr>
<tr>
<td colspan="2" style="text-align: center"><a href="/wiki/File:Fungi_collage.jpg" class="image" title="A collage of five fungi (clockwise from top-left): a mushroom with a flat, red top with white-spots, and a white stem growing on the ground; a red cup-shaped fungus growing on wood; a stack of green and white moldy bread slices on a plate; a microscopic, spherical grey semitransparent cell, with a smaller spherical cell beside it; a microscopic view of an elongated cellular structure shaped like a microphone, attached to the larger end is a number of smaller roughly circular elements that collectively form a mass around it"><img alt="A collage of five fungi (clockwise from top-left): a mushroom with a flat, red top with white-spots, and a white stem growing on the ground; a red cup-shaped fungus growing on wood; a stack of green and white moldy bread slices on a plate; a microscopic, spherical grey semitransparent cell, with a smaller spherical cell beside it; a microscopic view of an elongated cellular structure shaped like a microphone, attached to the larger end is a number of smaller roughly circular elements that collectively form a mass around it" src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fc/Fungi_collage.jpg/290px-Fungi_collage.jpg" width="290" height="252" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fc/Fungi_collage.jpg/435px-Fungi_collage.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fc/Fungi_collage.jpg/580px-Fungi_collage.jpg 2x" data-file-width="1220" data-file-height="1062" /></a></td>
</tr>
<tr>
<td colspan="2" style="text-align: center; font-size: 88%">
Clockwise from top left:
<div class="plainlist">
<ul>
<li><i><a href="/wiki/Amanita_muscaria" title="Amanita muscaria">Amanita muscaria</a></i>, a basidiomycete;</li>
<li><i><a href="/wiki/Sarcoscypha_coccinea" title="Sarcoscypha coccinea">Sarcoscypha coccinea</a></i>, an ascomycete;</li>
<li>bread covered in <a href="/wiki/Mold" title="Mold">mold</a>;</li>
<li>a <a href="/wiki/Chytridiomycota" title="Chytridiomycota">chytrid</a>;</li>
<li>an <i><a href="/wiki/Aspergillus" title="Aspergillus">Aspergillus</a></i> <a href="/wiki/Conidiophore" class="mw-redirect" title="Conidiophore">conidiophore</a>.</li>
</ul>
</div>
</td>
</tr>
<tr>
<th colspan="2" style="min-width:15em; text-align: center; background-color: rgb(145,250,250)"><a href="/wiki/Taxonomy_(biology)" title="Taxonomy (biology)">Scientific classification</a> <span class="plainlinks" style="font-size:smaller; float:right; padding-right:0.4em; margin-left:-3em;"><a href="/wiki/Template:Taxonomy/Fungi" title="e"><img alt="e" src="//upload.wikimedia.org/wikipedia/commons/7/74/Red_Pencil_Icon.png" width="16" height="16" data-file-width="16" data-file-height="16" /></a></span></th>
</tr>
<tr>
<td>(unranked):</td>
<td><a href="/wiki/Opisthokont" title="Opisthokont">Opisthokonta</a></td>
</tr>
<tr>
<td>(unranked):</td>
<td><a href="/wiki/Holomycota" title="Holomycota">Holomycota</a></td>
</tr>
<tr>
<td>(unranked):</td>
<td><a href="/wiki/Zoosporia" class="mw-redirect" title="Zoosporia">Zoosporia</a></td>
</tr>
<tr>
<td>Kingdom:</td>
<td><b>Fungi</b><br /><small>(<a href="/wiki/Carl_Linnaeus" title="Carl Linnaeus">L.</a>) <a href="/wiki/Royall_T._Moore" title="Royall T. Moore">R.T.Moore</a><sup id="cite_ref-Moore1980_1-0" class="reference"><a href="#cite_note-Moore1980-1">[1]</a></sup></small></td>
</tr>
<tr>
<th colspan="2" style="text-align: center; background-color: rgb(145,250,250)">Subkingdoms/Phyla/Subphyla<sup id="cite_ref-2" class="reference"><a href="#cite_note-2">[2]</a></sup></th>
</tr>
<tr>
<td colspan="2" style="text-align: left">
<dl>
<dd><a href="/wiki/Blastocladiomycota" title="Blastocladiomycota">Blastocladiomycota</a></dd>
<dd><a href="/wiki/Chytridiomycota" title="Chytridiomycota">Chytridiomycota</a></dd>
<dd><a href="/wiki/Glomeromycota" title="Glomeromycota">Glomeromycota</a></dd>
<dd><a href="/wiki/Microsporidia" title="Microsporidia">Microsporidia</a></dd>
<dd><a href="/wiki/Neocallimastigomycota" title="Neocallimastigomycota">Neocallimastigomycota</a></dd>
</dl>
<p><a href="/wiki/Dikarya" title="Dikarya">Dikarya</a> (inc. <a href="/wiki/Deuteromycota" class="mw-redirect" title="Deuteromycota">Deuteromycota</a>)</p>
<dl>
<dd>
<a href="/wiki/Ascomycota" title="Ascomycota">Ascomycota</a>
<dl>
<dd><a href="/wiki/Pezizomycotina" title="Pezizomycotina">Pezizomycotina</a></dd>
<dd><a href="/wiki/Saccharomycotina" title="Saccharomycotina">Saccharomycotina</a></dd>
<dd><a href="/wiki/Taphrinomycotina" title="Taphrinomycotina">Taphrinomycotina</a></dd>
</dl>
</dd>
<dd>
<a href="/wiki/Basidiomycota" title="Basidiomycota">Basidiomycota</a>
<dl>
<dd><a href="/wiki/Agaricomycotina" title="Agaricomycotina">Agaricomycotina</a></dd>
<dd><a href="/wiki/Pucciniomycotina" title="Pucciniomycotina">Pucciniomycotina</a></dd>
<dd><a href="/wiki/Ustilaginomycotina" title="Ustilaginomycotina">Ustilaginomycotina</a></dd>
</dl>
</dd>
</dl>
<p>Subphyla <i><a href="/wiki/Incertae_sedis" title="Incertae sedis">incertae sedis</a></i></p>
<dl>
<dd><a href="/wiki/Entomophthoromycotina" class="mw-redirect" title="Entomophthoromycotina">Entomophthoromycotina</a></dd>
<dd><a href="/wiki/Kickxellomycotina" title="Kickxellomycotina">Kickxellomycotina</a></dd>
<dd><a href="/wiki/Mucoromycotina" title="Mucoromycotina">Mucoromycotina</a></dd>
<dd><a href="/wiki/Zoopagomycotina" title="Zoopagomycotina">Zoopagomycotina</a></dd>
</dl>
</td>
</tr>
</tbody>
</table>
<p>A <b>fungus</b> (<a href="/wiki/Plural" title="Plural">plural</a>: <b>fungi</b><sup id="cite_ref-3" class="reference"><a href="#cite_note-3">[3]</a></sup> or <b>funguses</b><sup id="cite_ref-OxfordDictionary_4-0" class="reference"><a href="#cite_note-OxfordDictionary-4">[4]</a></sup>) is any member of the group of <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotic</a> organisms that includes microorganisms such as <a href="/wiki/Yeast" title="Yeast">yeasts</a> and <a href="/wiki/Mold" title="Mold">molds</a>, as well as the more familiar <a href="/wiki/Mushroom" title="Mushroom">mushrooms</a>. These organisms are classified as a <a href="/wiki/Kingdom_(biology)" title="Kingdom (biology)">kingdom</a>, <b>Fungi</b>, which is separate from the other eukaryotic life kingdoms of <a href="/wiki/Plant" title="Plant">plants</a> and <a href="/wiki/Animal" title="Animal">animals</a>.</p>
<p>A characteristic that places fungi in a different kingdom from plants, bacteria, and some protists is <a href="/wiki/Chitin" title="Chitin">chitin</a> in their <a href="/wiki/Cell_wall" title="Cell wall">cell walls</a>. Similar to animals, fungi are <a href="/wiki/Heterotroph" title="Heterotroph">heterotrophs</a>; they acquire their food by absorbing dissolved molecules, typically by secreting <a href="/wiki/Digestive_enzyme" title="Digestive enzyme">digestive enzymes</a> into their environment. Fungi do not <a href="/wiki/Photosynthesis" title="Photosynthesis">photosynthesise</a>. Growth is their means of mobility, except for spores (a few of which are flagellated), which may travel through the air or water. Fungi are the principal decomposers in ecological systems. These and other differences place fungi in a single group of related organisms, named the <i>Eumycota</i> (<i>true fungi</i> or <i>Eumycetes</i>), which share a <a href="/wiki/Common_ancestor" class="mw-redirect" title="Common ancestor">common ancestor</a> (form a <i><a href="/wiki/Monophyletic" class="mw-redirect" title="Monophyletic">monophyletic</a> group</i>), an interpretation that is also strongly supported by <a href="/wiki/Molecular_phylogenetics" title="Molecular phylogenetics">molecular phylogenetics</a>. This fungal group is distinct from the structurally similar <a href="/wiki/Mycetozoa" title="Mycetozoa">myxomycetes</a> (slime molds) and <a href="/wiki/Oomycete" title="Oomycete">oomycetes</a> (water molds). The discipline of <a href="/wiki/Biology" title="Biology">biology</a> devoted to the study of fungi is known as <a href="/wiki/Mycology" title="Mycology">mycology</a> (from the <a href="/wiki/Greek_language" title="Greek language">Greek</a> μύκης <i>mykes</i>, mushroom). In the past, mycology was regarded as a branch of <a href="/wiki/Botany" title="Botany">botany</a>, although it is now known fungi are genetically more closely related to animals than to plants.</p>
<p>Abundant worldwide, most fungi are inconspicuous because of the small size of their structures, and their <a href="/wiki/Crypsis" title="Crypsis">cryptic</a> lifestyles in soil or on dead matter. Fungi include <a href="/wiki/Symbiosis" title="Symbiosis">symbionts</a> of plants, animals, or other fungi and also <a href="/wiki/Parasites" class="mw-redirect" title="Parasites">parasites</a>. They may become noticeable when <a href="/wiki/Sporocarp_(fungi)" title="Sporocarp (fungi)">fruiting</a>, either as mushrooms or as molds. Fungi perform an essential role in the decomposition of organic matter and have fundamental roles in nutrient <a href="/wiki/Biogeochemical_cycle" title="Biogeochemical cycle">cycling</a> and exchange in the environment. They have long been used as a direct source of human food, in the form of mushrooms and <a href="/wiki/Tuber_(genus)" class="mw-redirect" title="Tuber (genus)">truffles</a>; as a <a href="/wiki/Bread#Leavening" title="Bread">leavening</a> agent for bread; and in the <a href="/wiki/Fermentation_(food)" class="mw-redirect" title="Fermentation (food)">fermentation</a> of various food products, such as <a href="/wiki/Wine" title="Wine">wine</a>, <a href="/wiki/Beer" title="Beer">beer</a>, and <a href="/wiki/Soy_sauce" title="Soy sauce">soy sauce</a>. Since the 1940s, fungi have been used for the production of <a href="/wiki/Antibiotic" title="Antibiotic">antibiotics</a>, and, more recently, various <a href="/wiki/Enzyme" title="Enzyme">enzymes</a> produced by fungi are used <a href="/wiki/Enzyme#Industrial_applications" title="Enzyme">industrially</a> and in <a href="/wiki/Protease#Occurrence" title="Protease">detergents</a>. Fungi are also used as <a href="/wiki/Biological_pesticide" class="mw-redirect" title="Biological pesticide">biological pesticides</a> to control weeds, plant diseases and insect pests. Many species produce <a href="/wiki/Bioactive_compound" class="mw-redirect" title="Bioactive compound">bioactive</a> compounds called <a href="/wiki/Mycotoxin" title="Mycotoxin">mycotoxins</a>, such as <a href="/wiki/Alkaloid" title="Alkaloid">alkaloids</a> and <a href="/wiki/Polyketide" title="Polyketide">polyketides</a>, that are toxic to animals including humans. The fruiting structures of a few species contain <a href="/wiki/Psychotropic" class="mw-redirect" title="Psychotropic">psychotropic</a> compounds and are consumed <a href="/wiki/Recreational_drug_use" title="Recreational drug use">recreationally</a> or in traditional <a href="/wiki/Entheogen" title="Entheogen">spiritual ceremonies</a>. Fungi can break down manufactured materials and buildings, and become significant <a href="/wiki/Pathogenic_fungi" class="mw-redirect" title="Pathogenic fungi">pathogens</a> of humans and other animals. Losses of crops due to fungal diseases (e.g., <a href="/wiki/Rice_blast_disease" class="mw-redirect" title="Rice blast disease">rice blast disease</a>) or food <a href="/wiki/Food_spoilage" title="Food spoilage">spoilage</a> can have a large impact on human <a href="/wiki/Food_security" title="Food security">food supplies</a> and local economies.</p>
<p>The fungus kingdom encompasses an enormous diversity of <a href="/wiki/Taxon" title="Taxon">taxa</a> with varied ecologies, <a href="/wiki/Biological_life_cycle" title="Biological life cycle">life cycle</a> strategies, and <a href="/wiki/Morphology_(biology)" title="Morphology (biology)">morphologies</a> ranging from unicellular aquatic <a href="/wiki/Chytrid" class="mw-redirect" title="Chytrid">chytrids</a> to large mushrooms. However, little is known of the true <a href="/wiki/Biodiversity" title="Biodiversity">biodiversity</a> of Kingdom Fungi, which has been estimated at 2.2 million to 3.8 million species.<sup id="cite_ref-Lucking2017_5-0" class="reference"><a href="#cite_note-Lucking2017-5">[5]</a></sup> Of these, only about 120,000 have been described, with over 8,000 species known to be detrimental to plants and at least 300 that can be pathogenic to humans.<sup id="cite_ref-6" class="reference"><a href="#cite_note-6">[6]</a></sup> Ever since the pioneering 18th and 19th century <a href="/wiki/Taxonomy_(biology)" title="Taxonomy (biology)">taxonomical</a> works of <a href="/wiki/Carl_Linnaeus" title="Carl Linnaeus">Carl Linnaeus</a>, <a href="/wiki/Christian_Hendrik_Persoon" class="mw-redirect" title="Christian Hendrik Persoon">Christian Hendrik Persoon</a>, and <a href="/wiki/Elias_Magnus_Fries" title="Elias Magnus Fries">Elias Magnus Fries</a>, fungi have been <a href="/wiki/Biological_classification" class="mw-redirect" title="Biological classification">classified</a> according to their morphology (e.g., characteristics such as spore color or microscopic features) or <a href="/wiki/Physiology" title="Physiology">physiology</a>. Advances in <a href="/wiki/Molecular_genetics" title="Molecular genetics">molecular genetics</a> have opened the way for <a href="/wiki/DNA_sequencing" title="DNA sequencing">DNA analysis</a> to be incorporated into taxonomy, which has sometimes challenged the historical groupings based on morphology and other traits. <a href="/wiki/Phylogenetic" class="mw-redirect" title="Phylogenetic">Phylogenetic</a> studies published in the last decade have helped reshape the classification within Kingdom Fungi, which is divided into one <a href="/wiki/Kingdom_(biology)#Definition_and_associated_terms" title="Kingdom (biology)">subkingdom</a>, seven <a href="/wiki/Phylum" title="Phylum">phyla</a>, and ten subphyla.</p>
<div id="toc" class="toc">
<input type="checkbox" role="button" id="toctogglecheckbox" class="toctogglecheckbox" style="display:none" />
<div class="toctitle" lang="en" dir="ltr">
<h2>Contents</h2>
<span class="toctogglespan"><label class="toctogglelabel" for="toctogglecheckbox"></label></span>
</div>
<ul>
<li class="toclevel-1 tocsection-1"><a href="#Etymology"><span class="tocnumber">1</span> <span class="toctext">Etymology</span></a></li>
<li class="toclevel-1 tocsection-2"><a href="#Characteristics"><span class="tocnumber">2</span> <span class="toctext">Characteristics</span></a></li>
<li class="toclevel-1 tocsection-3"><a href="#Diversity"><span class="tocnumber">3</span> <span class="toctext">Diversity</span></a></li>
<li class="toclevel-1 tocsection-4">
<a href="#Mycology"><span class="tocnumber">4</span> <span class="toctext">Mycology</span></a>
<ul>
<li class="toclevel-2 tocsection-5"><a href="#History"><span class="tocnumber">4.1</span> <span class="toctext">History</span></a></li>
</ul>
</li>
<li class="toclevel-1 tocsection-6">
<a href="#Morphology"><span class="tocnumber">5</span> <span class="toctext">Morphology</span></a>
<ul>
<li class="toclevel-2 tocsection-7"><a href="#Microscopic_structures"><span class="tocnumber">5.1</span> <span class="toctext">Microscopic structures</span></a></li>
<li class="toclevel-2 tocsection-8"><a href="#Macroscopic_structures"><span class="tocnumber">5.2</span> <span class="toctext">Macroscopic structures</span></a></li>
</ul>
</li>
<li class="toclevel-1 tocsection-9"><a href="#Growth_and_physiology"><span class="tocnumber">6</span> <span class="toctext">Growth and physiology</span></a></li>
<li class="toclevel-1 tocsection-10">
<a href="#Reproduction"><span class="tocnumber">7</span> <span class="toctext">Reproduction</span></a>
<ul>
<li class="toclevel-2 tocsection-11"><a href="#Asexual_reproduction"><span class="tocnumber">7.1</span> <span class="toctext">Asexual reproduction</span></a></li>
<li class="toclevel-2 tocsection-12"><a href="#Sexual_reproduction"><span class="tocnumber">7.2</span> <span class="toctext">Sexual reproduction</span></a></li>
<li class="toclevel-2 tocsection-13"><a href="#Spore_dispersal"><span class="tocnumber">7.3</span> <span class="toctext">Spore dispersal</span></a></li>
<li class="toclevel-2 tocsection-14"><a href="#Other_sexual_processes"><span class="tocnumber">7.4</span> <span class="toctext">Other sexual processes</span></a></li>
</ul>
</li>
<li class="toclevel-1 tocsection-15"><a href="#Evolution"><span class="tocnumber">8</span> <span class="toctext">Evolution</span></a></li>
<li class="toclevel-1 tocsection-16">
<a href="#Taxonomy"><span class="tocnumber">9</span> <span class="toctext">Taxonomy</span></a>
<ul>
<li class="toclevel-2 tocsection-17"><a href="#Taxonomic_groups"><span class="tocnumber">9.1</span> <span class="toctext">Taxonomic groups</span></a></li>
<li class="toclevel-2 tocsection-18"><a href="#Fungus-like_organisms"><span class="tocnumber">9.2</span> <span class="toctext">Fungus-like organisms</span></a></li>
</ul>
</li>
<li class="toclevel-1 tocsection-19">
<a href="#Ecology"><span class="tocnumber">10</span> <span class="toctext">Ecology</span></a>
<ul>
<li class="toclevel-2 tocsection-20">
<a href="#Symbiosis"><span class="tocnumber">10.1</span> <span class="toctext">Symbiosis</span></a>
<ul>
<li class="toclevel-3 tocsection-21"><a href="#With_plants"><span class="tocnumber">10.1.1</span> <span class="toctext">With plants</span></a></li>
<li class="toclevel-3 tocsection-22"><a href="#With_algae_and_cyanobacteria"><span class="tocnumber">10.1.2</span> <span class="toctext">With algae and cyanobacteria</span></a></li>
<li class="toclevel-3 tocsection-23"><a href="#With_insects"><span class="tocnumber">10.1.3</span> <span class="toctext">With insects</span></a></li>
<li class="toclevel-3 tocsection-24"><a href="#As_pathogens_and_parasites"><span class="tocnumber">10.1.4</span> <span class="toctext">As pathogens and parasites</span></a></li>
<li class="toclevel-3 tocsection-25"><a href="#As_targets_of_mycoparasites"><span class="tocnumber">10.1.5</span> <span class="toctext">As targets of mycoparasites</span></a></li>
</ul>
</li>
</ul>
</li>
<li class="toclevel-1 tocsection-26"><a href="#Mycotoxins"><span class="tocnumber">11</span> <span class="toctext">Mycotoxins</span></a></li>
<li class="toclevel-1 tocsection-27"><a href="#Pathogenic_mechanisms"><span class="tocnumber">12</span> <span class="toctext">Pathogenic mechanisms</span></a></li>
<li class="toclevel-1 tocsection-28">
<a href="#Human_use"><span class="tocnumber">13</span> <span class="toctext">Human use</span></a>
<ul>
<li class="toclevel-2 tocsection-29">
<a href="#Therapeutic_uses"><span class="tocnumber">13.1</span> <span class="toctext">Therapeutic uses</span></a>
<ul>
<li class="toclevel-3 tocsection-30"><a href="#Modern_chemotherapeutics"><span class="tocnumber">13.1.1</span> <span class="toctext">Modern chemotherapeutics</span></a></li>
<li class="toclevel-3 tocsection-31"><a href="#Traditional_and_folk_medicine"><span class="tocnumber">13.1.2</span> <span class="toctext">Traditional and folk medicine</span></a></li>
</ul>
</li>
<li class="toclevel-2 tocsection-32"><a href="#Cultured_foods"><span class="tocnumber">13.2</span> <span class="toctext">Cultured foods</span></a></li>
<li class="toclevel-2 tocsection-33"><a href="#In_food"><span class="tocnumber">13.3</span> <span class="toctext">In food</span></a></li>
<li class="toclevel-2 tocsection-34"><a href="#Poisonous_fungi"><span class="tocnumber">13.4</span> <span class="toctext">Poisonous fungi</span></a></li>
<li class="toclevel-2 tocsection-35"><a href="#Pest_control"><span class="tocnumber">13.5</span> <span class="toctext">Pest control</span></a></li>
<li class="toclevel-2 tocsection-36"><a href="#Bioremediation"><span class="tocnumber">13.6</span> <span class="toctext">Bioremediation</span></a></li>
<li class="toclevel-2 tocsection-37"><a href="#Model_organisms"><span class="tocnumber">13.7</span> <span class="toctext">Model organisms</span></a></li>
<li class="toclevel-2 tocsection-38"><a href="#Others"><span class="tocnumber">13.8</span> <span class="toctext">Others</span></a></li>
</ul>
</li>
<li class="toclevel-1 tocsection-39"><a href="#See_also"><span class="tocnumber">14</span> <span class="toctext">See also</span></a></li>
<li class="toclevel-1 tocsection-40">
<a href="#References"><span class="tocnumber">15</span> <span class="toctext">References</span></a>
<ul>
<li class="toclevel-2 tocsection-41"><a href="#Cited_literature"><span class="tocnumber">15.1</span> <span class="toctext">Cited literature</span></a></li>
</ul>
</li>
<li class="toclevel-1 tocsection-42"><a href="#External_links"><span class="tocnumber">16</span> <span class="toctext">External links</span></a></li>
</ul>
</div>
<h2><span class="mw-headline" id="Etymology">Etymology</span></h2>
<p>The English word <i>fungus</i> is directly adopted from the <a href="/wiki/Latin" title="Latin">Latin</a> <i>fungus</i> (mushroom), used in the writings of <a href="/wiki/Horace" title="Horace">Horace</a> and <a href="/wiki/Pliny_the_Elder" title="Pliny the Elder">Pliny</a>.<sup id="cite_ref-Simpson1979_7-0" class="reference"><a href="#cite_note-Simpson1979-7">[7]</a></sup> This in turn is derived from the <a href="/wiki/Ancient_Greek" title="Ancient Greek">Greek</a> word <i>sphongos</i> (σφογγος "sponge"), which refers to the <a href="/wiki/Macroscopic" class="mw-redirect" title="Macroscopic">macroscopic</a> structures and morphology of mushrooms and molds;<sup id="cite_ref-Ainsworth2_8-0" class="reference"><a href="#cite_note-Ainsworth2-8">[8]</a></sup> the root is also used in other languages, such as the German <i>Schwamm</i> ("sponge") and <i>Schimmel</i> ("mold").<sup id="cite_ref-Mitzka1960_9-0" class="reference"><a href="#cite_note-Mitzka1960-9">[9]</a></sup> The use of the word <i>mycology</i>, which is derived from the Greek <i>mykes</i> (μύκης "mushroom") and <i>logos</i> (λόγος "discourse"),<sup id="cite_ref-10" class="reference"><a href="#cite_note-10">[10]</a></sup> to denote the scientific study of fungi is thought to have originated in 1836 with English naturalist <a href="/wiki/Miles_Joseph_Berkeley" title="Miles Joseph Berkeley">Miles Joseph Berkeley</a>'s publication <i>The English Flora of Sir James Edward Smith, Vol. 5.</i><sup id="cite_ref-Ainsworth2_8-1" class="reference"><a href="#cite_note-Ainsworth2-8">[8]</a></sup> A group of all the fungi present in a particular area or geographic region is known as <a href="/wiki/Mycobiota" title="Mycobiota">mycobiota</a> (plural noun, no singular), e.g., "the mycobiota of Ireland".<sup id="cite_ref-11" class="reference"><a href="#cite_note-11">[11]</a></sup></p>
<h2><span class="mw-headline" id="Characteristics">Characteristics</span></h2>
<div class="thumb tright">
<div class="thumbinner" style="width:222px;">
<a href="/wiki/File:HYPHAE.png" class="image"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/4/46/HYPHAE.png/220px-HYPHAE.png" width="220" height="116" class="thumbimage" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/46/HYPHAE.png/330px-HYPHAE.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/46/HYPHAE.png/440px-HYPHAE.png 2x" data-file-width="1364" data-file-height="721" /></a>
<div class="thumbcaption">
<div class="magnify"><a href="/wiki/File:HYPHAE.png" class="internal" title="Enlarge"></a></div>
<b>Fungal hyphae cells</b>
<div class="hlist">
<div>
<ol>
<li>Hyphal wall</li>
<li><a href="/wiki/Septum" title="Septum">Septum</a></li>
<li><a href="/wiki/Mitochondrion" title="Mitochondrion">Mitochondrion</a></li>
<li><a href="/wiki/Vacuole" title="Vacuole">Vacuole</a></li>
<li><a href="/wiki/Ergosterol" title="Ergosterol">Ergosterol</a> crystal</li>
<li><a href="/wiki/Ribosome" title="Ribosome">Ribosome</a></li>
<li>Nucleus</li>
<li><a href="/wiki/Endoplasmic_reticulum" title="Endoplasmic reticulum">Endoplasmic reticulum</a></li>
<li><a href="/wiki/Lipid_body" class="mw-redirect" title="Lipid body">Lipid body</a></li>
<li><a href="/wiki/Plasma_membrane" class="mw-redirect" title="Plasma membrane">Plasma membrane</a></li>
<li><a href="/wiki/Spitzenk%C3%B6rper" title="Spitzenkörper">Spitzenkörper</a></li>
<li><a href="/wiki/Golgi_apparatus" title="Golgi apparatus">Golgi apparatus</a></li>
</ol>
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<p>Before the introduction of <a href="/wiki/Molecular_phylogenetics" title="Molecular phylogenetics">molecular methods</a> for phylogenetic analysis, <a href="/wiki/Taxonomy_(biology)" title="Taxonomy (biology)">taxonomists</a> considered fungi to be members of the <a href="/wiki/Plant" title="Plant">plant kingdom</a> because of similarities in lifestyle: both fungi and plants are mainly <a href="/wiki/Sessility_(zoology)" class="mw-redirect" title="Sessility (zoology)">immobile</a>, and have similarities in general morphology and growth habitat. Like plants, fungi often grow in soil and, in the case of <a href="/wiki/Mushroom" title="Mushroom">mushrooms</a>, form conspicuous <a href="/wiki/Fruit_bodies" class="mw-redirect" title="Fruit bodies">fruit bodies</a>, which sometimes resemble plants such as <a href="/wiki/Mosses" class="mw-redirect" title="Mosses">mosses</a>. The fungi are now considered a separate kingdom, distinct from both plants and animals, from which they appear to have <a href="/wiki/Genetic_divergence" title="Genetic divergence">diverged</a> around one billion years ago (around the start of the <a href="/wiki/Neoproterozoic" title="Neoproterozoic">Neoproterozoic</a> Era).<sup id="cite_ref-Bruns2006_12-0" class="reference"><a href="#cite_note-Bruns2006-12">[12]</a></sup><sup id="cite_ref-Baldauf1993_13-0" class="reference"><a href="#cite_note-Baldauf1993-13">[13]</a></sup> Some morphological, biochemical, and genetic features are shared with other organisms, while others are unique to the fungi, clearly separating them from the other kingdoms:</p>
<p>Shared features:</p>
<ul>
<li>With other <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotes</a>: Fungal cells contain <a href="/wiki/Membrane-bound" class="mw-redirect" title="Membrane-bound">membrane-bound</a> <a href="/wiki/Cell_nucleus" title="Cell nucleus">nuclei</a> with <a href="/wiki/Chromosomes" class="mw-redirect" title="Chromosomes">chromosomes</a> that contain DNA with <a href="/wiki/Noncoding_DNA" class="mw-redirect" title="Noncoding DNA">noncoding regions</a> called <a href="/wiki/Intron" title="Intron">introns</a> and coding regions called <a href="/wiki/Exons" class="mw-redirect" title="Exons">exons</a>. Fungi have membrane-bound cytoplasmic <a href="/wiki/Organelles" class="mw-redirect" title="Organelles">organelles</a> such as <a href="/wiki/Mitochondria" class="mw-redirect" title="Mitochondria">mitochondria</a>, <a href="/wiki/Sterol" title="Sterol">sterol</a>-containing membranes, and <a href="/wiki/Ribosomes" class="mw-redirect" title="Ribosomes">ribosomes</a> of the <a href="/wiki/80S" class="mw-redirect" title="80S">80S</a> type.<sup id="cite_ref-14" class="reference"><a href="#cite_note-14">[14]</a></sup> They have a characteristic range of soluble carbohydrates and storage compounds, including <a href="/wiki/Sugar_alcohol" title="Sugar alcohol">sugar alcohols</a> (e.g., <a href="/wiki/Mannitol" title="Mannitol">mannitol</a>), <a href="/wiki/Disaccharide" title="Disaccharide">disaccharides</a>, (e.g., <a href="/wiki/Trehalose" title="Trehalose">trehalose</a>), and <a href="/wiki/Polysaccharide" title="Polysaccharide">polysaccharides</a> (e.g., <a href="/wiki/Glycogen" title="Glycogen">glycogen</a>, which is also found in animals<sup id="cite_ref-Deacon4_15-0" class="reference"><a href="#cite_note-Deacon4-15">[15]</a></sup>).</li>
<li>With animals: Fungi lack <a href="/wiki/Chloroplast" title="Chloroplast">chloroplasts</a> and are <a href="/wiki/Heterotroph" title="Heterotroph">heterotrophic</a> organisms and so require preformed <a href="/wiki/Organic_compound" title="Organic compound">organic compounds</a> as energy sources.<sup id="cite_ref-16" class="reference"><a href="#cite_note-16">[16]</a></sup></li>
<li>With plants: Fungi have a cell wall<sup id="cite_ref-17" class="reference"><a href="#cite_note-17">[17]</a></sup> and <a href="/wiki/Vacuole" title="Vacuole">vacuoles</a>.<sup id="cite_ref-Shoji2006_18-0" class="reference"><a href="#cite_note-Shoji2006-18">[18]</a></sup> They reproduce by both sexual and asexual means, and like <a href="/wiki/Primitive_(phylogenetics)" title="Primitive (phylogenetics)">basal</a> plant groups (such as <a href="/wiki/Fern" title="Fern">ferns</a> and <a href="/wiki/Moss" title="Moss">mosses</a>) produce <a href="/wiki/Spore" title="Spore">spores</a>. Similar to mosses and algae, fungi typically have <a href="/wiki/Haploid" class="mw-redirect" title="Haploid">haploid</a> nuclei.<sup id="cite_ref-Deacon58_19-0" class="reference"><a href="#cite_note-Deacon58-19">[19]</a></sup></li>
<li>With <a href="/wiki/Euglenoid" class="mw-redirect" title="Euglenoid">euglenoids</a> and bacteria: Higher fungi, euglenoids, and some bacteria produce the <a href="/wiki/Amino_acid" title="Amino acid">amino acid</a> <small>L</small>-lysine in specific <a href="/wiki/Biosynthesis" title="Biosynthesis">biosynthesis</a> steps, called the <a href="/wiki/%CE%91-aminoadipate_pathway" class="mw-redirect" title="Α-aminoadipate pathway">α-aminoadipate pathway</a>.<sup id="cite_ref-Zabriskie2000_20-0" class="reference"><a href="#cite_note-Zabriskie2000-20">[20]</a></sup><sup id="cite_ref-Xu2006_21-0" class="reference"><a href="#cite_note-Xu2006-21">[21]</a></sup></li>
<li>The cells of most fungi grow as tubular, elongated, and thread-like (filamentous) structures called <a href="/wiki/Hypha" title="Hypha">hyphae</a>, which may contain multiple nuclei and extend by growing at their tips. Each tip contains a set of aggregated <a href="/wiki/Vesicle_(biology)" class="mw-redirect" title="Vesicle (biology)">vesicles</a>—cellular structures consisting of <a href="/wiki/Protein" title="Protein">proteins</a>, <a href="/wiki/Lipid" title="Lipid">lipids</a>, and other organic molecules—called the <a href="/wiki/Spitzenk%C3%B6rper" title="Spitzenkörper">Spitzenkörper</a>.<sup id="cite_ref-22" class="reference"><a href="#cite_note-22">[22]</a></sup> Both fungi and <a href="/wiki/Oomycete" title="Oomycete">oomycetes</a> grow as filamentous hyphal cells.<sup id="cite_ref-23" class="reference"><a href="#cite_note-23">[23]</a></sup> In contrast, similar-looking organisms, such as filamentous <a href="/wiki/Green_algae" title="Green algae">green algae</a>, grow by repeated cell division within a chain of cells.<sup id="cite_ref-Deacon4_15-1" class="reference"><a href="#cite_note-Deacon4-15">[15]</a></sup> There are also single-celled fungi (<a href="/wiki/Yeast" title="Yeast">yeasts</a>) that do not form hyphae, and some fungi have both hyphal and yeast forms.<sup id="cite_ref-Alexopoulos_et_al.,_p._30_24-0" class="reference"><a href="#cite_note-Alexopoulos_et_al.,_p._30-24">[24]</a></sup></li>
<li>In common with some plant and animal species, <a href="/wiki/List_of_bioluminescent_fungi" class="mw-redirect" title="List of bioluminescent fungi">more than 70 fungal species</a> display <a href="/wiki/Bioluminescence" title="Bioluminescence">bioluminescence</a>.<sup id="cite_ref-Desjardin_2010_25-0" class="reference"><a href="#cite_note-Desjardin_2010-25">[25]</a></sup></li>
</ul>
<p>Unique features:</p>
<ul>
<li>Some species grow as unicellular yeasts that reproduce by <a href="/wiki/Budding" title="Budding">budding</a> or <a href="/wiki/Binary_fission" class="mw-redirect" title="Binary fission">binary fission</a>. <a href="/wiki/Dimorphic_fungi" class="mw-redirect" title="Dimorphic fungi">Dimorphic fungi</a> can switch between a yeast phase and a hyphal phase in response to environmental conditions.<sup id="cite_ref-Alexopoulos_et_al.,_p._30_24-1" class="reference"><a href="#cite_note-Alexopoulos_et_al.,_p._30-24">[24]</a></sup></li>
<li>The fungal cell wall is composed of <a href="/wiki/Glucan" title="Glucan">glucans</a> and <a href="/wiki/Chitin" title="Chitin">chitin</a>; while glucans are also found in plants and chitin in the <a href="/wiki/Exoskeleton" title="Exoskeleton">exoskeleton</a> of <a href="/wiki/Arthropods" class="mw-redirect" title="Arthropods">arthropods</a>,<sup id="cite_ref-26" class="reference"><a href="#cite_note-26">[26]</a></sup><sup id="cite_ref-Bowman2006_27-0" class="reference"><a href="#cite_note-Bowman2006-27">[27]</a></sup> fungi are the only organisms that combine these two structural molecules in their cell wall. Unlike those of plants and oomycetes, fungal cell walls do not contain cellulose.<sup id="cite_ref-28" class="reference"><a href="#cite_note-28">[28]</a></sup></li>
</ul>
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<a href="/wiki/File:Omphalotus_nidiformis_Binnamittalong_2_email.jpg" class="image"><img alt="A whitish fan or funnel-shaped mushroom growing at the base of a tree." src="//upload.wikimedia.org/wikipedia/commons/thumb/d/dd/Omphalotus_nidiformis_Binnamittalong_2_email.jpg/220px-Omphalotus_nidiformis_Binnamittalong_2_email.jpg" width="220" height="208" class="thumbimage" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/dd/Omphalotus_nidiformis_Binnamittalong_2_email.jpg/330px-Omphalotus_nidiformis_Binnamittalong_2_email.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/dd/Omphalotus_nidiformis_Binnamittalong_2_email.jpg/440px-Omphalotus_nidiformis_Binnamittalong_2_email.jpg 2x" data-file-width="1152" data-file-height="1088" /></a>
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<i><a href="/wiki/Omphalotus_nidiformis" title="Omphalotus nidiformis">Omphalotus nidiformis</a></i>, a bioluminescent mushroom
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<p>Most fungi lack an efficient system for the long-distance transport of water and nutrients, such as the <a href="/wiki/Xylem" title="Xylem">xylem</a> and <a href="/wiki/Phloem" title="Phloem">phloem</a> in many plants. To overcome this limitation, some fungi, such as <i><a href="/wiki/Armillaria" title="Armillaria">Armillaria</a></i>, form <a href="/wiki/Mycelial_cord" title="Mycelial cord">rhizomorphs</a>,<sup id="cite_ref-Mikhail2005_29-0" class="reference"><a href="#cite_note-Mikhail2005-29">[29]</a></sup> which resemble and perform functions similar to the <a href="/wiki/Root" title="Root">roots</a> of plants. As eukaryotes, fungi possess a <a href="/wiki/Biochemical_pathway" class="mw-redirect" title="Biochemical pathway">biosynthetic pathway</a> for producing <a href="/wiki/Terpene" title="Terpene">terpenes</a> that uses <a href="/wiki/Mevalonic_acid" title="Mevalonic acid">mevalonic acid</a> and <a href="/wiki/Pyrophosphate" title="Pyrophosphate">pyrophosphate</a> as <a href="/wiki/Precursor_(chemistry)" title="Precursor (chemistry)">chemical building blocks</a>.<sup id="cite_ref-Keller2005_30-0" class="reference"><a href="#cite_note-Keller2005-30">[30]</a></sup> Plants and some other organisms have an additional terpene biosynthesis pathway in their chloroplasts, a structure fungi and animals do not have.<sup id="cite_ref-Wu2007_31-0" class="reference"><a href="#cite_note-Wu2007-31">[31]</a></sup> Fungi produce several <a href="/wiki/Secondary_metabolite" title="Secondary metabolite">secondary metabolites</a> that are similar or identical in structure to those made by plants.<sup id="cite_ref-Keller2005_30-1" class="reference"><a href="#cite_note-Keller2005-30">[30]</a></sup> Many of the plant and fungal enzymes that make these compounds differ from each other in <a href="/wiki/Peptide_sequence" class="mw-redirect" title="Peptide sequence">sequence</a> and other characteristics, which indicates separate origins and convergent evolution of these enzymes in the fungi and plants.<sup id="cite_ref-Keller2005_30-2" class="reference"><a href="#cite_note-Keller2005-30">[30]</a></sup><sup id="cite_ref-Tudzynski2005_32-0" class="reference"><a href="#cite_note-Tudzynski2005-32">[32]</a></sup></p>
<h2><span class="mw-headline" id="Diversity">Diversity</span></h2>
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<a href="/wiki/File:Fungus_in_a_Wood.JPG" class="image"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/0/08/Fungus_in_a_Wood.JPG/220px-Fungus_in_a_Wood.JPG" width="220" height="146" class="thumbimage" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/08/Fungus_in_a_Wood.JPG/330px-Fungus_in_a_Wood.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/08/Fungus_in_a_Wood.JPG/440px-Fungus_in_a_Wood.JPG 2x" data-file-width="4288" data-file-height="2848" /></a>
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<a href="/wiki/Bracket_fungus" class="mw-redirect" title="Bracket fungus">Bracket fungi</a> on a tree stump
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<p>Fungi have a worldwide distribution, and grow in a wide range of habitats, including extreme environments such as <a href="/wiki/Desert_fungi" title="Desert fungi">deserts</a> or areas with high salt concentrations<sup id="cite_ref-Vaupotic2008_33-0" class="reference"><a href="#cite_note-Vaupotic2008-33">[33]</a></sup> or <a href="/wiki/Ionizing_radiation" title="Ionizing radiation">ionizing radiation</a>,<sup id="cite_ref-Dadachova2007_34-0" class="reference"><a href="#cite_note-Dadachova2007-34">[34]</a></sup> as well as in <a href="/wiki/Deep_sea" title="Deep sea">deep sea</a> sediments.<sup id="cite_ref-Raghukumar1998_35-0" class="reference"><a href="#cite_note-Raghukumar1998-35">[35]</a></sup> Some can survive the intense <a href="/wiki/Ultraviolet_radiation" class="mw-redirect" title="Ultraviolet radiation">UV</a> and <a href="/wiki/Cosmic_radiation" class="mw-redirect" title="Cosmic radiation">cosmic radiation</a> encountered during space travel.<sup id="cite_ref-Sancho2007_36-0" class="reference"><a href="#cite_note-Sancho2007-36">[36]</a></sup> Most grow in terrestrial environments, though several species live partly or solely in aquatic habitats, such as the <a href="/wiki/Chytrid" class="mw-redirect" title="Chytrid">chytrid</a> fungus <i><a href="/wiki/Batrachochytrium_dendrobatidis" title="Batrachochytrium dendrobatidis">Batrachochytrium dendrobatidis</a></i>, a <a href="/wiki/Parasite" class="mw-redirect" title="Parasite">parasite</a> that has been responsible for a worldwide decline in <a href="/wiki/Amphibian" title="Amphibian">amphibian</a> populations. This organism spends part of its life cycle as a motile <a href="/wiki/Zoospore" title="Zoospore">zoospore</a>, enabling it to propel itself through water and enter its amphibian host.<sup id="cite_ref-Brem2008_37-0" class="reference"><a href="#cite_note-Brem2008-37">[37]</a></sup> Other examples of aquatic fungi include those living in <a href="/wiki/Hydrothermal" class="mw-redirect" title="Hydrothermal">hydrothermal</a> areas of the ocean.<sup id="cite_ref-LeCalvez2009_38-0" class="reference"><a href="#cite_note-LeCalvez2009-38">[38]</a></sup></p>
<p>Around 120,000 species of fungi have been <a href="/wiki/Species_description" title="Species description">described</a> by <a href="/wiki/Taxonomy_(biology)" title="Taxonomy (biology)">taxonomists</a>,<sup id="cite_ref-Mueller2006_39-0" class="reference"><a href="#cite_note-Mueller2006-39">[39]</a></sup> but the global biodiversity of the fungus kingdom is not fully understood.<sup id="cite_ref-Mueller2006_39-1" class="reference"><a href="#cite_note-Mueller2006-39">[39]</a></sup> A 2017 estimate suggests there may be between 2.2 and 3.8 million species.<sup id="cite_ref-Lucking2017_5-1" class="reference"><a href="#cite_note-Lucking2017-5">[5]</a></sup> In mycology, species have historically been distinguished by a variety of methods and concepts. Classification based on <a href="/wiki/Morphology_(biology)" title="Morphology (biology)">morphological</a> characteristics, such as the size and shape of spores or fruiting structures, has traditionally dominated fungal taxonomy.<sup id="cite_ref-Kirk_et_al.,_p._489_40-0" class="reference"><a href="#cite_note-Kirk_et_al.,_p._489-40">[40]</a></sup> Species may also be distinguished by their <a href="/wiki/Biochemistry" title="Biochemistry">biochemical</a> and <a href="/wiki/Physiology" title="Physiology">physiological</a> characteristics, such as their ability to metabolize certain biochemicals, or their reaction to <a href="/wiki/Chemical_tests_in_mushroom_identification" class="mw-redirect" title="Chemical tests in mushroom identification">chemical tests</a>. The <a href="/wiki/Species#The_isolation_species_concept_in_more_detail" title="Species">biological species concept</a> discriminates species based on their ability to <a href="/wiki/Mating_in_fungi" title="Mating in fungi">mate</a>. The application of <a href="/wiki/Molecular_biology" title="Molecular biology">molecular</a> tools, such as <a href="/wiki/DNA_sequencing" title="DNA sequencing">DNA sequencing</a> and phylogenetic analysis, to study diversity has greatly enhanced the resolution and added robustness to estimates of <a href="/wiki/Genetic_diversity" title="Genetic diversity">genetic diversity</a> within various taxonomic groups.<sup id="cite_ref-Hibbett2007_41-0" class="reference"><a href="#cite_note-Hibbett2007-41">[41]</a></sup></p>
<h2><span class="mw-headline" id="Mycology">Mycology</span></h2>
<p><a href="/wiki/Mycology" title="Mycology">Mycology</a> is the branch of <a href="/wiki/Biology" title="Biology">biology</a> concerned with the systematic study of fungi, including their genetic and biochemical properties, their taxonomy, and their use to humans as a source of medicine, food, and <a href="/wiki/Entheogen" title="Entheogen">psychotropic substances</a> consumed for religious purposes, as well as their dangers, such as poisoning or infection. The field of <a href="/wiki/Phytopathology" class="mw-redirect" title="Phytopathology">phytopathology</a>, the study of plant diseases, is closely related because many plant pathogens are fungi.<sup id="cite_ref-Struck2006_42-0" class="reference"><a href="#cite_note-Struck2006-42">[42]</a></sup></p>
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<a href="/wiki/File:Pier_Antonio_Micheli.jpg" class="image"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d1/Pier_Antonio_Micheli.jpg/170px-Pier_Antonio_Micheli.jpg" width="170" height="258" class="thumbimage" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d1/Pier_Antonio_Micheli.jpg/255px-Pier_Antonio_Micheli.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d1/Pier_Antonio_Micheli.jpg/340px-Pier_Antonio_Micheli.jpg 2x" data-file-width="428" data-file-height="650" /></a>
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In 1729, <a href="/wiki/Pier_Antonio_Micheli" title="Pier Antonio Micheli">Pier Antonio Micheli</a> first published descriptions of fungi.
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<p>The use of fungi by humans dates back to prehistory; <a href="/wiki/%C3%96tzi_the_Iceman" class="mw-redirect" title="Ötzi the Iceman">Ötzi the Iceman</a>, a well-preserved mummy of a 5,300-year-old <a href="/wiki/Neolithic" title="Neolithic">Neolithic</a> man found frozen in the Austrian Alps, carried two species of <a href="/wiki/Polypore" title="Polypore">polypore</a> mushrooms that may have been used as <a href="/wiki/Tinder" title="Tinder">tinder</a> (<i><a href="/wiki/Fomes_fomentarius" title="Fomes fomentarius">Fomes fomentarius</a></i>), or for medicinal purposes (<i><a href="/wiki/Piptoporus_betulinus" class="mw-redirect" title="Piptoporus betulinus">Piptoporus betulinus</a></i>).<sup id="cite_ref-Peintner1998_43-0" class="reference"><a href="#cite_note-Peintner1998-43">[43]</a></sup> Ancient peoples have used fungi as food sources–often unknowingly–for millennia, in the preparation of leavened bread and fermented juices. Some of the oldest written records contain references to the destruction of crops that were probably caused by pathogenic fungi.<sup id="cite_ref-44" class="reference"><a href="#cite_note-44">[44]</a></sup></p>
<h3><span class="mw-headline" id="History">History</span></h3>
<p>Mycology is a relatively new science that became systematic after the development of the <a href="/wiki/Microscope" title="Microscope">microscope</a> in the 17th century. Although fungal spores were first observed by <a href="/wiki/Giambattista_della_Porta" title="Giambattista della Porta">Giambattista della Porta</a> in 1588, the seminal work in the development of mycology is considered to be the publication of <a href="/wiki/Pier_Antonio_Micheli" title="Pier Antonio Micheli">Pier Antonio Micheli</a>'s 1729 work <i>Nova plantarum genera</i>.<sup id="cite_ref-45" class="reference"><a href="#cite_note-45">[45]</a></sup> Micheli not only observed spores but also showed that, under the proper conditions, they could be induced into growing into the same species of fungi from which they originated.<sup id="cite_ref-46" class="reference"><a href="#cite_note-46">[46]</a></sup> Extending the use of the <a href="/wiki/Binomial_nomenclature" title="Binomial nomenclature">binomial system of nomenclature</a> introduced by <a href="/wiki/Carl_Linnaeus" title="Carl Linnaeus">Carl Linnaeus</a> in his <i><a href="/wiki/Species_plantarum" class="mw-redirect" title="Species plantarum">Species plantarum</a></i> (1753), the Dutch <a href="/wiki/Christian_Hendrik_Persoon" class="mw-redirect" title="Christian Hendrik Persoon">Christian Hendrik Persoon</a> (1761–1836) established the first classification of mushrooms with such skill as to be considered a founder of modern mycology. Later, <a href="/wiki/Elias_Magnus_Fries" title="Elias Magnus Fries">Elias Magnus Fries</a> (1794–1878) further elaborated the <a href="/wiki/Biological_classification" class="mw-redirect" title="Biological classification">classification</a> of fungi, using spore color and microscopic characteristics, methods still used by taxonomists today. Other notable early contributors to mycology in the 17th–19th and early 20th centuries include <a href="/wiki/Miles_Joseph_Berkeley" title="Miles Joseph Berkeley">Miles Joseph Berkeley</a>, <a href="/wiki/August_Carl_Joseph_Corda" title="August Carl Joseph Corda">August Carl Joseph Corda</a>, <a href="/wiki/Anton_de_Bary" class="mw-redirect" title="Anton de Bary">Anton de Bary</a>, the brothers <a href="/wiki/Louis_Ren%C3%A9_Tulasne" class="mw-redirect" title="Louis René Tulasne">Louis René</a> and <a href="/wiki/Charles_Tulasne" title="Charles Tulasne">Charles Tulasne</a>, <a href="/wiki/Arthur_Henry_Reginald_Buller" title="Arthur Henry Reginald Buller">Arthur H. R. Buller</a>, <a href="/wiki/Curtis_Gates_Lloyd" title="Curtis Gates Lloyd">Curtis G. Lloyd</a>, and <a href="/wiki/Pier_Andrea_Saccardo" title="Pier Andrea Saccardo">Pier Andrea Saccardo</a>. The 20th century has seen a modernization of mycology that has come from advances in <a href="/wiki/Biochemistry" title="Biochemistry">biochemistry</a>, <a href="/wiki/Genetics" title="Genetics">genetics</a>, <a href="/wiki/Molecular_biology" title="Molecular biology">molecular biology</a>, and <a href="/wiki/Biotechnology" title="Biotechnology">biotechnology</a>. The use of <a href="/wiki/DNA_sequencing" title="DNA sequencing">DNA sequencing</a> technologies and phylogenetic analysis has provided new insights into fungal relationships and <a href="/wiki/Biodiversity" title="Biodiversity">biodiversity</a>, and has challenged traditional morphology-based groupings in fungal <a href="/wiki/Taxonomy_(biology)" title="Taxonomy (biology)">taxonomy</a>.<sup id="cite_ref-Hawksworth2006_47-0" class="reference"><a href="#cite_note-Hawksworth2006-47">[47]</a></sup></p>
<h2><span class="mw-headline" id="Morphology">Morphology</span></h2>
<h3><span class="mw-headline" id="Microscopic_structures">Microscopic structures</span></h3>
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<a href="/wiki/File:Penicillium_labeled_cropped.jpg" class="image"><img alt="Monochrome micrograph showing Penicillium hyphae as long, transparent, tube-like structures a few micrometres across. Conidiophores branch out laterally from the hyphae, terminating in bundles of phialides on which spherical condidiophores are arranged like beads on a string. Septa are faintly visible as dark lines crossing the hyphae." src="//upload.wikimedia.org/wikipedia/commons/thumb/3/30/Penicillium_labeled_cropped.jpg/330px-Penicillium_labeled_cropped.jpg" width="330" height="194" class="thumbimage" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/30/Penicillium_labeled_cropped.jpg/495px-Penicillium_labeled_cropped.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/3/30/Penicillium_labeled_cropped.jpg 2x" data-file-width="648" data-file-height="380" /></a>
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An environmental isolate of <i><a href="/wiki/Penicillium" title="Penicillium">Penicillium</a></i>
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<ol>
<li><a href="/wiki/Hypha" title="Hypha">hypha</a></li>
<li><a href="/wiki/Conidiophore" class="mw-redirect" title="Conidiophore">conidiophore</a></li>
<li><a href="/wiki/Phialide" title="Phialide">phialide</a></li>
<li><a href="/wiki/Conidia" class="mw-redirect" title="Conidia">conidia</a></li>
<li><a href="/wiki/Septum" title="Septum">septa</a></li>
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<p>Most fungi grow as <a href="/wiki/Hypha" title="Hypha">hyphae</a>, which are cylindrical, thread-like structures 2–10 <a href="/wiki/Micrometres" class="mw-redirect" title="Micrometres">µm</a> in diameter and up to several centimeters in length. Hyphae grow at their tips (apices); new hyphae are typically formed by emergence of new tips along existing hyphae by a process called <i>branching</i>, or occasionally growing hyphal tips fork, giving rise to two parallel-growing hyphae.<sup id="cite_ref-Harris2008_48-0" class="reference"><a href="#cite_note-Harris2008-48">[48]</a></sup> Hyphae also sometimes fuse when they come into contact, a process called hyphal fusion (or <a href="/wiki/Anastamosis" class="mw-redirect" title="Anastamosis">anastamosis</a>). These growth processes lead to the development of a <a href="/wiki/Mycelium" title="Mycelium">mycelium</a>, an interconnected network of hyphae.<sup id="cite_ref-Alexopoulos_et_al.,_p._30_24-2" class="reference"><a href="#cite_note-Alexopoulos_et_al.,_p._30-24">[24]</a></sup> Hyphae can be either <a href="/wiki/Septum" title="Septum">septate</a> or <a href="/wiki/Coenocytic" class="mw-redirect" title="Coenocytic">coenocytic</a>. Septate hyphae are divided into compartments separated by cross walls (internal cell walls, called septa, that are formed at <a href="/wiki/Right_angle" title="Right angle">right angles</a> to the cell wall giving the hypha its shape), with each compartment containing one or more nuclei; coenocytic hyphae are not compartmentalized.<sup id="cite_ref-49" class="reference"><a href="#cite_note-49">[49]</a></sup> Septa have <a href="/wiki/Pit_connection#Characteristics" title="Pit connection">pores</a> that allow <a href="/wiki/Cytoplasm" title="Cytoplasm">cytoplasm</a>, <a href="/wiki/Organelle" title="Organelle">organelles</a>, and sometimes nuclei to pass through; an example is the dolipore septum in fungi of the phylum Basidiomycota.<sup id="cite_ref-50" class="reference"><a href="#cite_note-50">[50]</a></sup> Coenocytic hyphae are in essence <a href="/wiki/Multinucleate" title="Multinucleate">multinucleate</a> supercells.<sup id="cite_ref-Chang2004_51-0" class="reference"><a href="#cite_note-Chang2004-51">[51]</a></sup></p>
<p>Many species have developed specialized hyphal structures for nutrient uptake from living hosts; examples include <a href="/wiki/Haustoria" class="mw-redirect" title="Haustoria">haustoria</a> in plant-parasitic species of most fungal phyla, and <a href="/wiki/Arbuscular_mycorrhiza" title="Arbuscular mycorrhiza">arbuscules</a> of several <a href="/wiki/Mycorrhiza" title="Mycorrhiza">mycorrhizal</a> fungi, which penetrate into the host cells to consume nutrients.<sup id="cite_ref-Parniske2008_52-0" class="reference"><a href="#cite_note-Parniske2008-52">[52]</a></sup></p>
<p>Although fungi are <a href="/wiki/Opisthokont" title="Opisthokont">opisthokonts</a>—a grouping of evolutionarily related organisms broadly characterized by a single posterior <a href="/wiki/Flagellum" title="Flagellum">flagellum</a>—all phyla except for the <a href="/wiki/Chytrids" class="mw-redirect" title="Chytrids">chytrids</a> have lost their posterior flagella.<sup id="cite_ref-Steenkamp2006_53-0" class="reference"><a href="#cite_note-Steenkamp2006-53">[53]</a></sup> Fungi are unusual among the eukaryotes in having a cell wall that, in addition to <a href="/wiki/Glucan" title="Glucan">glucans</a> (e.g., β-1,3-glucan) and other typical components, also contains the <a href="/wiki/Biopolymer" title="Biopolymer">biopolymer</a> chitin.<sup id="cite_ref-Stevens2006_54-0" class="reference"><a href="#cite_note-Stevens2006-54">[54]</a></sup></p>
<h3><span class="mw-headline" id="Macroscopic_structures">Macroscopic structures</span></h3>
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<a href="/wiki/File:Armillaria_ostoyae_MO.jpg" class="image"><img alt="A cluster of large, thick-stem, light-brown gilled mushrooms growing at the base of a tree" src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0b/Armillaria_ostoyae_MO.jpg/220px-Armillaria_ostoyae_MO.jpg" width="220" height="165" class="thumbimage" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0b/Armillaria_ostoyae_MO.jpg/330px-Armillaria_ostoyae_MO.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/0b/Armillaria_ostoyae_MO.jpg/440px-Armillaria_ostoyae_MO.jpg 2x" data-file-width="3648" data-file-height="2736" /></a>
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<i><a href="/wiki/Armillaria_solidipes" class="mw-redirect" title="Armillaria solidipes">Armillaria solidipes</a></i>
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<p>Fungal mycelia can become visible to the naked eye, for example, on various surfaces and <a href="/wiki/Substrate_(biology)" title="Substrate (biology)">substrates</a>, such as damp walls and spoiled food, where they are commonly called <a href="/wiki/Mold" title="Mold">molds</a>. Mycelia grown on solid <a href="/wiki/Agar" title="Agar">agar</a> media in laboratory <a href="/wiki/Petri_dish" title="Petri dish">petri dishes</a> are usually referred to as <a href="/wiki/Colony_(biology)" title="Colony (biology)">colonies</a>. These colonies can exhibit growth shapes and colors (due to spores or <a href="/wiki/Biological_pigment" title="Biological pigment">pigmentation</a>) that can be used as diagnostic features in the identification of species or groups.<sup id="cite_ref-55" class="reference"><a href="#cite_note-55">[55]</a></sup> Some individual fungal colonies can reach extraordinary dimensions and ages as in the case of a <a href="/wiki/Clone_(cell_biology)" title="Clone (cell biology)">clonal</a> colony of <i><a href="/wiki/Armillaria_solidipes" class="mw-redirect" title="Armillaria solidipes">Armillaria solidipes</a></i>, which extends over an area of more than 900 <a href="/wiki/Hectare" title="Hectare">ha</a> (3.5 square miles), with an estimated age of nearly 9,000 years.<sup id="cite_ref-Ferguson2003_56-0" class="reference"><a href="#cite_note-Ferguson2003-56">[56]</a></sup></p>
<p>The <a href="/wiki/Apothecium" class="mw-redirect" title="Apothecium">apothecium</a>—a specialized structure important in <a href="/wiki/Sexual_reproduction" title="Sexual reproduction">sexual reproduction</a> in the ascomycetes—is a cup-shaped fruit body that is often macroscopic and holds the <a href="/wiki/Hymenium" title="Hymenium">hymenium</a>, a layer of tissue containing the spore-bearing cells.<sup id="cite_ref-57" class="reference"><a href="#cite_note-57">[57]</a></sup> The fruit bodies of the basidiomycetes (<a href="/wiki/Basidiocarp" title="Basidiocarp">basidiocarps</a>) and some ascomycetes can sometimes grow very large, and many are well known as <a href="/wiki/Mushroom" title="Mushroom">mushrooms</a>.</p>
<h2><span class="mw-headline" id="Growth_and_physiology">Growth and physiology</span></h2>
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<a href="/wiki/File:DecayingPeachSmall.gif" class="image"><img alt="Time-lapse photography sequence of a peach becoming progressively discolored and disfigured" src="//upload.wikimedia.org/wikipedia/commons/2/27/DecayingPeachSmall.gif" width="200" height="190" class="thumbimage" data-file-width="200" data-file-height="190" /></a>
<div class="thumbcaption"><a href="/wiki/Mold" title="Mold">Mold</a> growth covering a decaying <a href="/wiki/Peach" title="Peach">peach</a>. The frames were taken approximately 12 hours apart over a period of six days.</div>
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<p>The growth of fungi as hyphae on or in solid substrates or as single cells in aquatic environments is adapted for the efficient extraction of nutrients, because these growth forms have high <a href="/wiki/Surface_area_to_volume_ratio" class="mw-redirect" title="Surface area to volume ratio">surface area to volume ratios</a>.<sup id="cite_ref-Moss1986_58-0" class="reference"><a href="#cite_note-Moss1986-58">[58]</a></sup> Hyphae are specifically adapted for growth on solid surfaces, and to invade <a href="/wiki/Substrate_(biology)" title="Substrate (biology)">substrates</a> and tissues.<sup id="cite_ref-Penalva2002_59-0" class="reference"><a href="#cite_note-Penalva2002-59">[59]</a></sup> They can exert large penetrative mechanical forces; for example, many <a href="/wiki/Plant_pathogen" class="mw-redirect" title="Plant pathogen">plant pathogens</a>, including <i><a href="/wiki/Magnaporthe_grisea" title="Magnaporthe grisea">Magnaporthe grisea</a></i>, form a structure called an <a href="/wiki/Appressorium" title="Appressorium">appressorium</a> that evolved to puncture plant tissues.<sup id="cite_ref-Howard1991_60-0" class="reference"><a href="#cite_note-Howard1991-60">[60]</a></sup> The pressure generated by the appressorium, directed against the plant <a href="/wiki/Epidermis_(botany)" title="Epidermis (botany)">epidermis</a>, can exceed 8 <a href="/wiki/Pascal_(unit)" title="Pascal (unit)">megapascals</a> (1,200 psi).<sup id="cite_ref-Howard1991_60-1" class="reference"><a href="#cite_note-Howard1991-60">[60]</a></sup> The filamentous fungus <i><a href="/wiki/Paecilomyces_lilacinus" class="mw-redirect" title="Paecilomyces lilacinus">Paecilomyces lilacinus</a></i> uses a similar structure to penetrate the eggs of <a href="/wiki/Nematode" title="Nematode">nematodes</a>.<sup id="cite_ref-Money1997_61-0" class="reference"><a href="#cite_note-Money1997-61">[61]</a></sup></p>
<p>The mechanical pressure exerted by the appressorium is generated from physiological processes that increase intracellular <a href="/wiki/Turgor" class="mw-redirect" title="Turgor">turgor</a> by producing <a href="/wiki/Osmolyte" title="Osmolyte">osmolytes</a> such as <a href="/wiki/Glycerol" title="Glycerol">glycerol</a>.<sup id="cite_ref-Wang2005_62-0" class="reference"><a href="#cite_note-Wang2005-62">[62]</a></sup> Adaptations such as these are complemented by <a href="/wiki/Cellulase" title="Cellulase">hydrolytic enzymes</a> secreted into the environment to digest large organic molecules—such as <a href="/wiki/Polysaccharide" title="Polysaccharide">polysaccharides</a>, <a href="/wiki/Protein" title="Protein">proteins</a>, and <a href="/wiki/Lipid" title="Lipid">lipids</a>—into smaller molecules that may then be absorbed as nutrients.<sup id="cite_ref-Pereira2007_63-0" class="reference"><a href="#cite_note-Pereira2007-63">[63]</a></sup><sup id="cite_ref-Schaller2007_64-0" class="reference"><a href="#cite_note-Schaller2007-64">[64]</a></sup><sup id="cite_ref-Farrar1985_65-0" class="reference"><a href="#cite_note-Farrar1985-65">[65]</a></sup> The vast majority of filamentous fungi grow in a polar fashion (extending in one direction) by elongation at the tip (apex) of the hypha.<sup id="cite_ref-Fischer2008_66-0" class="reference"><a href="#cite_note-Fischer2008-66">[66]</a></sup> Other forms of fungal growth include intercalary extension (longitudinal expansion of hyphal compartments that are below the apex) as in the case of some <a href="/wiki/Endophyte" title="Endophyte">endophytic</a> fungi,<sup id="cite_ref-Christensen2008_67-0" class="reference"><a href="#cite_note-Christensen2008-67">[67]</a></sup> or growth by volume expansion during the development of mushroom <a href="/wiki/Stipe_(mycology)" title="Stipe (mycology)">stipes</a> and other large organs.<sup id="cite_ref-Money2002_68-0" class="reference"><a href="#cite_note-Money2002-68">[68]</a></sup> Growth of fungi as <a href="/wiki/Multicellularity" class="mw-redirect" title="Multicellularity">multicellular structures</a> consisting of <a href="/wiki/Somatic_(biology)" title="Somatic (biology)">somatic</a> and reproductive cells—a feature independently evolved in animals and plants<sup id="cite_ref-Willensdorfer2009_69-0" class="reference"><a href="#cite_note-Willensdorfer2009-69">[69]</a></sup>—has several functions, including the development of fruit bodies for dissemination of sexual spores (see above) and <a href="/wiki/Biofilm" title="Biofilm">biofilms</a> for substrate colonization and <a href="/wiki/Intercellular_communication" class="mw-redirect" title="Intercellular communication">intercellular communication</a>.<sup id="cite_ref-Daniels2006_70-0" class="reference"><a href="#cite_note-Daniels2006-70">[70]</a></sup></p>
<p>The fungi are traditionally considered <a href="/wiki/Heterotroph" title="Heterotroph">heterotrophs</a>, organisms that rely solely on <a href="/wiki/Carbon_fixation" title="Carbon fixation">carbon fixed</a> by other organisms for <a href="/wiki/Metabolism" title="Metabolism">metabolism</a>. Fungi have <a href="/wiki/Evolution" title="Evolution">evolved</a> a high degree of metabolic versatility that allows them to use a diverse range of organic substrates for growth, including simple compounds such as <a href="/wiki/Nitrate" title="Nitrate">nitrate</a>, <a href="/wiki/Ammonia" title="Ammonia">ammonia</a>, <a href="/wiki/Acetate" title="Acetate">acetate</a>, or <a href="/wiki/Ethanol" title="Ethanol">ethanol</a>.<sup id="cite_ref-Marzluf1981_71-0" class="reference"><a href="#cite_note-Marzluf1981-71">[71]</a></sup><sup id="cite_ref-Heynes1994_72-0" class="reference"><a href="#cite_note-Heynes1994-72">[72]</a></sup> In some species the pigment <a href="/wiki/Melanin" title="Melanin">melanin</a> may play a role in extracting energy from <a href="/wiki/Ionizing_radiation" title="Ionizing radiation">ionizing radiation</a>, such as <a href="/wiki/Gamma_rays" class="mw-redirect" title="Gamma rays">gamma radiation</a>. This form of <a href="/wiki/Radiotrophic_fungus" title="Radiotrophic fungus">"radiotrophic"</a> growth has been described for only a few species, the effects on growth rates are small, and the underlying <a href="/wiki/Biophysics" title="Biophysics">biophysical</a> and biochemical processes are not well known.<sup id="cite_ref-Dadachova2007_34-1" class="reference"><a href="#cite_note-Dadachova2007-34">[34]</a></sup> This process might bear similarity to <a href="/wiki/Carbon_fixation" title="Carbon fixation">CO<sub>2</sub> fixation</a> via <a href="/wiki/Visible_spectrum" title="Visible spectrum">visible light</a>, but instead uses ionizing radiation as a source of energy.<sup id="cite_ref-Dadachova2008_73-0" class="reference"><a href="#cite_note-Dadachova2008-73">[73]</a></sup></p>
<h2><span class="mw-headline" id="Reproduction">Reproduction</span></h2>
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<a href="/wiki/File:Polyporus_squamosus_Molter.jpg" class="image"><img alt="Two thickly stemmed brownish mushrooms with scales on the upper surface, growing out of a tree trunk" src="//upload.wikimedia.org/wikipedia/commons/thumb/3/36/Polyporus_squamosus_Molter.jpg/220px-Polyporus_squamosus_Molter.jpg" width="220" height="165" class="thumbimage" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/36/Polyporus_squamosus_Molter.jpg/330px-Polyporus_squamosus_Molter.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/36/Polyporus_squamosus_Molter.jpg/440px-Polyporus_squamosus_Molter.jpg 2x" data-file-width="2612" data-file-height="1959" /></a>
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<i><a href="/wiki/Polyporus_squamosus" class="mw-redirect" title="Polyporus squamosus">Polyporus squamosus</a></i>
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<p>Fungal reproduction is complex, reflecting the differences in lifestyles and genetic makeup within this diverse kingdom of organisms.<sup id="cite_ref-74" class="reference"><a href="#cite_note-74">[74]</a></sup> It is estimated that a third of all fungi reproduce using more than one method of propagation; for example, reproduction may occur in two well-differentiated stages within the <a href="/wiki/Biological_life_cycle" title="Biological life cycle">life cycle</a> of a species, the <a href="/wiki/Teleomorph" class="mw-redirect" title="Teleomorph">teleomorph</a> and the <a href="/wiki/Anamorph" class="mw-redirect" title="Anamorph">anamorph</a>.<sup id="cite_ref-75" class="reference"><a href="#cite_note-75">[75]</a></sup> Environmental conditions trigger genetically determined developmental states that lead to the creation of specialized structures for sexual or asexual reproduction. These structures aid reproduction by efficiently dispersing spores or spore-containing <a href="/wiki/Propagule" title="Propagule">propagules</a>.</p>
<h3><span class="mw-headline" id="Asexual_reproduction">Asexual reproduction</span></h3>
<p><a href="/wiki/Asexual_reproduction" title="Asexual reproduction">Asexual reproduction</a> occurs via vegetative spores (<a href="/wiki/Conidium" title="Conidium">conidia</a>) or through <a href="/wiki/Fragmentation_(reproduction)" title="Fragmentation (reproduction)">mycelial fragmentation</a>. Mycelial fragmentation occurs when a fungal mycelium separates into pieces, and each component grows into a separate mycelium. Mycelial fragmentation and vegetative spores maintain <a href="/wiki/Clone_(genetics)" class="mw-redirect" title="Clone (genetics)">clonal</a> populations adapted to a specific <a href="/wiki/Ecological_niche" title="Ecological niche">niche</a>, and allow more rapid dispersal than sexual reproduction.<sup id="cite_ref-Heitman2005_76-0" class="reference"><a href="#cite_note-Heitman2005-76">[76]</a></sup> The "Fungi imperfecti" (fungi lacking the perfect or sexual stage) or <a href="/wiki/Deuteromycota" class="mw-redirect" title="Deuteromycota">Deuteromycota</a> comprise all the species that lack an observable sexual cycle.<sup id="cite_ref-Alcamo2004_77-0" class="reference"><a href="#cite_note-Alcamo2004-77">[77]</a></sup> <a href="/wiki/Deuteromycota" class="mw-redirect" title="Deuteromycota">Deuteromycota</a> is not an accepted taxonomic clade, and is now taken to mean simply fungi that lack a known sexual stage.</p>
<h3><span class="mw-headline" id="Sexual_reproduction">Sexual reproduction</span></h3>
<div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Mating_in_fungi" title="Mating in fungi">Mating in fungi</a> and <a href="/wiki/Sexual_selection_in_fungi" title="Sexual selection in fungi">Sexual selection in fungi</a></div>
<p>Sexual reproduction with <a href="/wiki/Meiosis" title="Meiosis">meiosis</a> has been directly observed in all fungal phyla except <a href="/wiki/Glomeromycota" title="Glomeromycota">Glomeromycota</a> <sup id="cite_ref-Redecker2006_78-0" class="reference"><a href="#cite_note-Redecker2006-78">[78]</a></sup> (genetic analysis suggests meiosis in Glomeromycota as well). It differs in many aspects from sexual reproduction in animals or plants. Differences also exist between fungal groups and can be used to discriminate species by morphological differences in sexual structures and reproductive strategies.<sup id="cite_ref-Guarro1999_79-0" class="reference"><a href="#cite_note-Guarro1999-79">[79]</a></sup><sup id="cite_ref-Taylor2000_80-0" class="reference"><a href="#cite_note-Taylor2000-80">[80]</a></sup> Mating experiments between fungal isolates may identify species on the basis of biological species concepts.<sup id="cite_ref-Taylor2000_80-1" class="reference"><a href="#cite_note-Taylor2000-80">[80]</a></sup> The major fungal groupings have initially been delineated based on the morphology of their sexual structures and spores; for example, the spore-containing structures, <a href="/wiki/Ascus" title="Ascus">asci</a> and <a href="/wiki/Basidium" title="Basidium">basidia</a>, can be used in the identification of ascomycetes and basidiomycetes, respectively. Some species may allow mating only between individuals of opposite <a href="/wiki/Mating_type" title="Mating type">mating type</a>, whereas others can mate and sexually reproduce with any other individual or itself. Species of the former <a href="/wiki/Mating_system" title="Mating system">mating system</a> are called <a href="/wiki/Heterothallic" class="mw-redirect" title="Heterothallic">heterothallic</a>, and of the latter <a href="/wiki/Homothallic" class="mw-redirect" title="Homothallic">homothallic</a>.<sup id="cite_ref-Metzenberg1990_81-0" class="reference"><a href="#cite_note-Metzenberg1990-81">[81]</a></sup></p>
<p>Most fungi have both a <a href="/wiki/Haploid" class="mw-redirect" title="Haploid">haploid</a> and a <a href="/wiki/Diploid" class="mw-redirect" title="Diploid">diploid</a> stage in their life cycles. In sexually reproducing fungi, compatible individuals may combine by fusing their hyphae together into an interconnected network; this process, <a href="/wiki/Anastomosis" title="Anastomosis">anastomosis</a>, is required for the initiation of the sexual cycle. Many ascomycetes and basidiomycetes go through a <a href="/wiki/Dikaryotic" class="mw-redirect" title="Dikaryotic">dikaryotic</a> stage, in which the nuclei inherited from the two parents do not combine immediately after cell fusion, but remain separate in the hyphal cells (see <a href="/wiki/Heterokaryosis" class="mw-redirect" title="Heterokaryosis">heterokaryosis</a>).<sup id="cite_ref-82" class="reference"><a href="#cite_note-82">[82]</a></sup></p>
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<a href="/wiki/File:Morelasci.jpg" class="image"><img alt="Microscopic view of numerous translucent or transparent elongated sac-like structures each containing eight spheres lined up in a row" src="//upload.wikimedia.org/wikipedia/commons/thumb/6/61/Morelasci.jpg/220px-Morelasci.jpg" width="220" height="165" class="thumbimage" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/61/Morelasci.jpg/330px-Morelasci.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/61/Morelasci.jpg/440px-Morelasci.jpg 2x" data-file-width="1388" data-file-height="1040" /></a>
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The 8-spore asci of <i><a href="/wiki/Morchella_elata" title="Morchella elata">Morchella elata</a></i>, viewed with <a href="/wiki/Phase_contrast_microscopy" class="mw-redirect" title="Phase contrast microscopy">phase contrast microscopy</a>
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<p>In ascomycetes, dikaryotic hyphae of the <a href="/wiki/Hymenium" title="Hymenium">hymenium</a> (the spore-bearing tissue layer) form a characteristic <i>hook</i> at the hyphal septum. During <a href="/wiki/Cell_division" title="Cell division">cell division</a>, formation of the hook ensures proper distribution of the newly divided nuclei into the apical and basal hyphal compartments. An ascus (plural <i>asci</i>) is then formed, in which <a href="/wiki/Karyogamy" title="Karyogamy">karyogamy</a> (nuclear fusion) occurs. Asci are embedded in an <a href="/wiki/Ascocarp" title="Ascocarp">ascocarp</a>, or fruiting body. Karyogamy in the asci is followed immediately by meiosis and the production of <a href="/wiki/Ascospore" title="Ascospore">ascospores</a>. After dispersal, the ascospores may germinate and form a new haploid mycelium.<sup id="cite_ref-83" class="reference"><a href="#cite_note-83">[83]</a></sup></p>
<p>Sexual reproduction in basidiomycetes is similar to that of the ascomycetes. Compatible haploid hyphae fuse to produce a dikaryotic mycelium. However, the dikaryotic phase is more extensive in the basidiomycetes, often also present in the vegetatively growing mycelium. A specialized anatomical structure, called a <a href="/wiki/Clamp_connection" title="Clamp connection">clamp connection</a>, is formed at each hyphal septum. As with the structurally similar hook in the ascomycetes, the clamp connection in the basidiomycetes is required for controlled transfer of nuclei during cell division, to maintain the dikaryotic stage with two genetically different nuclei in each hyphal compartment.<sup id="cite_ref-84" class="reference"><a href="#cite_note-84">[84]</a></sup> A <a href="/wiki/Basidiocarp" title="Basidiocarp">basidiocarp</a> is formed in which club-like structures known as <a href="/wiki/Basidia" class="mw-redirect" title="Basidia">basidia</a> generate haploid <a href="/wiki/Basidiospores" class="mw-redirect" title="Basidiospores">basidiospores</a> after karyogamy and meiosis.<sup id="cite_ref-85" class="reference"><a href="#cite_note-85">[85]</a></sup> The most commonly known basidiocarps are mushrooms, but they may also take other forms (see <a href="/wiki/Fungus#Morphology" title="Fungus">Morphology</a> section).</p>
<p>In glomeromycetes (formerly zygomycetes), haploid hyphae of two individuals fuse, forming a <a href="/wiki/Gametangium" title="Gametangium">gametangium</a>, a specialized cell structure that becomes a fertile <a href="/wiki/Gamete" title="Gamete">gamete</a>-producing cell. The gametangium develops into a <a href="/wiki/Zygospore" title="Zygospore">zygospore</a>, a thick-walled spore formed by the union of gametes. When the zygospore germinates, it undergoes <a href="/wiki/Meiosis" title="Meiosis">meiosis</a>, generating new haploid hyphae, which may then form asexual <a href="/wiki/Sporangiospore" class="mw-redirect" title="Sporangiospore">sporangiospores</a>. These sporangiospores allow the fungus to rapidly disperse and germinate into new genetically identical haploid fungal mycelia.<sup id="cite_ref-86" class="reference"><a href="#cite_note-86">[86]</a></sup></p>
<h3><span class="mw-headline" id="Spore_dispersal">Spore dispersal</span></h3>
<p>
Both asexual and sexual spores or sporangiospores are often actively dispersed by forcible ejection from their reproductive structures. This ejection ensures exit of the spores from the reproductive structures as well as traveling through the air over long distances.
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The bird's nest fungus <i><a href="/wiki/Cyathus_stercoreus" title="Cyathus stercoreus">Cyathus stercoreus</a></i>
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<p> Specialized mechanical and physiological mechanisms, as well as spore surface structures (such as <a href="/wiki/Hydrophobin" title="Hydrophobin">hydrophobins</a>), enable efficient spore ejection.<sup id="cite_ref-Linder2005_87-0" class="reference"><a href="#cite_note-Linder2005-87">[87]</a></sup> For example, the structure of the <a href="/wiki/Ascus" title="Ascus">spore-bearing cells</a> in some ascomycete species is such that the buildup of <a href="/wiki/Osmolyte" title="Osmolyte">substances</a> affecting cell volume and fluid balance enables the explosive discharge of spores into the air.<sup id="cite_ref-Trail2007_88-0" class="reference"><a href="#cite_note-Trail2007-88">[88]</a></sup> The forcible discharge of single spores termed <i>ballistospores</i> involves formation of a small drop of water (Buller's drop), which upon contact with the spore leads to its projectile release with an initial acceleration of more than 10,000 <a href="/wiki/G-force" title="G-force">g</a>;<sup id="cite_ref-Pringle2005_89-0" class="reference"><a href="#cite_note-Pringle2005-89">[89]</a></sup> the net result is that the spore is ejected 0.01–0.02 cm, sufficient distance for it to fall through the gills or pores into the air below.<sup id="cite_ref-90" class="reference"><a href="#cite_note-90">[90]</a></sup> Other fungi, like the <a href="/wiki/Puffballs" class="mw-redirect" title="Puffballs">puffballs</a>, rely on alternative mechanisms for spore release, such as external mechanical forces. The <a href="/wiki/Nidulariaceae" title="Nidulariaceae">bird's nest fungi</a> use the force of falling water drops to liberate the spores from cup-shaped fruiting bodies.<sup id="cite_ref-Brodie1975_91-0" class="reference"><a href="#cite_note-Brodie1975-91">[91]</a></sup> Another strategy is seen in the <a href="/wiki/Stinkhorns" class="mw-redirect" title="Stinkhorns">stinkhorns</a>, a group of fungi with lively colors and putrid odor that attract insects to disperse their spores.<sup id="cite_ref-92" class="reference"><a href="#cite_note-92">[92]</a></sup></p>
<h3><span class="mw-headline" id="Other_sexual_processes">Other sexual processes</span></h3>
<p>Besides regular sexual reproduction with meiosis, certain fungi, such as those in the genera <i><a href="/wiki/Penicillium" title="Penicillium">Penicillium</a></i> and <i><a href="/wiki/Aspergillus" title="Aspergillus">Aspergillus</a></i>, may exchange genetic material via <a href="/wiki/Parasexuality" class="mw-redirect" title="Parasexuality">parasexual</a> processes, initiated by anastomosis between hyphae and <a href="/wiki/Plasmogamy" title="Plasmogamy">plasmogamy</a> of fungal cells.<sup id="cite_ref-93" class="reference"><a href="#cite_note-93">[93]</a></sup> The frequency and relative importance of parasexual events is unclear and may be lower than other sexual processes. It is known to play a role in intraspecific hybridization<sup id="cite_ref-Furlaneto1992_94-0" class="reference"><a href="#cite_note-Furlaneto1992-94">[94]</a></sup> and is likely required for hybridization between species, which has been associated with major events in fungal evolution.<sup id="cite_ref-Schardl2003_95-0" class="reference"><a href="#cite_note-Schardl2003-95">[95]</a></sup></p>
<h2><span class="mw-headline" id="Evolution">Evolution</span></h2>
<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Evolution_of_fungi" title="Evolution of fungi">Evolution of fungi</a></div>
<p>In contrast to <a href="/wiki/Evolutionary_history_of_plants" title="Evolutionary history of plants">plants</a> and <a href="/wiki/Evolutionary_history_of_life" title="Evolutionary history of life">animals</a>, the early fossil record of the fungi is meager. Factors that likely contribute to the under-representation of fungal species among fossils include the nature of fungal <a href="/wiki/Sporocarp_(fungi)" title="Sporocarp (fungi)">fruiting bodies</a>, which are soft, fleshy, and easily degradable tissues and the microscopic dimensions of most fungal structures, which therefore are not readily evident. Fungal fossils are difficult to distinguish from those of other microbes, and are most easily identified when they resemble <a href="/wiki/Extant_taxon" class="mw-redirect" title="Extant taxon">extant</a> fungi.<sup id="cite_ref-Donoghue2004_96-0" class="reference"><a href="#cite_note-Donoghue2004-96">[96]</a></sup> Often recovered from a <a href="/wiki/Permineralization" title="Permineralization">permineralized</a> plant or animal host, these samples are typically studied by making thin-section preparations that can be examined with <a href="/wiki/Optical_microscope" title="Optical microscope">light microscopy</a> or <a href="/wiki/Transmission_electron_microscopy" title="Transmission electron microscopy">transmission electron microscopy</a>.<sup id="cite_ref-97" class="reference"><a href="#cite_note-97">[97]</a></sup> Researchers study <a href="/wiki/Compression_fossil" title="Compression fossil">compression fossils</a> by dissolving the surrounding matrix with acid and then using light or <a href="/wiki/Scanning_electron_microscopy" class="mw-redirect" title="Scanning electron microscopy">scanning electron microscopy</a> to examine surface details.<sup id="cite_ref-98" class="reference"><a href="#cite_note-98">[98]</a></sup></p>
<p>The earliest fossils possessing features typical of fungi date to the <a href="/wiki/Paleoproterozoic" title="Paleoproterozoic">Paleoproterozoic</a> era, some <span class="nourlexpansion plainlinks" style="white-space:nowrap;"><a class="external text" href="http://tools.wmflabs.org/timescale/?Ma=2,400">2,400</a> million years ago</span> (<a href="/wiki/Annum" class="mw-redirect" title="Annum">Ma</a>); these multicellular <a href="/wiki/Benthic" class="mw-redirect" title="Benthic">benthic</a> organisms had filamentous structures capable of <a href="/wiki/Anastomosis" title="Anastomosis">anastomosis</a>.<sup id="cite_ref-99" class="reference"><a href="#cite_note-99">[99]</a></sup> Other studies (2009) estimate the arrival of fungal organisms at about 760–1060 Ma on the basis of comparisons of the rate of evolution in closely related groups.<sup id="cite_ref-Lucking2009_100-0" class="reference"><a href="#cite_note-Lucking2009-100">[100]</a></sup> For much of the <a href="/wiki/Paleozoic" title="Paleozoic">Paleozoic</a> Era (542–251 Ma), the fungi appear to have been aquatic and consisted of organisms similar to the extant <a href="/wiki/Chytrid" class="mw-redirect" title="Chytrid">chytrids</a> in having flagellum-bearing spores.<sup id="cite_ref-James2006b_101-0" class="reference"><a href="#cite_note-James2006b-101">[101]</a></sup> The evolutionary adaptation from an aquatic to a terrestrial lifestyle necessitated a diversification of ecological strategies for obtaining nutrients, including <a href="/wiki/Parasitism" title="Parasitism">parasitism</a>, <a href="/wiki/Saphrotrophic_nutrition" class="mw-redirect" title="Saphrotrophic nutrition">saprobism</a>, and the development of <a href="/wiki/Mutualism_(biology)" title="Mutualism (biology)">mutualistic</a> relationships such as <a href="/wiki/Mycorrhiza" title="Mycorrhiza">mycorrhiza</a> and lichenization.<sup id="cite_ref-102" class="reference"><a href="#cite_note-102">[102]</a></sup> Recent (2009) studies suggest that the ancestral ecological state of the <a href="/wiki/Ascomycota" title="Ascomycota">Ascomycota</a> was saprobism, and that independent <a href="/wiki/Lichen" title="Lichen">lichenization</a> events have occurred multiple times.<sup id="cite_ref-Schoch2009_103-0" class="reference"><a href="#cite_note-Schoch2009-103">[103]</a></sup></p>
<p>It is presumed that the fungi colonized the land during the <a href="/wiki/Cambrian" title="Cambrian">Cambrian</a> (542–488.3 Ma), long before land plants.<sup id="cite_ref-Brundrett2002_104-0" class="reference"><a href="#cite_note-Brundrett2002-104">[104]</a></sup> Fossilized hyphae and spores recovered from the <a href="/wiki/Ordovician" title="Ordovician">Ordovician</a> of Wisconsin (460 Ma) resemble modern-day <a href="/wiki/Glomerales" title="Glomerales">Glomerales</a>, and existed at a time when the land flora likely consisted of only non-vascular <a href="/wiki/Bryophyte" title="Bryophyte">bryophyte</a>-like plants.<sup id="cite_ref-Redecker2000_105-0" class="reference"><a href="#cite_note-Redecker2000-105">[105]</a></sup> <a href="/wiki/Prototaxites" title="Prototaxites">Prototaxites</a>, which was probably a fungus or lichen, would have been the tallest organism of the late <a href="/wiki/Silurian" title="Silurian">Silurian</a>. Fungal fossils do not become common and uncontroversial until the early <a href="/wiki/Devonian" title="Devonian">Devonian</a> (416–359.2 Ma), when they occur abundantly in the <a href="/wiki/Rhynie_chert" title="Rhynie chert">Rhynie chert</a>, mostly as <a href="/wiki/Zygomycota" title="Zygomycota">Zygomycota</a> and <a href="/wiki/Chytridiomycota" title="Chytridiomycota">Chytridiomycota</a>.<sup id="cite_ref-Brundrett2002_104-1" class="reference"><a href="#cite_note-Brundrett2002-104">[104]</a></sup><sup id="cite_ref-Taylor1996_106-0" class="reference"><a href="#cite_note-Taylor1996-106">[106]</a></sup><sup id="cite_ref-Dotzler2009_107-0" class="reference"><a href="#cite_note-Dotzler2009-107">[107]</a></sup> At about this same time, approximately 400 Ma, the Ascomycota and Basidiomycota diverged,<sup id="cite_ref-Taylor2006_108-0" class="reference"><a href="#cite_note-Taylor2006-108">[108]</a></sup> and all modern <a href="/wiki/Class_(biology)" title="Class (biology)">classes</a> of fungi were present by the Late <a href="/wiki/Carboniferous" title="Carboniferous">Carboniferous</a> (<a href="/wiki/Pennsylvanian_(geology)" title="Pennsylvanian (geology)">Pennsylvanian</a>, 318.1–299 Ma).<sup id="cite_ref-urlFungi_109-0" class="reference"><a href="#cite_note-urlFungi-109">[109]</a></sup></p>
<p><a href="/wiki/Lichen" title="Lichen">Lichen</a>-like fossils have been found in the <a href="/wiki/Doushantuo_Formation" title="Doushantuo Formation">Doushantuo Formation</a> in southern China dating back to 635–551 Ma.<sup id="cite_ref-Yuan2005_110-0" class="reference"><a href="#cite_note-Yuan2005-110">[110]</a></sup> Lichens formed a component of the early terrestrial ecosystems, and the estimated age of the oldest terrestrial lichen fossil is 400 Ma;<sup id="cite_ref-Karatygin2009_111-0" class="reference"><a href="#cite_note-Karatygin2009-111">[111]</a></sup> this date corresponds to the age of the oldest known <a href="/wiki/Sporocarp_(fungi)" title="Sporocarp (fungi)">sporocarp</a> fossil, a <i>Paleopyrenomycites</i> species found in the Rhynie Chert.<sup id="cite_ref-Taylor2005_112-0" class="reference"><a href="#cite_note-Taylor2005-112">[112]</a></sup> The oldest fossil with microscopic features resembling modern-day basidiomycetes is <i>Palaeoancistrus</i>, found permineralized with a <a href="/wiki/Fern" title="Fern">fern</a> from the Pennsylvanian.<sup id="cite_ref-Dennis1970_113-0" class="reference"><a href="#cite_note-Dennis1970-113">[113]</a></sup> Rare in the fossil record are the Homobasidiomycetes (a <a href="/wiki/Taxon" title="Taxon">taxon</a> roughly equivalent to the mushroom-producing species of the <a href="/wiki/Agaricomycetes" title="Agaricomycetes">Agaricomycetes</a>). Two <a href="/wiki/Amber" title="Amber">amber</a>-preserved specimens provide evidence that the earliest known mushroom-forming fungi (the extinct species <i><a href="/wiki/Archaeomarasmius_leggetti" class="mw-redirect" title="Archaeomarasmius leggetti">Archaeomarasmius leggetti</a></i>) appeared during the late <a href="/wiki/Cretaceous" title="Cretaceous">Cretaceous</a>, 90 Ma.<sup id="cite_ref-Hibbett1995_114-0" class="reference"><a href="#cite_note-Hibbett1995-114">[114]</a></sup><sup id="cite_ref-Hibbett1997_115-0" class="reference"><a href="#cite_note-Hibbett1997-115">[115]</a></sup></p>
<p>Some time after the <a href="/wiki/Permian%E2%80%93Triassic_extinction_event" title="Permian–Triassic extinction event">Permian–Triassic extinction event</a> (251.4 Ma), a fungal spike (originally thought to be an extraordinary abundance of fungal spores in <a href="/wiki/Sediment" title="Sediment">sediments</a>) formed, suggesting that fungi were the dominant life form at this time, representing nearly 100% of the available <a href="/wiki/Fossil_record" class="mw-redirect" title="Fossil record">fossil record</a> for this period.<sup id="cite_ref-Eshet1995_116-0" class="reference"><a href="#cite_note-Eshet1995-116">[116]</a></sup> However, the relative proportion of fungal spores relative to spores formed by <a href="/wiki/Algae" title="Algae">algal</a> species is difficult to assess,<sup id="cite_ref-Foster2002_117-0" class="reference"><a href="#cite_note-Foster2002-117">[117]</a></sup> the spike did not appear worldwide,<sup id="cite_ref-LopezGomez2005_118-0" class="reference"><a href="#cite_note-LopezGomez2005-118">[118]</a></sup><sup id="cite_ref-Looy2005_119-0" class="reference"><a href="#cite_note-Looy2005-119">[119]</a></sup> and in many places it did not fall on the Permian–Triassic boundary.<sup id="cite_ref-Ward2005_120-0" class="reference"><a href="#cite_note-Ward2005-120">[120]</a></sup></p>
<p>65 million years ago, immediately after the Cretaceous–Paleogene extinction that famously killed off most dinosaurs, there is a dramatic increase in evidence of fungi, apparently the death of most plant and animal species leading to a huge fungal bloom like "a massive compost heap".<sup id="cite_ref-121" class="reference"><a href="#cite_note-121">[121]</a></sup></p>
<h2><span class="mw-headline" id="Taxonomy">Taxonomy</span></h2>
<p>Although commonly included in botany curricula and textbooks, fungi are more closely related to <a href="/wiki/Animal" title="Animal">animals</a> than to plants and are placed with the animals in the <a href="/wiki/Monophyletic" class="mw-redirect" title="Monophyletic">monophyletic</a> group of <a href="/wiki/Opisthokont" title="Opisthokont">opisthokonts</a>.<sup id="cite_ref-ShalchianTabrizi2008_122-0" class="reference"><a href="#cite_note-ShalchianTabrizi2008-122">[122]</a></sup> Analyses using <a href="/wiki/Molecular_phylogenetics" title="Molecular phylogenetics">molecular phylogenetics</a> support a <a href="/wiki/Monophyletic_group" class="mw-redirect" title="Monophyletic group">monophyletic</a> origin of the Fungi.<sup id="cite_ref-Hibbett2007_41-1" class="reference"><a href="#cite_note-Hibbett2007-41">[41]</a></sup> The <a href="/wiki/Taxonomy_(biology)" title="Taxonomy (biology)">taxonomy</a> of the Fungi is in a state of constant flux, especially due to recent research based on DNA comparisons. These current phylogenetic analyses often overturn classifications based on older and sometimes less discriminative methods based on morphological features and biological species concepts obtained from experimental <a href="/wiki/Mating" title="Mating">matings</a>.<sup id="cite_ref-123" class="reference"><a href="#cite_note-123">[123]</a></sup></p>
<p>There is no unique generally accepted system at the higher taxonomic levels and there are frequent name changes at every level, from species upwards. Efforts among researchers are now underway to establish and encourage usage of a unified and more consistent <a href="/wiki/Botanical_nomenclature" title="Botanical nomenclature">nomenclature</a>.<sup id="cite_ref-Hibbett2007_41-2" class="reference"><a href="#cite_note-Hibbett2007-41">[41]</a></sup><sup id="cite_ref-Celio2006_124-0" class="reference"><a href="#cite_note-Celio2006-124">[124]</a></sup> Fungal species can also have multiple scientific names depending on their life cycle and mode (sexual or asexual) of reproduction. Web sites such as <a href="/wiki/Index_Fungorum" title="Index Fungorum">Index Fungorum</a> and <a href="/wiki/Integrated_Taxonomic_Information_System" title="Integrated Taxonomic Information System">ITIS</a> list current names of fungal species (with cross-references to older synonyms).</p>
<p>The 2007 classification of Kingdom Fungi is the result of a large-scale collaborative research effort involving dozens of mycologists and other scientists working on fungal taxonomy.<sup id="cite_ref-Hibbett2007_41-3" class="reference"><a href="#cite_note-Hibbett2007-41">[41]</a></sup> It recognizes seven <a href="/wiki/Phylum" title="Phylum">phyla</a>, two of which—the Ascomycota and the Basidiomycota—are contained within a branch representing <a href="/wiki/Subkingdom" class="mw-redirect" title="Subkingdom">subkingdom</a> Dikarya, the most species rich and familiar group, including all the mushrooms, most food-spoilage molds, most plant pathogenic fungi, and the beer, wine, and bread yeasts. The accompanying <a href="/wiki/Cladogram" title="Cladogram">cladogram</a> depicts the major fungal <a href="/wiki/Taxon" title="Taxon">taxa</a> and their relationship to opisthokont and unikont organisms, based on the work of Philippe Silar,<sup id="cite_ref-Silar2016_125-0" class="reference"><a href="#cite_note-Silar2016-125">[125]</a></sup> "The Mycota: A Comprehensive Treatise on Fungi as Experimental Systems for Basic and Applied Research"<sup id="cite_ref-MycotaVIIS&E_126-0" class="reference"><a href="#cite_note-MycotaVIIS&E-126">[126]</a></sup> and Tedersoo et al. 2018.<sup id="cite_ref-127" class="reference"><a href="#cite_note-127">[127]</a></sup> The lengths of the branches are not proportional to <a href="/wiki/Evolutionary" class="mw-redirect" title="Evolutionary">evolutionary</a> distances.</p>
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<td rowspan="2" style="border:0;padding:0;">
<p><i><a href="/w/index.php?title=Nucleophaga&action=edit&redlink=1" class="new" title="Nucleophaga (page does not exist)">Nucleophaga</a></i></p>
</td>
</tr>
<tr>
<td style="border:0;border-left:1px solid;vertical-align:top;text-align:center;"><br /></td>
</tr>
<tr>
<td style="border:0;padding:0 0.2em;border-left:1px solid;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><br /></td>
<td rowspan="2" style="border:0;padding:0;">
<p class="mw-empty-elt"></p>
<table style="border-spacing:0;margin:0;">
<tbody>
<tr>
<td style="width:1em;border:0;padding:0 0.2em;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><br /></td>
<td rowspan="2" style="border:0;padding:0;">
<p><a href="/wiki/Metchnikovellea" class="mw-redirect" title="Metchnikovellea">Metchnikovellea</a></p>
</td>
</tr>
<tr>
<td style="border:0;border-left:1px solid;vertical-align:top;text-align:center;"><br /></td>
</tr>
<tr>
<td style="border:0;padding:0 0.2em;border-left:1px solid;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><br /></td>
<td rowspan="2" style="border:0;padding:0;">
<p><a href="/wiki/Microsporea" class="mw-redirect" title="Microsporea">Microsporea</a></p>
</td>
</tr>
<tr>
<td style="border:0;vertical-align:top;text-align:center;"><br /></td>
</tr>
</tbody>
</table>
</td>
</tr>
<tr>
<td style="border:0;vertical-align:top;text-align:center;"><br /></td>
</tr>
</tbody>
</table>
</td>
</tr>
<tr>
<td style="border:0;vertical-align:top;text-align:center;"><br /></td>
</tr>
</tbody>
</table>
</td>
</tr>
<tr>
<td style="border:0;vertical-align:top;text-align:center;"><br /></td>
</tr>
</tbody>
</table>
</td>
</tr>
<tr>
<td style="border:0;vertical-align:top;text-align:center;"><br /></td>
</tr>
</tbody>
</table>
</td>
</tr>
<tr>
<td style="border:0;border-left:1px solid;vertical-align:top;text-align:center;"><br /></td>
</tr>
<tr>
<td style="border:0;padding:0 0.2em;border-left:1px solid;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><br /></td>
<td rowspan="2" style="border:0;padding:0;">
<p class="mw-empty-elt"></p>
<table style="border-spacing:0;margin:0;">
<tbody>
<tr>
<td style="width:1em;border:0;padding:0 0.2em;border-bottom:1px solid;vertical-align:bottom;text-align:center;">Aphelidiomyceta</td>
<td rowspan="2" style="border:0;padding:0;">
<p class="mw-empty-elt"></p>
<table style="border-spacing:0;margin:0;">
<tbody>
<tr>
<td style="width:1em;border:0;padding:0 0.2em;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><a href="/wiki/Aphelida" title="Aphelida">Aphelidiomycota</a></td>
<td rowspan="2" style="border:0;padding:0;">
<p><a href="/wiki/Aphelida" title="Aphelida">Aphelidiomycetes</a></p>
</td>
</tr>
<tr>
<td style="border:0;vertical-align:top;text-align:center;"><br /></td>
</tr>
</tbody>
</table>
</td>
</tr>
<tr>
<td style="border:0;border-left:1px solid;vertical-align:top;text-align:center;"><br /></td>
</tr>
<tr>
<td style="border:0;padding:0 0.2em;border-left:1px solid;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><a href="/wiki/Eumycota" class="mw-redirect" title="Eumycota">Eumycota</a></td>
<td rowspan="2" style="border:0;padding:0;">
<p class="mw-empty-elt"></p>
<table style="border-spacing:0;margin:0;">
<tbody>
<tr>
<td style="width:1em;border:0;padding:0 0.2em;border-bottom:1px solid;vertical-align:bottom;text-align:center;">Chytridiomyceta</td>
<td rowspan="2" style="border:0;padding:0;">
<p class="mw-empty-elt"></p>
<table style="border-spacing:0;margin:0;">
<tbody>
<tr>
<td style="width:1em;border:0;padding:0 0.2em;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><a href="/wiki/Neocallimastigomycota" title="Neocallimastigomycota">Neocallimastigomycota</a></td>
<td rowspan="2" style="border:0;padding:0;">
<p><a href="/wiki/Neocallimastigomycetes" class="mw-redirect" title="Neocallimastigomycetes">Neocallimastigomycetes</a></p>
</td>
</tr>
<tr>
<td style="border:0;border-left:1px solid;vertical-align:top;text-align:center;"><br /></td>
</tr>
<tr>
<td style="border:0;padding:0 0.2em;border-left:1px solid;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><a href="/wiki/Chytridiomycota" title="Chytridiomycota">Chytridiomycota</a></td>
<td rowspan="2" style="border:0;padding:0;">
<p class="mw-empty-elt"></p>
<table style="border-spacing:0;margin:0;">
<tbody>
<tr>
<td style="width:1em;border:0;padding:0 0.2em;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><a href="/w/index.php?title=Monoblepharomycotina&action=edit&redlink=1" class="new" title="Monoblepharomycotina (page does not exist)">Monoblepharomycotina</a></td>
<td rowspan="2" style="border:0;padding:0;">
<p class="mw-empty-elt"></p>
<table style="border-spacing:0;margin:0;">
<tbody>
<tr>
<td style="width:1em;border:0;padding:0 0.2em;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><br /></td>
<td rowspan="2" style="border:0;padding:0;">
<p><a href="/wiki/Hyaloraphidiomycetes" class="mw-redirect" title="Hyaloraphidiomycetes">Hyaloraphidiomycetes</a></p>
</td>
</tr>
<tr>
<td style="border:0;border-left:1px solid;vertical-align:top;text-align:center;"><br /></td>
</tr>
<tr>
<td style="border:0;padding:0 0.2em;border-left:1px solid;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><br /></td>
<td rowspan="2" style="border:0;padding:0;">
<p><a href="/wiki/Monoblepharidomycetes" title="Monoblepharidomycetes">Monoblepharidomycetes</a></p>
</td>
</tr>
<tr>
<td style="border:0;border-left:1px solid;vertical-align:top;text-align:center;"><br /></td>
</tr>
<tr>
<td style="border:0;padding:0 0.2em;border-left:1px solid;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><br /></td>
<td rowspan="2" style="border:0;padding:0;">
<p><a href="/w/index.php?title=Sanchytriomycetes&action=edit&redlink=1" class="new" title="Sanchytriomycetes (page does not exist)">Sanchytriomycetes</a></p>
</td>
</tr>
<tr>
<td style="border:0;vertical-align:top;text-align:center;"><br /></td>
</tr>
</tbody>
</table>
</td>
</tr>
<tr>
<td style="border:0;border-left:1px solid;vertical-align:top;text-align:center;"><br /></td>
</tr>
<tr>
<td style="border:0;padding:0 0.2em;border-left:1px solid;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><a href="/w/index.php?title=Chytridiomycotina&action=edit&redlink=1" class="new" title="Chytridiomycotina (page does not exist)">Chytridiomycotina</a></td>
<td rowspan="2" style="border:0;padding:0;">
<p class="mw-empty-elt"></p>
<table style="border-spacing:0;margin:0;">
<tbody>
<tr>
<td style="width:1em;border:0;padding:0 0.2em;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><br /></td>
<td rowspan="2" style="border:0;padding:0;">
<p><a href="/w/index.php?title=Mesochytriomycetes&action=edit&redlink=1" class="new" title="Mesochytriomycetes (page does not exist)">Mesochytriomycetes</a></p>
</td>
</tr>
<tr>
<td style="border:0;border-left:1px solid;vertical-align:top;text-align:center;"><br /></td>
</tr>
<tr>
<td style="border:0;padding:0 0.2em;border-left:1px solid;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><br /></td>
<td rowspan="2" style="border:0;padding:0;">
<p><a href="/wiki/Chytridiomycetes" title="Chytridiomycetes">Chytridiomycetes</a></p>
</td>
</tr>
<tr>
<td style="border:0;vertical-align:top;text-align:center;"><br /></td>
</tr>
</tbody>
</table>
</td>
</tr>
<tr>
<td style="border:0;vertical-align:top;text-align:center;"><br /></td>
</tr>
</tbody>
</table>
</td>
</tr>
<tr>
<td style="border:0;vertical-align:top;text-align:center;"><br /></td>
</tr>
</tbody>
</table>
</td>
</tr>
<tr>
<td style="border:0;border-left:1px solid;vertical-align:top;text-align:center;"><br /></td>
</tr>
<tr>
<td style="border:0;padding:0 0.2em;border-left:1px solid;border-bottom:1px solid;vertical-align:bottom;text-align:center;">Amastigomycota</td>
<td rowspan="2" style="border:0;padding:0;">
<p class="mw-empty-elt"></p>
<table style="border-spacing:0;margin:0;">
<tbody>
<tr>
<td style="width:1em;border:0;padding:0 0.2em;border-bottom:1px solid;vertical-align:bottom;text-align:center;">Blastocladiomyceta</td>
<td rowspan="2" style="border:0;padding:0;">
<p class="mw-empty-elt"></p>
<table style="border-spacing:0;margin:0;">
<tbody>
<tr>
<td style="width:1em;border:0;padding:0 0.2em;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><a href="/wiki/Blastocladiomycota" title="Blastocladiomycota">Blastocladiomycota</a></td>
<td rowspan="2" style="border:0;padding:0;">
<p class="mw-empty-elt"></p>
<table style="border-spacing:0;margin:0;">
<tbody>
<tr>
<td style="width:1em;border:0;padding:0 0.2em;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><br /></td>
<td rowspan="2" style="border:0;padding:0;">
<p><a href="/wiki/Blastocladiomycetes" class="mw-redirect" title="Blastocladiomycetes">Blastocladiomycetes</a></p>
</td>
</tr>
<tr>
<td style="border:0;border-left:1px solid;vertical-align:top;text-align:center;"><br /></td>
</tr>
<tr>
<td style="border:0;padding:0 0.2em;border-left:1px solid;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><br /></td>
<td rowspan="2" style="border:0;padding:0;">
<p><a href="/w/index.php?title=Physodermatomycetes&action=edit&redlink=1" class="new" title="Physodermatomycetes (page does not exist)">Physodermatomycetes</a></p>
</td>
</tr>
<tr>
<td style="border:0;vertical-align:top;text-align:center;"><br /></td>
</tr>
</tbody>
</table>
</td>
</tr>
<tr>
<td style="border:0;vertical-align:top;text-align:center;"><br /></td>
</tr>
</tbody>
</table>
</td>
</tr>
<tr>
<td style="border:0;border-left:1px solid;vertical-align:top;text-align:center;"><br /></td>
</tr>
<tr>
<td style="border:0;padding:0 0.2em;border-left:1px solid;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><br /></td>
<td rowspan="2" style="border:0;padding:0;">
<p class="mw-empty-elt"></p>
<table style="border-spacing:0;margin:0;">
<tbody>
<tr>
<td style="width:1em;border:0;padding:0 0.2em;border-bottom:1px solid;vertical-align:bottom;text-align:center;">Zoopagomyceta</td>
<td rowspan="2" style="border:0;padding:0;">
<p class="mw-empty-elt"></p>
<table style="border-spacing:0;margin:0;">
<tbody>
<tr>
<td style="width:1em;border:0;padding:0 0.2em;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><a href="/wiki/Basidiobolomycota" class="mw-redirect" title="Basidiobolomycota">Basidiobolomycota</a></td>
<td rowspan="2" style="border:0;padding:0;">
<p class="mw-empty-elt"></p>
<table style="border-spacing:0;margin:0;">
<tbody>
<tr>
<td style="width:1em;border:0;padding:0 0.2em;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><br /></td>
<td rowspan="2" style="border:0;padding:0;">
<p><a href="/wiki/Basidiobolomycetes" title="Basidiobolomycetes">Basidiobolomycetes</a></p>
</td>
</tr>
<tr>
<td style="border:0;border-left:1px solid;vertical-align:top;text-align:center;"><br /></td>
</tr>
<tr>
<td style="border:0;padding:0 0.2em;border-left:1px solid;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><br /></td>
<td rowspan="2" style="border:0;padding:0;">
<p><a href="/wiki/Olpidiomycetes" class="mw-redirect" title="Olpidiomycetes">Olpidiomycetes</a></p>
</td>
</tr>
<tr>
<td style="border:0;vertical-align:top;text-align:center;"><br /></td>
</tr>
</tbody>
</table>
</td>
</tr>
<tr>
<td style="border:0;border-left:1px solid;vertical-align:top;text-align:center;"><br /></td>
</tr>
<tr>
<td style="border:0;padding:0 0.2em;border-left:1px solid;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><br /></td>
<td rowspan="2" style="border:0;padding:0;">
<p class="mw-empty-elt"></p>
<table style="border-spacing:0;margin:0;">
<tbody>
<tr>
<td style="width:1em;border:0;padding:0 0.2em;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><a href="/wiki/Entomophthoromycota" title="Entomophthoromycota">Entomophthoromycota</a></td>
<td rowspan="2" style="border:0;padding:0;">
<p class="mw-empty-elt"></p>
<table style="border-spacing:0;margin:0;">
<tbody>
<tr>
<td style="width:1em;border:0;padding:0 0.2em;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><br /></td>
<td rowspan="2" style="border:0;padding:0;">
<p><a href="/wiki/Neozygitomycetes" class="mw-redirect" title="Neozygitomycetes">Neozygitomycetes</a></p>
</td>
</tr>
<tr>
<td style="border:0;border-left:1px solid;vertical-align:top;text-align:center;"><br /></td>
</tr>
<tr>
<td style="border:0;padding:0 0.2em;border-left:1px solid;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><br /></td>
<td rowspan="2" style="border:0;padding:0;">
<p><a href="/wiki/Entomophthoromycetes" class="mw-redirect" title="Entomophthoromycetes">Entomophthoromycetes</a></p>
</td>
</tr>
<tr>
<td style="border:0;vertical-align:top;text-align:center;"><br /></td>
</tr>
</tbody>
</table>
</td>
</tr>
<tr>
<td style="border:0;border-left:1px solid;vertical-align:top;text-align:center;"><br /></td>
</tr>
<tr>
<td style="border:0;padding:0 0.2em;border-left:1px solid;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><a href="/w/index.php?title=Kickxellomycota&action=edit&redlink=1" class="new" title="Kickxellomycota (page does not exist)">Kickxellomycota</a></td>
<td rowspan="2" style="border:0;padding:0;">
<p class="mw-empty-elt"></p>
<table style="border-spacing:0;margin:0;">
<tbody>
<tr>
<td style="width:1em;border:0;padding:0 0.2em;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><a href="/wiki/Zoopagomycotina" title="Zoopagomycotina">Zoopagomycotina</a></td>
<td rowspan="2" style="border:0;padding:0;">
<p><a href="/wiki/Zoopagomycetes" class="mw-redirect" title="Zoopagomycetes">Zoopagomycetes</a></p>
</td>
</tr>
<tr>
<td style="border:0;border-left:1px solid;vertical-align:top;text-align:center;"><br /></td>
</tr>
<tr>
<td style="border:0;padding:0 0.2em;border-left:1px solid;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><a href="/wiki/Kickxellomycotina" title="Kickxellomycotina">Kickxellomycotina</a></td>
<td rowspan="2" style="border:0;padding:0;">
<p class="mw-empty-elt"></p>
<table style="border-spacing:0;margin:0;">
<tbody>
<tr>
<td style="width:1em;border:0;padding:0 0.2em;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><br /></td>
<td rowspan="2" style="border:0;padding:0;">
<p><a href="/w/index.php?title=Dimargaritomycetes&action=edit&redlink=1" class="new" title="Dimargaritomycetes (page does not exist)">Dimargaritomycetes</a></p>
</td>
</tr>
<tr>
<td style="border:0;border-left:1px solid;vertical-align:top;text-align:center;"><br /></td>
</tr>
<tr>
<td style="border:0;padding:0 0.2em;border-left:1px solid;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><br /></td>
<td rowspan="2" style="border:0;padding:0;">
<p><a href="/wiki/Kickxellomycetes" class="mw-redirect" title="Kickxellomycetes">Kickxellomycetes</a></p>
</td>
</tr>
<tr>
<td style="border:0;vertical-align:top;text-align:center;"><br /></td>
</tr>
</tbody>
</table>
</td>
</tr>
<tr>
<td style="border:0;vertical-align:top;text-align:center;"><br /></td>
</tr>
</tbody>
</table>
</td>
</tr>
<tr>
<td style="border:0;vertical-align:top;text-align:center;"><br /></td>
</tr>
</tbody>
</table>
</td>
</tr>
<tr>
<td style="border:0;vertical-align:top;text-align:center;"><br /></td>
</tr>
</tbody>
</table>
</td>
</tr>
<tr>
<td style="border:0;border-left:1px solid;vertical-align:top;text-align:center;"><br /></td>
</tr>
<tr>
<td style="border:0;padding:0 0.2em;border-left:1px solid;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><br /></td>
<td rowspan="2" style="border:0;padding:0;">
<p class="mw-empty-elt"></p>
<table style="border-spacing:0;margin:0;">
<tbody>
<tr>
<td style="width:1em;border:0;padding:0 0.2em;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><a href="/w/index.php?title=Mortierellomycota&action=edit&redlink=1" class="new" title="Mortierellomycota (page does not exist)">Mortierellomycota</a></td>
<td rowspan="2" style="border:0;padding:0;">
<p><a href="/wiki/Mortierellomycetes" class="mw-redirect" title="Mortierellomycetes">Mortierellomycetes</a></p>
</td>
</tr>
<tr>
<td style="border:0;border-left:1px solid;vertical-align:top;text-align:center;"><br /></td>
</tr>
<tr>
<td style="border:0;padding:0 0.2em;border-left:1px solid;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><br /></td>
<td rowspan="2" style="border:0;padding:0;">
<p class="mw-empty-elt"></p>
<table style="border-spacing:0;margin:0;">
<tbody>
<tr>
<td style="width:1em;border:0;padding:0 0.2em;border-bottom:1px solid;vertical-align:bottom;text-align:center;">Mucoromyceta</td>
<td rowspan="2" style="border:0;padding:0;">
<p class="mw-empty-elt"></p>
<table style="border-spacing:0;margin:0;">
<tbody>
<tr>
<td style="width:1em;border:0;padding:0 0.2em;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><a href="/w/index.php?title=Calcarisporiellomycota&action=edit&redlink=1" class="new" title="Calcarisporiellomycota (page does not exist)">Calcarisporiellomycota</a></td>
<td rowspan="2" style="border:0;padding:0;">
<p><a href="/w/index.php?title=Calcarisporiellomycetes&action=edit&redlink=1" class="new" title="Calcarisporiellomycetes (page does not exist)">Calcarisporiellomycetes</a></p>
</td>
</tr>
<tr>
<td style="border:0;border-left:1px solid;vertical-align:top;text-align:center;"><br /></td>
</tr>
<tr>
<td style="border:0;padding:0 0.2em;border-left:1px solid;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><a href="/w/index.php?title=Mucoromycota&action=edit&redlink=1" class="new" title="Mucoromycota (page does not exist)">Mucoromycota</a></td>
<td rowspan="2" style="border:0;padding:0;">
<p class="mw-empty-elt"></p>
<table style="border-spacing:0;margin:0;">
<tbody>
<tr>
<td style="width:1em;border:0;padding:0 0.2em;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><br /></td>
<td rowspan="2" style="border:0;padding:0;">
<p><a href="/w/index.php?title=Umbelopsidomycetes&action=edit&redlink=1" class="new" title="Umbelopsidomycetes (page does not exist)">Umbelopsidomycetes</a></p>
</td>
</tr>
<tr>
<td style="border:0;border-left:1px solid;vertical-align:top;text-align:center;"><br /></td>
</tr>
<tr>
<td style="border:0;padding:0 0.2em;border-left:1px solid;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><br /></td>
<td rowspan="2" style="border:0;padding:0;">
<p><a href="/wiki/Mucoromycetes" class="mw-redirect" title="Mucoromycetes">Mucoromycetes</a></p>
</td>
</tr>
<tr>
<td style="border:0;vertical-align:top;text-align:center;"><br /></td>
</tr>
</tbody>
</table>
</td>
</tr>
<tr>
<td style="border:0;vertical-align:top;text-align:center;"><br /></td>
</tr>
</tbody>
</table>
</td>
</tr>
<tr>
<td style="border:0;border-left:1px solid;vertical-align:top;text-align:center;"><br /></td>
</tr>
<tr>
<td style="border:0;padding:0 0.2em;border-left:1px solid;border-bottom:1px solid;vertical-align:bottom;text-align:center;">Symbiomycota</td>
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<p class="mw-empty-elt"></p>
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<td style="width:1em;border:0;padding:0 0.2em;border-bottom:1px solid;vertical-align:bottom;text-align:center;"><a href="/wiki/Glomeromycota" title="Glomeromycota">Glomeromycota</a></td>
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