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font-size: 100%"> <tbody><tr> <th colspan="2" style="color:inherit; text-align: center; background-color: rgb(235,235,210)">Cloudinidae<br /><div style="font-size: 85%;">Temporal range: <span class="noprint"><span style="display:inline-block;"></span><span style="display:inline-block;">555–529 <a href="/wiki/Megaannum" class="mw-redirect" title="Megaannum">Ma</a></span><sup id="cite_ref-Yang2016_1-0" class="reference"><a href="#cite_note-Yang2016-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> <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; white-space:nowrap; border:1px #666; border-style:solid none; position:relative; z-index:0; font-size:97%;"> <div style="position:absolute; height:100%; left:0px; width:207.23076923077px; padding-left:5px; text-align:left; background-color:rgb(254,217,106); 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">PreꞒ</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(127,160,86); left:37.636923076923px; width:17.583076923077px;"><a href="/wiki/Cambrian" title="Cambrian">Ꞓ</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(0,146,112); left:55.22px; width:14.807692307692px;"><a href="/wiki/Ordovician" title="Ordovician">O</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(179,225,182); left:70.027692307692px; width:7.9470769230769px;"><a href="/wiki/Silurian" title="Silurian">S</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(203,140,55); left:77.974769230769px; width:20.564923076923px;"><a href="/wiki/Devonian" title="Devonian">D</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(103,165,153); left:98.539692307692px; width:20.294153846154px;"><a href="/wiki/Carboniferous" title="Carboniferous">C</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(240,64,40); 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(129,43,146); left:134.74086153846px; width:17.092984615385px;"><a href="/wiki/Triassic" title="Triassic">T</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(52,178,201); left:151.83384615385px; width:19.732307692308px;"><a href="/wiki/Jurassic" title="Jurassic">J</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(127,198,78); left:171.56615384615px; width:26.095384615385px;"><a href="/wiki/Cretaceous" title="Cretaceous">K</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(253,154,82); left:197.66153846154px; width:14.540307692308px;"><a href="/wiki/Paleogene" title="Paleogene">Pg</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(255,230,25); left:212.20184615385px; width:6.9249230769231px;"><a href="/wiki/Neogene" title="Neogene">N</a></div> <div id="end-border" style="position:absolute; height:100%; background-color:#666; width:1px; left:219px"></div><div style="margin:0 auto; line-height:0; clear:both; width:220px; padding:0px; height:8px; overflow:visible; background-color:transparent; position:relative; top:-4px; z-index:100;"><div style="position:absolute; height:8px; left:32.153846153846px; width:8.8px; background-color:#360; opacity:0.42;"></div> <div style="position:absolute; height:8px; left:32.153846153846px; width:8.8px; background-color:#360; opacity:1;"></div> <div style="position:absolute; height:6px; top:1px; left:33.153846153846px; width:6.8px; background-color:#6c3;"></div> </div> </div></span></div> </th></tr> <tr> <td colspan="2" style="text-align: center"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/wiki/File:Cloudina.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/14/Cloudina.svg/110px-Cloudina.svg.png" decoding="async" width="110" height="309" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/14/Cloudina.svg/165px-Cloudina.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/14/Cloudina.svg/220px-Cloudina.svg.png 2x" data-file-width="264" data-file-height="741" /></a></span> </td></tr> <tr> <th colspan="2" style="color:inherit; min-width:15em; text-align: center; background-color: rgb(235,235,210)"><a href="/wiki/Taxonomy_(biology)" title="Taxonomy (biology)">Scientific classification</a> <span class="plainlinks taxobox-edit-taxonomy skin-invert" style="font-size:smaller; float:right; padding-right:0.4em; margin-left:-3em;"><span typeof="mw:File"><a href="/wiki/Template:Taxonomy/Cloudinidae" title="Edit this classification"><img alt="Edit this classification" src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/15px-OOjs_UI_icon_edit-ltr.svg.png" decoding="async" width="15" height="15" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/23px-OOjs_UI_icon_edit-ltr.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/30px-OOjs_UI_icon_edit-ltr.svg.png 2x" data-file-width="20" data-file-height="20" /></a></span></span> </th></tr> <tr> <td>Domain: </td> <td><a href="/wiki/Eukaryote" title="Eukaryote">Eukaryota</a> </td></tr> <tr> <td>Kingdom: </td> <td><a href="/wiki/Animal" title="Animal">Animalia</a> </td></tr> <tr> <td>Phylum: </td> <td><a href="/wiki/Incertae_sedis" title="Incertae sedis"><i>incertae sedis</i></a> </td></tr> <tr> <td>Family: </td> <td><span style="font-style:normal;font-weight:normal;">†</span><a class="mw-selflink selflink">Cloudinidae</a><br /><small>Hahn and Pflug, 1985</small> </td></tr> <tr> <th colspan="2" style="color:inherit; text-align: center; background-color: rgb(235,235,210)">Species </th></tr> <tr> <td colspan="2" style="text-align: left"> <ul style="margin-left: 0"><li style="width:100%; clear:both; list-style: none; padding-left:0.2em;"> <i>Cloudina </i> <small> Germs, 1972 </small></li><div style="padding-left:0.5em;"><ul class="taxonlist"><li><i>Cl. hartmannae</i> <small>Germs 1972 </small></li><li><i>Cl. riemkeae</i> <small>Germs 1972 </small></li><li><i>Cl. lucianoi</i> <small>Beurlen & Sommer, 1957) Zaine & Fairchild, 1985 </small></li><li><i>Cl. sinensis</i> <small>Zhang, Li et Dung, 1992 </small></li><li><i>Cl. carinata</i> <small>Cortijo, Musa, Jensena et Palacios, 2009 </small></li><li><i>Cl. ningqiangensis</i> <small>Cai <i>et al.</i>, 2017 </small></li><li><i>Cl. xuanjiangpingensis</i> <small>Cai <i>et al.</i>, 2017 </small></li></ul> </div><li style="width:100%; clear:both; list-style: none; padding-left:0.2em;"> <i>Conotubus</i> <small> Zhang and Lin 1986 </small></li><div style="padding-left:0.5em;"><ul class="taxonlist"><li><i>Co. hemiannulatus</i> <small>Zhang and Lin 1986 </small></li><li><i><span style="font-style:normal;">(</span>others?<span style="font-style:normal;">)</span></i></li></ul> </div><li style="width:100%; clear:both; list-style: none; padding-left:0.2em;"><i>Acuticocloudina</i> <small>Hahn and Pflug, 1985 </small></li><div style="padding-left:0.5em;"></div></ul> </td></tr> <tr> <th colspan="2" style="color:inherit; text-align: center; background-color: rgb(235,235,210)"><a href="/wiki/Synonym_(taxonomy)" title="Synonym (taxonomy)">Synonyms</a> </th></tr> <tr> <td colspan="2" style="text-align: left"> <ul><li><i>Aulophycus lucianoi</i> <small>Beurlen & Sommer 1957</small> = <i>C. waldei</i> <small>Hahn & Pflug, 1985</small> = <i><b>C. lucianoi</b></i> <small>Zaine & Fairchild, 1985</small></li></ul> </td></tr> </tbody></table><style data-mw-deduplicate="TemplateStyles:r1238732961">@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox.biota tr{background:transparent!important}html.skin-theme-clientpref-night .mw-parser-output .infobox.biota img{background:transparent}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox.biota tr{background:transparent!important}html.skin-theme-clientpref-os .mw-parser-output .infobox.biota img{background:white}}.mw-parser-output .infobox.biota .taxobox-edit-taxonomy img{background:transparent!important}body.skin-vector .mw-parser-output table.biota.infobox{margin-top:0.5em}body.skin--responsive .mw-parser-output table.biota.infobox tr.taxonrow td{padding:2px 10px}</style> <p>The <b>cloudinids</b>, an early <a href="/wiki/Metazoan" class="mw-redirect" title="Metazoan">metazoan</a> <a href="/wiki/Family_(biology)" title="Family (biology)">family</a> containing the <a href="/wiki/Genus" title="Genus">genera</a> <i><b>Acuticocloudina</b></i>, <i><b>Cloudina</b></i> and <i><b>Conotubus</b></i>, lived in the late <a href="/wiki/Ediacaran" title="Ediacaran">Ediacaran</a> <a href="/wiki/Period_(geology)" class="mw-redirect" title="Period (geology)">period</a> about 550 million years ago<sup id="cite_ref-NYT-20200110_2-0" class="reference"><a href="#cite_note-NYT-20200110-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-NAT-20200110_3-0" class="reference"><a href="#cite_note-NAT-20200110-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> and became extinct at the base of the <a href="/wiki/Cambrian" title="Cambrian">Cambrian</a>.<sup id="cite_ref-Yang2016_1-1" class="reference"><a href="#cite_note-Yang2016-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> They formed millimetre-scale conical <a href="/wiki/Fossil" title="Fossil">fossils</a> consisting of <a href="/wiki/Calcareous" title="Calcareous">calcareous</a> cones nested within one another; the appearance of the organism itself remains unknown. The name <i>Cloudina</i> honors the 20th-century geologist and paleontologist <a href="/wiki/Preston_Cloud" title="Preston Cloud">Preston Cloud</a>.<sup id="cite_ref-Description_4-0" class="reference"><a href="#cite_note-Description-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p><p>Cloudinids comprise two genera: <i>Cloudina</i> itself is mineralized, whereas <i>Conotubus</i> is at best weakly mineralized, whilst sharing the same "funnel-in-funnel" construction.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>Cloudinids had a wide geographic range, reflected in the present distribution of localities in which their fossils are found, and are an abundant component of some deposits. They never appear in the same layers as soft-bodied <a href="/wiki/Ediacaran_biota" title="Ediacaran biota">Ediacaran biota</a>, but the fact that some sequences contain cloudinids and Ediacaran biota in alternating layers suggests that these groups had different environmental preferences. It has been suggested that cloudinids lived embedded in <a href="/wiki/Microbial_mat" title="Microbial mat">microbial mats</a>, growing new cones to avoid being buried by silt. However no specimens have been found embedded in mats, and their mode of life is still an unresolved question. </p><p>The <a href="/wiki/Taxonomy_(biology)" title="Taxonomy (biology)">classification</a> of the cloudinids has proved difficult: they were initially regarded as <a href="/wiki/Polychaete" title="Polychaete">polychaete</a> worms, and then as coral-like <a href="/wiki/Cnidarian" class="mw-redirect" title="Cnidarian">cnidarians</a> on the basis of what look like <a href="/wiki/Budding" title="Budding">buds</a> on some specimens. Current scientific opinion is divided between classifying them as polychaetes and regarding it as unsafe to classify them as members of any broader grouping. In 2020, a new study of pyritized specimens from the <a href="/wiki/Wood_Canyon_Formation" title="Wood Canyon Formation">Wood Canyon Formation</a> in Nevada showed the presence of <a href="/wiki/Nephrozoa" title="Nephrozoa">Nephrozoan</a> type <a href="/wiki/Gastrointestinal_tract" title="Gastrointestinal tract">guts</a>, the oldest on record, supporting the <a href="/wiki/Bilateria" title="Bilateria">bilaterian</a> interpretation.<sup id="cite_ref-NAT-20200110_3-1" class="reference"><a href="#cite_note-NAT-20200110-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p><p>Cloudinids are important in the history of animal evolution for two reasons. They are among the earliest and most abundant of the <a href="/wiki/Small_shelly_fossils" class="mw-redirect" title="Small shelly fossils">small shelly fossils</a> with <a href="/wiki/Mineralization_(biology)" class="mw-redirect" title="Mineralization (biology)">mineralized</a> <a href="/wiki/Skeleton" title="Skeleton">skeletons</a>, and therefore feature in the debate about why such skeletons first appeared in the Late Ediacaran. The most widely supported answer is that their shells are a defense against predators, as some <i>Cloudina</i> specimens from China bear the marks of multiple attacks, which suggests they survived at least a few of them. The holes made by predators are approximately proportional to the size of the <i>Cloudina</i> specimens, and <i><a href="/wiki/Sinotubulites" title="Sinotubulites">Sinotubulites</a></i> fossils, which are often found in the same beds, have so far shown no such holes. These two points suggest that predators attacked in a selective manner, and the <a href="/wiki/Evolutionary_arms_race" title="Evolutionary arms race">evolutionary arms race</a> which this indicates is commonly cited as a cause of the <a href="/wiki/Cambrian_explosion" title="Cambrian explosion">Cambrian explosion</a> of animal <a href="/wiki/Biodiversity" title="Biodiversity">diversity</a> and complexity. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Morphology">Morphology</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cloudinidae&action=edit&section=1" title="Edit section: Morphology"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Cloudina_internal_01.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0e/Cloudina_internal_01.png/100px-Cloudina_internal_01.png" decoding="async" width="100" height="217" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/0/0e/Cloudina_internal_01.png 1.5x" data-file-width="133" data-file-height="289" /></a><figcaption> Cutaway diagram of <i>Cloudina</i> showing "living space" within the shell.</figcaption></figure> <p><i>Cloudina</i> varies in size from a diameter of 0.3 to 6.5 mm, and 8 to 150 mm in length.<sup id="cite_ref-Description_4-1" class="reference"><a href="#cite_note-Description-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Fossils consist of a series of stacked vase-like <a href="/wiki/Calcite" title="Calcite">calcite</a> tubes, whose original mineral composition is unknown,<sup id="cite_ref-Porter2007_6-0" class="reference"><a href="#cite_note-Porter2007-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> but inferred to be high-magnesium calcite.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> Each cone traps a significant pore space beneath it, and stacks eccentrically in the one below. This results in a ridged external appearance. The overall tube is curved or sinuous, and occasionally branches. The tube walls are 8 to 50 micrometers thick, usually lying in the range 10 to 25 μm.<sup id="cite_ref-Grant1990_8-0" class="reference"><a href="#cite_note-Grant1990-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> Although it used to be thought that the tubes had test-tube like bases,<sup id="cite_ref-Description_4-2" class="reference"><a href="#cite_note-Description-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> detailed three-dimensional reconstruction has shown that the tubes had an open base.<sup id="cite_ref-Miller_9-0" class="reference"><a href="#cite_note-Miller-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> There is evidence that the tube was flexible.<sup id="cite_ref-Brain2001_10-0" class="reference"><a href="#cite_note-Brain2001-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Classification">Classification</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cloudinidae&action=edit&section=2" title="Edit section: Classification"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><i>Cloudina</i> was originally classified in 1972 as a member of the Cribricyathea, a class known from the Early Cambrian.<sup id="cite_ref-Description_4-3" class="reference"><a href="#cite_note-Description-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Glaessner (1976) accepted this classification and also proposed that <i>Cloudina</i> was similar to the <a href="/wiki/Annelid" title="Annelid">annelid</a> worms, particularly <a href="/wiki/Serpulid" class="mw-redirect" title="Serpulid">serpulid</a> <a href="/wiki/Polychaete" title="Polychaete">polychaetes</a>.<sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> However, Hahn & Pflug (1985) and Conway Morris <i>et al.</i>. (1990) doubted both Germs' and Glaessner's suggested relationships, and were unwilling to classify it to anything more than its own <a href="/wiki/Family_(biology)" title="Family (biology)">family</a>, Cloudinidae.<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Conway1990_13-0" class="reference"><a href="#cite_note-Conway1990-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> Some specimens of <i>Cloudina hartmannae</i> display budding,<sup id="cite_ref-Description_4-4" class="reference"><a href="#cite_note-Description-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> which implies asexual reproduction.<sup id="cite_ref-Hua2005_14-0" class="reference"><a href="#cite_note-Hua2005-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> On this basis Grant (1990) classified <i>Cloudina</i> as a coral-like <a href="/wiki/Cnidarian" class="mw-redirect" title="Cnidarian">cnidarian</a>.<sup id="cite_ref-Grant1990_8-1" class="reference"><a href="#cite_note-Grant1990-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> Since the tubes had an open base, creating a single living space rather than a series of separate chambers, <i>Cloudina</i> is more likely to be a <a href="/wiki/Stem_group" class="mw-redirect" title="Stem group">stem group</a> polychaete worm,<sup id="cite_ref-Miller_9-1" class="reference"><a href="#cite_note-Miller-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> in other words an evolutionary "aunt" or "cousin" of more recent polychaetes. This interpretation is reinforced by the even distribution of bore-holes made by predators.<sup id="cite_ref-HUA2003_15-0" class="reference"><a href="#cite_note-HUA2003-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bengtson1992_16-0" class="reference"><a href="#cite_note-Bengtson1992-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> However, as with so many <a href="/wiki/Ediacaran" title="Ediacaran">Ediacaran</a> life forms, there is great debate surrounding its position in the tree of life, and classification between the <a href="/wiki/Kingdom_(biology)" title="Kingdom (biology)">kingdom</a> and family level may be unwise.<sup id="cite_ref-Grant1990_8-2" class="reference"><a href="#cite_note-Grant1990-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Conway1990_13-1" class="reference"><a href="#cite_note-Conway1990-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-VinnZaton2012_17-0" class="reference"><a href="#cite_note-VinnZaton2012-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Ecology">Ecology</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cloudinidae&action=edit&section=3" title="Edit section: Ecology"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Cloudina_NT.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/28/Cloudina_NT.jpg/220px-Cloudina_NT.jpg" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/28/Cloudina_NT.jpg/330px-Cloudina_NT.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/28/Cloudina_NT.jpg/440px-Cloudina_NT.jpg 2x" data-file-width="800" data-file-height="600" /></a><figcaption> Restoration of <i>Cloudina hartmannae</i> with speculative mouth parts</figcaption></figure> <p><i>Cloudina</i> is frequently found in association with <a href="/wiki/Stromatolites" class="mw-redirect" title="Stromatolites">stromatolites</a>, which are limited to shallow water; their <a href="/wiki/Isotope_analysis" title="Isotope analysis">isotopic composition</a><sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> suggests that water temperatures were relatively cool. They have also been found in normal sea-floor sediments, suggesting that they were not only restricted to dwelling on microbial mounds.<sup id="cite_ref-domke2009_19-0" class="reference"><a href="#cite_note-domke2009-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> On the other hand, <i>Cloudina</i> has never been found in the same layers as the soft-bodied <a href="/wiki/Ediacara_biota" class="mw-redirect" title="Ediacara biota">Ediacara biota</a>, but <i>Cloudina</i> and Ediacara biota have been found in alternating layers. This suggests that the two groups of organisms had different environmental preferences.<sup id="cite_ref-Miller_9-2" class="reference"><a href="#cite_note-Miller-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p><p>In many <i>Cloudina</i> specimens the ridges formed by the cones are of varying width, which suggests the organisms grew at a variable rate. <a href="/wiki/Adolf_Seilacher" title="Adolf Seilacher">Adolf Seilacher</a> suggests that they adhered to <a href="/wiki/Microbial_mat" title="Microbial mat">microbial mats</a>, and that the growth phases represented the organism keeping pace with sedimentation—growing through new material deposited on it that would otherwise bury it. Kinks in the developing tube are easily explained by the mat falling slightly from the horizontal.<sup id="cite_ref-Seilacher1999_20-0" class="reference"><a href="#cite_note-Seilacher1999-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> Because of its small size, <i>Cloudina</i> would be expected to be found <i>in situ</i> in the microbial mat, especially if, as Seilacher suggests, sedimentation built up around it during its lifetime. But all the many specimens discovered to date have only been found having been washed out of their places of growth. A further argument against Seilacher's hypothesis is that the predatory borings found in many specimens are not concentrated at what would be the top end, as one would expect if the animal was mainly buried. An alternative is that the organism dwelt on seaweeds,<sup id="cite_ref-Miller_9-3" class="reference"><a href="#cite_note-Miller-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> but until a specimen unquestionably <i>in situ</i> is discovered, its mode of life remains open to debate. </p><p>The tubes often appear to form colonies, although they are sometimes found in more isolated situations. The frequent appearance of large and sometimes single-species colonies has been attributed to the lack of significant predation.<sup id="cite_ref-Description_4-5" class="reference"><a href="#cite_note-Description-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> On the other hand, in some locations up to 20% of <i>Cloudina</i> fossils contain <a href="/wiki/Predator" class="mw-redirect" title="Predator">predatory</a> borings ranging from 15 to 400 μm in diameter.<sup id="cite_ref-HUA2003_15-1" class="reference"><a href="#cite_note-HUA2003-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bengtson1992_16-1" class="reference"><a href="#cite_note-Bengtson1992-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> The boreholes are rather evenly distributed along the tube length, and some tubes had been bored multiple times—hence the organism could survive attacks, since predators do not attack empty shells. This may indicate that the animal could vary its position in the tube in response to predation, or that it occupied the full length—but not the full width—of the tube. The even distribution is perhaps difficult to reconcile with an <a href="/wiki/Infauna" class="mw-redirect" title="Infauna">infaunal</a> lifestyle, mainly buried in a microbial mat, and adds weight to Miller's suggestion that the animal lived on seaweeds or in a reef environment. If modern-day molluscs are a suitable analogy, the size distribution of the borings suggests that the predator was similar in size to <i>Cloudina</i>.<sup id="cite_ref-Brain2001_10-1" class="reference"><a href="#cite_note-Brain2001-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> </p><p>Fossil findings in the <a href="/wiki/Nama_Group" title="Nama Group">Nama Group</a>, <a href="/wiki/Namibia" title="Namibia">Namibia</a>, suggest that <i>Cloudina</i> was one of the first reef-building animals,<sup id="cite_ref-natnews2014_21-0" class="reference"><a href="#cite_note-natnews2014-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-penny2014_22-0" class="reference"><a href="#cite_note-penny2014-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> but machine-learning facilitated 3D tomography indicates that the 'reef-forming' fossils are in fact simply aggregations of solitary individuals.<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Fossil_locations">Fossil locations</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cloudinidae&action=edit&section=4" title="Edit section: Fossil locations"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><i>Cloudina</i> occurred in <a href="/wiki/Calcium_carbonate" title="Calcium carbonate">calcium carbonate</a> rich areas of <a href="/wiki/Stromatolite" title="Stromatolite">stromatolite</a> reefs. It is found in association with <i><a href="/wiki/Namacalathus" title="Namacalathus">Namacalathus</a></i>, which like <i>Cloudina</i> was "weakly skeletal" and solitary, and <i><a href="/wiki/Namapoikia" title="Namapoikia">Namapoikia</a></i>, which was "robustly skeletal" and formed sheets on open surfaces.<sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> </p><p>First found in the <a href="/wiki/Nama_Group" title="Nama Group">Nama Group</a> in <a href="/wiki/Namibia" title="Namibia">Namibia</a>,<sup id="cite_ref-Description_4-6" class="reference"><a href="#cite_note-Description-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> <i>Cloudina</i> has also been reported in <a href="/wiki/Oman" title="Oman">Oman</a>,<sup id="cite_ref-Conway1990_13-2" class="reference"><a href="#cite_note-Conway1990-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> <a href="/wiki/China" title="China">China</a>'s <a href="/wiki/Dengying_Formation" title="Dengying Formation">Dengying Formation</a>,<sup id="cite_ref-Conway1990_13-3" class="reference"><a href="#cite_note-Conway1990-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Bengtson1992_16-2" class="reference"><a href="#cite_note-Bengtson1992-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Canada" title="Canada">Canada</a>,<sup id="cite_ref-Hofmann2001_25-0" class="reference"><a href="#cite_note-Hofmann2001-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Uruguay" title="Uruguay">Uruguay</a>,<sup id="cite_ref-Gaucher1998_26-0" class="reference"><a href="#cite_note-Gaucher1998-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Gaucher2000_27-0" class="reference"><a href="#cite_note-Gaucher2000-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Argentina" title="Argentina">Argentina</a>,<sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Antarctica" title="Antarctica">Antarctica</a>,<sup id="cite_ref-Yochelson1977_29-0" class="reference"><a href="#cite_note-Yochelson1977-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Brazil" title="Brazil">Brazil</a>,<sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Zaine1985_31-0" class="reference"><a href="#cite_note-Zaine1985-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Nevada" title="Nevada">Nevada</a>,<sup id="cite_ref-HAGADORN2000_32-0" class="reference"><a href="#cite_note-HAGADORN2000-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> central <a href="/wiki/Spain" title="Spain">Spain</a>, northwest <a href="/wiki/Mexico" title="Mexico">Mexico</a> and <a href="/wiki/California" title="California">California</a>,<sup id="cite_ref-Grant1990_8-3" class="reference"><a href="#cite_note-Grant1990-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> in west and south <a href="/wiki/Siberia" title="Siberia">Siberia</a>. The <i>Cloudina</i> fossils found in association with late Precambrian-Early Cambrian <a href="/wiki/Anabarites" title="Anabarites">anabaritids</a> SSF and tubular agglutinated skeletal fossils <i><a href="/wiki/Platysolenites" title="Platysolenites">Platysolenites</a></i> and <i><a href="/w/index.php?title=Spirosolenites&action=edit&redlink=1" class="new" title="Spirosolenites (page does not exist)">Spirosolenites</a></i> in Siberia.<sup id="cite_ref-Kontorovich2008_33-0" class="reference"><a href="#cite_note-Kontorovich2008-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Zhuravlev_2009_34-0" class="reference"><a href="#cite_note-Zhuravlev_2009-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Paleontological_importance">Paleontological importance</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cloudinidae&action=edit&section=5" title="Edit section: Paleontological importance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Although not the first <a href="/wiki/Small_shelly_fossils" class="mw-redirect" title="Small shelly fossils">small shelly fossil</a> to be found, <i>Cloudina</i> is one of the earliest and most abundant.<sup id="cite_ref-Bengtson2004_35-0" class="reference"><a href="#cite_note-Bengtson2004-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> The evolution of external shells in the Late <a href="/wiki/Ediacaran" title="Ediacaran">Ediacaran</a> is thought to be a defence against predators, marking the start of an evolutionary arms race.<sup id="cite_ref-Bengtson2004_35-1" class="reference"><a href="#cite_note-Bengtson2004-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Dzik2007VerdunSyndrome_36-0" class="reference"><a href="#cite_note-Dzik2007VerdunSyndrome-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> While predatory borings are common in <i>Cloudina</i> specimens, no such borings have been found in <i><a href="/wiki/Sinotubulites" title="Sinotubulites">Sinotubulites</a></i>, a similar shelly fossil sometimes found in the same beds. In addition, the diameters of borings in <i>Cloudina</i> are proportional to the sizes of specimens, which suggests that predators were selective about the size of their prey. These two indications that predators attacked selectively suggest the possibility of <a href="/wiki/Speciation" title="Speciation">speciation</a> in response to predation, which is often postulated as a potential cause of the <a href="/wiki/Cambrian_explosion" title="Cambrian explosion">rapid diversification of animals in the Early Cambrian</a>.<sup id="cite_ref-Bengtson1992_16-3" class="reference"><a href="#cite_note-Bengtson1992-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cloudinidae&action=edit&section=6" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><i><a href="/wiki/Anabarites" title="Anabarites">Anabarites</a></i></li> <li><i><a href="/wiki/Corumbella" title="Corumbella">Corumbella</a></i></li> <li><i><a href="/wiki/Saarina" title="Saarina">Saarina</a></i></li> <li><i><a href="/wiki/Sinotubulites" title="Sinotubulites">Sinotubulites</a></i></li> <li><i><a href="/wiki/Somatohelix" title="Somatohelix">Somatohelix</a></i></li> <li><a href="/wiki/List_of_Ediacaran_genera" title="List of Ediacaran genera">List of Ediacaran genera</a></li></ul> <p><br /> </p> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Cloudinidae&action=edit&section=7" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width" style="column-width: 25em;"> <ol class="references"> <li id="cite_note-Yang2016-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-Yang2016_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Yang2016_1-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><style 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no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output 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"Grupo Arroyo del Soldado: paleontologia, edad y correlaciones (Vendiano-Cámbrico Inferior, Uruguay)". <i>Actas II Congreso Uruguaya de Geologia, Montevideo, Sociedad Uruguaya de Geologia — Facultad de Ciencias</i> (in Spanish): <span class="nowrap">183–</span>187.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Actas+II+Congreso+Uruguaya+de+Geologia%2C+Montevideo%2C+Sociedad+Uruguaya+de+Geologia+%E2%80%94+Facultad+de+Ciencias&rft.atitle=Grupo+Arroyo+del+Soldado%3A+paleontologia%2C+edad+y+correlaciones+%28Vendiano-C%C3%A1mbrico+Inferior%2C+Uruguay%29&rft.pages=%3Cspan+class%3D%22nowrap%22%3E183-%3C%2Fspan%3E187&rft.date=1998&rft.au=Gaucher%2C+C.&rft.au=Sprechmann%2C+P.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACloudinidae" class="Z3988"></span></span> </li> <li id="cite_note-Gaucher2000-27"><span class="mw-cite-backlink"><b><a href="#cite_ref-Gaucher2000_27-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFGaucher,_C.2000" class="citation book cs1">Gaucher, C. 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See Grant 1990 for reference and discussion.</span> </li> <li id="cite_note-Yochelson1977-29"><span class="mw-cite-backlink"><b><a href="#cite_ref-Yochelson1977_29-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFYochelson,_E.L.Stump,_E.1977" class="citation journal cs1">Yochelson, E.L.; Stump, E. (1977). 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Retrieved <span class="nowrap">7 July</span> 2024</span> – via Elsevier Science Direct.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+South+American+Earth+Sciences&rft.atitle=Ediacaran+ramp+depositional+model+of+the+Tamengo+Formation%2C+Brazil&rft.volume=96&rft.pages=102348&rft.date=2019-12&rft_id=info%3Adoi%2F10.1016%2Fj.jsames.2019.102348&rft.aulast=Oliveira&rft.aufirst=Rick+Souza+de&rft.au=Nogueira%2C+Afonso+C%C3%A9sar+Rodrigues&rft.au=Romero%2C+Guilherme+Raffaeli&rft.au=Truckenbrodt%2C+Werner&rft.au=da+Silva+Bandeira%2C+Jos%C3%A9+Cavalcante&rft_id=https%3A%2F%2Flinkinghub.elsevier.com%2Fretrieve%2Fpii%2FS0895981119300197&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACloudinidae" class="Z3988"></span></span> </li> <li id="cite_note-Zaine1985-31"><span class="mw-cite-backlink"><b><a href="#cite_ref-Zaine1985_31-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222" /><cite id="CITEREFZaine,_M.F.Fairchild,_T.R.1985" class="citation journal cs1">Zaine, M.F.; Fairchild, T.R. 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(2007). <a rel="nofollow" class="external text" href="http://www.paleo.pan.pl/people/Dzik/Publications/Verdun.pdf">"The Verdun Syndrome: Simultaneous origin of protective armour and infaunal shelters at the Precambrian–Cambrian Transition"</a> <span class="cs1-format">(PDF)</span>. <i>Geological Society, London, Special Publications</i>. <b>286</b> (1): <span class="nowrap">405–</span>414. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2007GSLSP.286..405D">2007GSLSP.286..405D</a>. <a href="/wiki/CiteSeerX_(identifier)" class="mw-redirect" title="CiteSeerX (identifier)">CiteSeerX</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://citeseerx.ist.psu.edu/viewdoc/summary?doi=10.1.1.693.9187">10.1.1.693.9187</a></span>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1144%2FSP286.30">10.1144/SP286.30</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:33112819">33112819</a>. <a rel="nofollow" class="external text" href="https://ghostarchive.org/archive/20221010/http://www.paleo.pan.pl/people/Dzik/Publications/Verdun.pdf">Archived</a> <span class="cs1-format">(PDF)</span> from the original on 10 October 2022<span class="reference-accessdate">. Retrieved <span class="nowrap">30 July</span> 2008</span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Geological+Society%2C+London%2C+Special+Publications&rft.atitle=The+Verdun+Syndrome%3A+Simultaneous+origin+of+protective+armour+and+infaunal+shelters+at+the+Precambrian%E2%80%93Cambrian+Transition&rft.volume=286&rft.issue=1&rft.pages=%3Cspan+class%3D%22nowrap%22%3E405-%3C%2Fspan%3E414&rft.date=2007&rft_id=https%3A%2F%2Fciteseerx.ist.psu.edu%2Fviewdoc%2Fsummary%3Fdoi%3D10.1.1.693.9187%23id-name%3DCiteSeerX&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A33112819%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1144%2FSP286.30&rft_id=info%3Abibcode%2F2007GSLSP.286..405D&rft.au=Dzik%2C+J.&rft_id=http%3A%2F%2Fwww.paleo.pan.pl%2Fpeople%2FDzik%2FPublications%2FVerdun.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ACloudinidae" class="Z3988"></span></span> </li> </ol></div> <div class="navbox-styles"><style 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