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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">EGEOL</journal-id>
<journal-title-group>
<journal-title>Estudios Geol&#x00F3;gicos</journal-title>
<abbrev-journal-title>Estud. Geol.</abbrev-journal-title>
</journal-title-group>
<issn pub-type="epub">0367-0449</issn>
<publisher>
<publisher-name>Consejo Superior de Investigaciones Cientificas</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="publisher-id">EG201919</article-id>
<article-id pub-id-type="doi">10.3989/egeol.43590.559</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Articles</subject>
</subj-group>
</article-categories>
<title-group>
<article-title>Taphonomic bias in <italic>Cloudina</italic> distribution data from Siberia</article-title>
<trans-title-group xml:lang="es">
<trans-title>Sesgo tafon&#x00F3;mico en los datos de distribuci&#x00F3;n de Cloudina en Siberia</trans-title>
</trans-title-group>
<alt-title alt-title-type="running-head">Taphonomic bias in Cloudina distribution data from Siberia</alt-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Markov</surname>
<given-names>G.</given-names>
</name>
<xref ref-type="aff" rid="aff0001">1</xref>
<xref ref-type="aff" rid="aff0002">2</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Rogov</surname>
<given-names>V.</given-names>
</name>
<xref ref-type="aff" rid="aff0001">1</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Karlova</surname>
<given-names>G.</given-names>
</name>
<xref ref-type="aff" rid="aff0001">1</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Grazhdankin</surname>
<given-names>D.</given-names>
</name>
<xref ref-type="aff" rid="aff0001">1</xref>
<xref ref-type="aff" rid="aff0002">2</xref>
</contrib>
</contrib-group>
<aff id="aff0001"><label>1</label>Trofimuk Institute of Petroleum Geology and Geophysics, Novosibirsk, Russia. Email: <email xlink:href="markovge@ipgg.sbras.ru">markovge@ipgg.sbras.ru</email>; ORCID ID: <ext-link ext-link-type="uri" xlink:href="https://orcid.org/0000-0002-7391-5629">https://orcid.org/0000-0002-7391-5629</ext-link>, <ext-link ext-link-type="uri" xlink:href="https://orcid.org/0000-0003-1880-5635">https://orcid.org/0000-0003-1880-5635</ext-link>, <ext-link ext-link-type="uri" xlink:href="https://orcid.org/0000-0001-5914-6870">https://orcid.org/0000-0001-5914-6870</ext-link>, <ext-link ext-link-type="uri" xlink:href="https://orcid.org/0000-0003-0797-1347">https://orcid.org/0000-0003-0797-1347</ext-link></aff>
<aff id="aff0002"><label>2</label>Novosibirsk State University, Novosibirsk, Russia</aff>
<pub-date pub-type="epub">
<day>31</day>
<month>12</month>
<year>2019</year>
</pub-date>
<pub-date pub-type="collection">
<year>2019</year>
</pub-date>
<volume>75</volume>
<issue>2</issue>
<elocation-id>10.3989/egeol.43590.559</elocation-id>
<history>
<date date-type="received">
<day>02</day>
<month>05</month>
<year>2019</year>
</date>
<date date-type="accepted">
<day>01</day>
<month>07</month>
<year>2019</year>
</date>
<date date-type="Publicado on-line">
<day>21</day>
<month>11</month>
<year>2019</year>
</date>
</history>
<permissions>
<copyright-statement>&#x00A9; 2019 CSIC</copyright-statement>
<copyright-year>2019</copyright-year>
<license license-type="open-access" xlink:href="https://creativecommons.org/licenses/by-nc/4.0/">
<license-p>This is an open-access article distributed under the terms of the Creative Commons Attribution-Non Commercial (by-nc) Spain 4.0 License.</license-p>
</license>
</permissions>
<abstract>
<title>ABSTRACT</title>
<p><italic>Cloudina</italic>-morph fossils in Siberia have been traditionally regarded as a taphonomic mode of <italic>Anabarites</italic> tests inserted one into another under specific hydrodynamic conditions. Clusters of telescoped conical tests are ubiquitous in the Kessyusa Group and coeval strata across Siberia and not all of them can be easily interpreted as a result of simple mechanical stacking. It remains to be confirmed whether any of these clusters actually represents a life association of a <italic>Cloudina</italic>-morph structure.</p>
</abstract>
<trans-abstract xml:lang="es">
<title>RESUMEN</title>
<p>En Siberia los morfotipos de <italic>Cloudina</italic> han sido tradicionalmente considerados como una variedad tafon&#x00F3;mica de conchas de <italic>Anabarites</italic>, insertadas una dentro de otra, bajo condiciones hidrodin&#x00E1;micas espec&#x00ED;ficas. Las asociaciones de conchas c&#x00F3;nicas telesc&#x00F3;picas son omnipresentes en el Grupode Kessyusa y en estratos contempor&#x00E1;neos a lo largo de Siberia, y no todos ellos pueden ser f&#x00E1;cilmente interpretados como resultado de un simple apilamiento mec&#x00E1;nico. Queda por ver si alguna de estas bioacumulaciones representa una asociaci&#x00F3;n de vida de una estructura morfot&#x00ED;pica de tipo <italic>Cloudina</italic>.</p>
</trans-abstract>
<kwd-group xml:lang="en">
<title>Keywords</title>
<kwd><italic>Cloudina</italic></kwd>
<kwd><italic>Anabarites</italic></kwd>
<kwd>Taphonomy</kwd>
<kwd>Ediacaran</kwd>
<kwd>Siberia</kwd>
</kwd-group>
<kwd-group xml:lang="en">
<title>Palabras clave</title>
<kwd><italic>Cloudina</italic></kwd>
<kwd><italic>Anabarites</italic></kwd>
<kwd>Tafonom&#x00ED;a</kwd>
<kwd>Ediac&#x00E1;rico</kwd>
<kwd>Siberia</kwd>
</kwd-group>
</article-meta>
</front>
<body>
<sec id="sec1" sec-type="intro">
<title>Introduction</title>
<p>Representatives of the fossil genus <italic>Cloudina</italic>, a potential terminal Ediacaran index-taxon, are preserved in the form of conico-tubular calcareous structure built of numerous nested funnel-shaped elements, with concentrically placed circular to ellipsoid layers in cross section. It is one of the most geographically widespread and numerically abundant fossil in uppermost Ediacaran strata; however, the paucity of this taxon in Siberia has been puzzling (Zhuravlev <italic>et al.</italic>, <xref ref-type="bibr" rid="cit0004">2012</xref>; Grazhdankin <italic>et al.</italic>, <xref ref-type="bibr" rid="cit0001">2015</xref>). Fossiliferous strata in the north-western slope of the Olenek Uplift, arctic Siberia offer an exceptional view of terminal Ediacaran and Terreneuvian palaeobiology. In particular, the Ediacaran strata contain the youngest assemblage of Ediacaran Avalon-type macrofossils preserved as mouldic imprints in limestones; diverse carbonaceous compression macrofossils; probably the planet&#x2019;s oldest known occurrence of diverse taxonomically identifiable skeletal fossils; taxonomically, numerically and ecologically abundant trace fossils; and key radiometric and chemostratigraphic records &#x2013; all in a single 400-m-thick continuous succession (<xref ref-type="fig" rid="f0001">Fig. 1</xref>). Fragments of conico-tubular calcareous moulds found throughout the Turkut Formation and predating the lowest occurrence of fossil anabaritids in the section have been traditionally regarded as poorly preserved fossils of <italic>Cambrotubulus decurvatus</italic> (Nagovitsin <italic>et al.</italic>, <xref ref-type="bibr" rid="cit0002">2015</xref>).</p>
<fig id="f0001">
<label>Figure 1</label>
<caption>
<p>Stratigraphic range of the earliest &#x2018;<italic>Cambrotubulus</italic>&#x2019; fossils in the northwestern slope of the Olenek Uplift, Siberia postdates a diverse assemblage of Ediacaran macrofossils including meniscate trace fossils <italic>Nenoxites curvus</italic> that is often confused with compressed tubular fossils of <italic>Shaanxilithes</italic>, and predates the first occurrence of small skeletal fossils <italic>Anabarites trisulcatus</italic> found in clasts of the stratiform breccia of the Tas-Yuryakh Volcanic Complex (Rogov <italic>et al.</italic>, <xref ref-type="bibr" rid="cit0003">2015</xref>).</p>
</caption>
<graphic xlink:href="EG201919-104-g001.tif" xmlns:xlink="http://www.w3.org/1999/xlink"/>
</fig>
</sec>
<sec id="sec2" sec-type="results|discussion">
<title>Results and discussion</title>
<p>The first occurrence of <italic>Cloudina</italic>-morph small skeletal fossils in the Olenek Uplift is found in the uppermost Syhargalakh Formation of the Kessyusa Group (<xref ref-type="fig" rid="f0001">Fig. 1</xref>). Here the <italic>Cloudina</italic>-morph fossils are all allochthonous (<xref ref-type="fig" rid="f0002">Fig. 2A-D</xref>) forming a laterally continuous coquina deposit that rests upon the stratiform breccia of the Tas-Yuryakh Volcanic Complex and appears to be coeval with the lowest stratigraphic occurrence in the section of the Cambrian index fossil <italic>Treptichnus pedum</italic>. Furthermore, the coquina deposit is immediately overlain by alternating sandstone and siltstone of the lowermost Mattaia Formation that yielded arthropod trace fossils <italic>Rusophycus avalonensis</italic> (<xref ref-type="fig" rid="f0001">Fig. 1</xref>). <italic>Cloudina</italic>-morphs in South China, Maly Karatau and Siberia have been known to occur together with small skeletal fossils of early Cambrian appearance (Zhuravlev <italic>et al.</italic>, <xref ref-type="bibr" rid="cit0004">2012</xref>; Yang <italic>et al.</italic>, <xref ref-type="bibr" rid="cit0005">2016</xref>). The section of the Olenek Uplift in Siberia provides the first record of the <italic>Cloudina</italic>-morph fossils postdating the Cambrian index-taxon <italic>Treptichnus pedum</italic>. Supposing our interpretation of the earliest &#x2018;<italic>Cambrotubulus</italic>&#x2019; fossils from the Turkut Formation is correct, the stratigraphic range of <italic>Cloudina</italic>-morph fossils in Siberia commences with the strata predating the first occurrence of <italic>Anabarites trisulcatus</italic> and extends into the <italic>Purella antiqua</italic> Assemblage Zone. Some of the <italic>Cloudina</italic>-morph fossils occur in strata that are demonstrably part of the Cambrian Stage 2 (<xref ref-type="fig" rid="f0002">Fig. 2E-G</xref>).</p>
<fig id="f0002">
<label>Figure 2</label>
<caption>
<p><italic>Cloudina</italic>-morph fossils from the lower (A-D), middle (E) and uppermost strata (F-G) of the Kessyusa Group, Terreneuvian, Olenek Uplift, Siberia.</p>
</caption>
<graphic xlink:href="EG201919-104-g002.tif" xmlns:xlink="http://www.w3.org/1999/xlink"/>
</fig>
</sec>
<sec id="sec3" sec-type="conclusions">
<title>Concluding remarks</title>
<p><italic>Cloudina</italic>-morph fossils in Siberia have always been interpreted as a taphonomic mode of <italic>Anabarites</italic> tests inserted one into another under specific hydrodynamic conditions. Clusters of telescoped conical tests are ubiquitous in the Kessyusa Group and coeval strata across Siberia but not all of them can be easily interpreted as a result of simple mechanical stacking (<xref ref-type="fig" rid="f0003">Fig. 3</xref>). We suggest that these fossils represent internal moulds of disarticulated <italic>Cloudina</italic> specimens. Disarticulation, internal mould formation, diagenetic dissolution and recrystallisation of tubes represent an under-explored taphonomic pathway in the preservation of <italic>Cloudina</italic>. If borne out by future studies, the entire early Nemakit-Daldynian fossil record of Siberia must be reevaluated to remove the bias in the <italic>Cloudina</italic> distribution data.</p>
<fig id="f0003">
<label>Figure 3</label>
<caption>
<p>Clusters of telescoped tests of <italic>Anabarites</italic> in lateral (A-D; scale: 1000 &#x00B5;m) and cross-sectional (A&#x2019;-D&#x2019;; scale: 500 &#x00B5;m) view from the lowermost Mattaia Formation. A hemispherical base of one of the tests in lateral and plane view showing a central cavity (E; scale: 1000 &#x00B5;m) from the Syhargalakh Formation.</p>
</caption>
<graphic xlink:href="EG201919-104-g003.tif" xmlns:xlink="http://www.w3.org/1999/xlink"/>
</fig>
</sec>
</body>
<back>
<ack>
<title>ACKNOWLEDGEMENTS</title>
<p>This study was supported by the Russian Foundation for Basic Research Grant 18-05-70110. The findings and conclusions have immediate implications for understanding early evolution of Metazoa (Russian Science Foundation Grant 17-17-01241). Fieldwork was conducted with the financial support of the National Geographic Society&#x2019;s Committee for Research and Exploration (grants 8227-07, 8637-09, 9031-11, NGS-372R-18).</p>
</ack>
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