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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">EG201932</article-id>
<article-id pub-id-type="doi">10.3989/egeol.43586.572</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Articles</subject>
</subj-group>
</article-categories>
<title-group>
<article-title>A critical reappraisal of the holotype of <italic>Beothukis mistakensis</italic>, a unique exceptionally preserved rangeomorph organism from Mistaken Point, Newfoundland, Canada</article-title>
<trans-title-group xml:lang="es">
<trans-title>Reevaluaci&#x00F3;n cr&#x00ED;tica del holotipo de Beothukis mistakensis, &#x00FA;nico organismo rangeomorfo excepcionalmente preservado en Mistaken Point, Terranova, Canad&#x00E1;</trans-title>
</trans-title-group>
<alt-title alt-title-type="running-head">A critical reappraisal of the holotype of Beothukis mistakensis</alt-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Taylor</surname>
<given-names>R.S.</given-names>
</name>
<xref ref-type="aff" rid="aff0001">1</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Hawco</surname>
<given-names>J.B.</given-names>
</name>
<xref ref-type="aff" rid="aff0001">1</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Nichols</surname>
<given-names>R.</given-names>
</name>
<xref ref-type="aff" rid="aff0002">2</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>McIlroy</surname>
<given-names>D.</given-names>
</name>
<xref ref-type="aff" rid="aff0001">1</xref>
</contrib>
</contrib-group>
<aff id="aff0001"><label>1</label>Department of Earth Sciences, Memorial University of Newfoundland, St. John&#x2019;s, NL, Canada. Email: <email xlink:href="rodt@mun.ca">rodt@mun.ca</email>; ORCID ID: <ext-link ext-link-type="uri" xlink:href="http://orcid.org/0000-0003-3072-6040">http://orcid.org/0000-0003-3072-6040</ext-link>, <ext-link ext-link-type="uri" xlink:href="http://orcid.org/0000-0001-9162-2013">http://orcid.org/0000-0001-9162-2013</ext-link>, <ext-link ext-link-type="uri" xlink:href="http://orcid.org/0000-0001-9521-499X">http://orcid.org/0000-0001-9521-499X</ext-link></aff>
<aff id="aff0002"><label>2</label>Palaeocreations, Bristol, UK. ORCID ID: <ext-link ext-link-type="uri" xlink:href="http://orcid.org/0000-0001-8980-7297">http://orcid.org/0000-0001-8980-7297</ext-link></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.43586.572</elocation-id>
<history>
<date date-type="received">
<day>01</day>
<month>05</month>
<year>2019</year>
</date>
<date date-type="accepted">
<day>03</day>
<month>07</month>
<year>2019</year>
</date>
<date date-type="Publicado on-line">
<day>29</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>The holotype of <italic>Beothukis mistakensis</italic> in Mistaken Point, Newfoundland, is the only currently known specimen that can be definitely placed within the species. Morphological differences between the holotype of <italic>B. mistakensis</italic> and the well-constrained species <italic>plumosa</italic> suggest that the two are not congeneric and <italic>Culmofrons plumosa</italic> should be retained as a valid taxon as originally established.</p>
</abstract>
<trans-abstract xml:lang="es">
<title>RESUMEN</title>
<p>El holotipo de <italic>Beothukis mistakensis</italic> en Mistaken Point, Terranova, es el &#x00FA;nico ejemplar actualmente conocido que puede ser colocado definitivamente dentro de la especie. Las diferencias morfol&#x00F3;gicas entre el holotipo de B. <italic>mistakensis</italic> y la especie plumosa sugieren que ambas no son cong&#x00E9;neres y que <italic>Culmofrons plumosa</italic> deber&#x00ED;a conservarse como un tax&#x00F3;n v&#x00E1;lido tal como se estableci&#x00F3; originalmente.</p>
</trans-abstract>
<kwd-group xml:lang="en">
<title>Keywords</title>
<kwd><italic>Beothukis</italic></kwd>
<kwd>Rangeomorph</kwd>
<kwd>Mistaken Point</kwd>
<kwd>Ediacaran</kwd>
<kwd>Newfoundland</kwd>
</kwd-group>
<kwd-group xml:lang="en">
<title>Palabras clave</title>
<kwd><italic>Beothukis</italic></kwd>
<kwd>Rangeomorfo</kwd>
<kwd>Mistaken Point</kwd>
<kwd>Ediac&#x00E1;rico</kwd>
<kwd>Terranova</kwd>
</kwd-group>
</article-meta>
</front>
<body>
<sec id="sec1" sec-type="intro">
<title>Introduction</title>
<p>The Mistaken Point Ecological Reserve is a UNESCO World Heritage Site situated on the southeast coast of Newfoundland, Canada. It is home to multiple localities preserving complex accumulations of a group of late Ediacaran fossils known as the Avalonian Assemblage (Waggoner, <xref ref-type="bibr" rid="cit0008">2003</xref>). These fossiliferous sites represent whole communities of deepsea seafloor-dwelling organisms that were preserved as &#x2018;snapshots in time&#x2019; through burial by volcanic ash (Clapham <italic>et al</italic>., <xref ref-type="bibr" rid="cit0002">2003</xref>; Ichaso <italic>et al</italic>., <xref ref-type="bibr" rid="cit0003">2007</xref>) ranging in age from 570.94 &#x00B1; 0.38 to 566.25 &#x00B1; 0.35 Ma (Pu <italic>et al</italic>., <xref ref-type="bibr" rid="cit0007">2016</xref>). <italic>Beothukis mistakensis</italic> (<xref ref-type="fig" rid="f0001">Fig. 1A</xref>) is one of the most recognizable Ediacaran fossils from Mistaken Point. It has been illustrated in numerous publications, although it represents an exceptionally small component of the total fossil assemblage. It is perhaps best known for its occurrence on the Mistaken Point &#x2018;E surface&#x2019; in a small area colloquially known as &#x2018;Seilacher&#x2019;s Corner&#x2019;, in close proximity to other well-known Ediacaran taxa (e.g. <italic>Fractofusus</italic>, <italic>Charniodiscus</italic>) in a highly photogenic portion of the surface.</p>
<fig id="f0001">
<label>Figure 1</label>
<caption>
<p>A. Plastotype of the <italic>Beothukis mistakensis</italic> holotype, Mistaken Point Ecological Reserve. B. Interpretive drawing showing the first order units, labelled L1-9 and R1-8 (second order rangeomorph units are coloured so as to be progressively darker closer to the margin of the fossil); the orange coloured features are arcuate units with non-rangeomorph sub-units. (c) Secondary growth occurs at the tips of some first order units; the first arcuate unit on the left overgrows the secondary growths, implying its later development relative to secondary growth of unit L2. Scale bars = 1 cm.</p>
</caption>
<graphic xlink:href="EG201932-117-g001.tif" xmlns:xlink="http://www.w3.org/1999/xlink"/>
</fig>
<p><italic>Beothukis mistakensis</italic> was a uniterminal, frond-shaped organism with an oval to spatulate outline. Like all rangeomorphs, <italic>B. mistakensis</italic> is composed of a complex modular architecture, with each component unit being made up of similar but smaller units arranged in a fractal-like manner (Narbonne, <xref ref-type="bibr" rid="cit0006">2004</xref>; Brasier <italic>et al</italic>., <xref ref-type="bibr" rid="cit0001">2012</xref>). The taxonomic status of the genus <italic>Beothukis</italic> is currently uncertain: it has been amended to include specimens with a stem and basal disc (Brasier <italic>et al</italic>. <xref ref-type="bibr" rid="cit0001">2012</xref>), although these features are not preserved on the holotype. Liu <italic>et al</italic>. (<xref ref-type="bibr" rid="cit0005">2016</xref>) reconsidered <italic>Beothukis mistakensis</italic> in the context of new discoveries of <italic>Culmofrons plumosa</italic> (LaFlamme <italic>et al</italic>., <xref ref-type="bibr" rid="cit0004">2012</xref>; <xref ref-type="fig" rid="f0002">Fig. 2</xref>). <italic>C. plumosa</italic> was considered to possess a frond with similar gross morphology to <italic>Beothukis</italic>, but also a stem and holdfast. <italic>Culmofrons plumosa</italic> was thus considered to be a junior synonym of <italic>B. mistakensis</italic>, thereby creating the new taxon <italic>Beothukis</italic> (<italic>Culmofrons</italic>) <italic>plumosa</italic>. Part of the rationale for this decision was on the basis that continuous characters (i.e., stem length) are unsuitable for genus-level differentiation of taxa (Liu <italic>et al</italic>., <xref ref-type="bibr" rid="cit0005">2016</xref>), the authors instead recommended utilizing gross architecture, the presence/absence of morphological features, and overall growth programme. The number of second order rangeomorph branches (8-12 in <italic>B</italic>. (<italic>C</italic>.) <italic>plumosa versus</italic> 5-8 in <italic>B. mistakensis</italic>) is an important feature used by Liu <italic>et al</italic>. (<xref ref-type="bibr" rid="cit0005">2016</xref>) to distinguish between the two species. The objectives of this study are to re-assess the morphology of the holotype of <italic>B. mistakensis</italic>, and to re-examine the relationship between <italic>B. mistakensis</italic> and <italic>B</italic>. (<italic>C</italic>.) <italic>plumosa</italic>.</p>
<fig id="f0002">
<label>Figure 2</label>
<caption>
<p>Holotype of <italic>Beothukis</italic> (<italic>Culmofrons</italic>) <italic>plumosa</italic> from Lower Mistaken Point, Mistaken Point Ecological Reserve. Scale bar: 2 cm.</p>
</caption>
<graphic xlink:href="EG201932-117-g002.tif" xmlns:xlink="http://www.w3.org/1999/xlink"/>
</fig>
</sec>
<sec id="sec2" sec-type="results|discussion">
<title>Results and discussion</title>
<p>Careful study of the <italic>B. mistakensis</italic> holotype reveals a considerably more complex morphology than previously recognized. The frond is asymmetrical, with the left side being made up of nine primary order units (&#x2018;branches&#x2019;) while the right side consists of only eight such units (<xref ref-type="fig" rid="f0001">Fig. 1B</xref>). Primary order units were added sequentially from the distal tip of the frond towards its base, our interpretation of the primary order units differs from earlier authors but is based on retention of equivalent number of orders of &#x201C;branching&#x201D; within all primary order units. Although the overall outline of the organism is approximately symmetrical across the midline, the units making up the left and right sides show dissimilar sizes and orientations. The number of secondary order rangeomorph units within each primary order unit varies across the medial axis, compensating for the variability in size and shape of the primary units to maintain the overall frond shape. The angle of the primary order units relative to the midline increases from being acute at the distal tip of the frond to being approximately perpendicular near the base of the frond. Tertiary order rangeomorph branches are rotated and furled near the central axis but displayed and furled when bound proximally and distally by secondary growth. The tertiary units located at the lateral margin of the frond show variable morphologies.</p>
<p>Secondary growth occurs around the fringes of several of the first order units on the left side of the organism (<xref ref-type="fig" rid="f0001">Fig. 1C</xref>). Arcuate units without typical rangeomorph branching are also visible on the left-hand side of the holotype, in places overgrowing the secondary fringe growth. Several first order units without rangeomorph branching are located near the basal region of the organism, each composed of multiple rectangular second-order units and obliquely adjoining the frond&#x2019;s central axis.</p>
<p>The paratype of <italic>B. mistakensis</italic> was also examined as a part of this study, revealing that there are 3 vanes of growth within the fossil. We thus determine that it is not congeneric with the holotype and thus needs to be reassigned to a different taxon. Of the many fossils previously described as <italic>Beothukis</italic>, none are morphologically comparable to the holotype, beyond perhaps having a spatulate outline. The other material is thus in need of reassessment, which is the focus of a separate morphometric study.</p>
<p>Several important morphological differences between <italic>B</italic>. <italic>mistakensis</italic> (<xref ref-type="fig" rid="f0001">Fig. 1A</xref>) and <italic>B</italic>. (<italic>C</italic>.) <italic>plumosa</italic> (<xref ref-type="fig" rid="f0002">Fig. 2</xref>) have been identified as a result of this study:</p>
<list list-type="bullet">
<list-item><p>The primary order branches of <italic>B. mistakensis</italic> are unequal and asymmetrical across a straight central axis, while the type specimen of <italic>B</italic>. (<italic>C</italic>.) <italic>plumosa</italic> possesses up to five pairs of equal but alternating primary order branches across its two rows, producing a zig-zag axis.</p></list-item>
<list-item><p>The primary branches of <italic>B. mistakensis</italic> are small near the base and do not attach to a stem, while the primary branches are largest close to the stem in <italic>B</italic>. (<italic>C</italic>.) <italic>plumosa</italic> and attach directly to it.</p></list-item>
<list-item><p>The largest second-order units in <italic>B. mistakensis</italic> are located close to the frond&#x2019;s edge and in the middle of the organism, while in <italic>B</italic>. (<italic>C</italic>.) <italic>plumosa</italic> the largest second-order units are situated near the axis and close to the base of the frond.</p></list-item>
<list-item><p>The morphology of the third-order branches in B. mistakensis vary depending on their position within the frond, while in <italic>B</italic>. (<italic>C</italic>.) <italic>plumosa</italic> most of the tertiary branches appear to be rotated and furled.</p></list-item>
<list-item><p>Lateral branches are found at the base of <italic>B. mistakensis</italic> but are not observed in <italic>B</italic>. (<italic>C</italic>.) <italic>plumosa</italic>.</p></list-item>
</list>
</sec>
<sec id="sec3" sec-type="conclusions">
<title>Conclusions</title>
<p>The holotype of <italic>Beothukis mistakensis</italic> is the only currently known specimen that can be definitely placed within the species. Morphological differences between the holotype of <italic>B. mistakensis</italic> and the well-constrained species plumosa suggest that the two are not congeneric and <italic>C. plumosa</italic> should be retained as a valid taxon as originally established. We hope that further study of the Avalonian Assemblage in Newfoundland will reveal more material belonging to <italic>B. mistakensis</italic>, and that morphometric analysis of the spatulate fronds will provide rationale for discrimination of the diversity within that morphospace.</p>
</sec>
</body>
<back>
<ref-list>
<title>References</title>
<ref id="cit0001">
<nlm-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Brasier</surname>
<given-names>M.D.</given-names>
</name>
<name>
<surname>Antcliffe</surname>
<given-names>J.B.</given-names>
</name>
<name>
<surname>Liu</surname>
<given-names>A.G.</given-names>
</name>
</person-group>
<article-title>The architecture of Ediacaran fronds</article-title>
<source>Palaeontology</source>
<year>2012</year>
<volume>55</volume>
<fpage>1105</fpage>
<lpage>1124</lpage>
<comment>
<ext-link ext-link-type="uri" xlink:href="https://doi.org/10.1111/j.1475-4983.2012.01164.x">https://doi.org/10.1111/j.1475-4983.2012.01164.x</ext-link>
</comment>
</nlm-citation>
</ref>
<ref id="cit0002">
<nlm-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Clapham</surname>
<given-names>M.E.</given-names>
</name>
<name>
<surname>Narbonne</surname>
<given-names>G.M.</given-names>
</name>
<name>
<surname>Gehling</surname>
<given-names>J.G.</given-names>
</name>
</person-group>
<article-title>Paleoecology of the oldest known animal communities: Ediacaran assemblages at Mistaken Point, Newfoundland</article-title>
<source>Paleobiology</source>
<year>2003</year>
<volume>29</volume>
<fpage>527</fpage>
<lpage>544</lpage>
<comment>
<ext-link ext-link-type="uri" xlink:href="https://doi.org/10.1666/0094-8373(2003)029&#x003C;0527:POTOKA&#x003E;2.0.CO;2">https://doi.org/10.1666/0094-8373(2003)029&#x003C;0527:POTOKA&#x003E;2.0.CO;2</ext-link>
</comment>
</nlm-citation>
</ref>
<ref id="cit0003">
<nlm-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Ichaso</surname>
<given-names>A.A.</given-names>
</name>
<name>
<surname>Dalrymple</surname>
<given-names>R.W.</given-names>
</name>
<name>
<surname>Narbonne</surname>
<given-names>G.M.</given-names>
</name>
</person-group>
<article-title>Paleoenvironmental and basin analysis of the Late Neoproterozoic (Ediacaran) Upper Conception and St. John&#x2019;s Groups, West Conception Bay, Newfoundland</article-title>
<source>Canadian Journal of Earth Sciences</source>
<year>2007</year>
<volume>44</volume>
<fpage>25</fpage>
<lpage>41</lpage>
<comment>
<ext-link ext-link-type="uri" xlink:href="https://doi.org/10.1139/e06-098">https://doi.org/10.1139/e06-098</ext-link>
</comment>
</nlm-citation>
</ref>
<ref id="cit0004">
<nlm-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>LaFlamme</surname>
<given-names>M.</given-names>
</name>
<name>
<surname>Flude</surname>
<given-names>L.I.</given-names>
</name>
<name>
<surname>Narbonne</surname>
<given-names>G.Y.</given-names>
</name>
</person-group>
<article-title>Ecological tiering and the evolution of a stem: the oldest stemmed frond from the Ediacaran of Newfoundland, Canada</article-title>
<source>Journal of Palaeontology</source>
<year>2012</year>
<volume>86</volume>
<fpage>193</fpage>
<lpage>200</lpage>
<comment>
<ext-link ext-link-type="uri" xlink:href="https://doi.org/10.1666/11-044.1">https://doi.org/10.1666/11-044.1</ext-link>
</comment>
</nlm-citation>
</ref>
<ref id="cit0005">
<nlm-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Liu</surname>
<given-names>A.G.</given-names>
</name>
<name>
<surname>Matthews</surname>
<given-names>J.J.</given-names>
</name>
<name>
<surname>McIlroy</surname>
<given-names>D.</given-names>
</name>
</person-group>
<article-title>The Beothukis/Culmofrons problem and its bearing on Ediacaran macrofossil taxonomy: evidence from an exceptional new fossil locality</article-title>
<source>Palaeontology</source>
<year>2016</year>
<volume>59</volume>
<fpage>45</fpage>
<lpage>58</lpage>
<comment>
<ext-link ext-link-type="uri" xlink:href="https://doi.org/10.1111/pala.12206">https://doi.org/10.1111/pala.12206</ext-link>
</comment>
</nlm-citation>
</ref>
<ref id="cit0006">
<nlm-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Narbonne</surname>
<given-names>G.M.</given-names>
</name>
</person-group>
<article-title>Modular construction in the Ediacaran Biota</article-title>
<source>Science</source>
<year>2004</year>
<volume>305</volume>
<fpage>1141</fpage>
<lpage>1144</lpage>
<comment>
<ext-link ext-link-type="uri" xlink:href="https://doi.org/10.1126/science.1099727">https://doi.org/10.1126/science.1099727</ext-link>
</comment>
</nlm-citation>
</ref>
<ref id="cit0007">
<nlm-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Pu</surname>
<given-names>J.P.</given-names>
</name>
<name>
<surname>Bowring</surname>
<given-names>S.A.</given-names>
</name>
<name>
<surname>Ramezani</surname>
<given-names>J.</given-names>
</name>
<name>
<surname>Myrow</surname>
<given-names>P.</given-names>
</name>
<name>
<surname>Raub</surname>
<given-names>T.D.</given-names>
</name>
<name>
<surname>Landing</surname>
<given-names>E.</given-names>
</name>
<name>
<surname>Mills</surname>
<given-names>A.</given-names>
</name>
<name>
<surname>Hodgin</surname>
<given-names>E.</given-names>
</name>
<name>
<surname>Macdonald</surname>
<given-names>F. A.</given-names>
</name>
</person-group>
<article-title>Dodging Snowballs: Geochronology of the Gaskiers Glaciation and the first appearance of the Ediacaran Biota</article-title>
<source>Geology</source>
<year>2016</year>
<volume>44</volume>
<fpage>955</fpage>
<lpage>958</lpage>
<comment>
<ext-link ext-link-type="uri" xlink:href="https://doi.org/10.1130/G38284.1">https://doi.org/10.1130/G38284.1</ext-link>
</comment>
</nlm-citation>
</ref>
<ref id="cit0008">
<nlm-citation publication-type="journal">
<person-group person-group-type="author">
<name>
<surname>Waggoner</surname>
<given-names>B.</given-names>
</name>
</person-group>
<article-title>The Ediacaran Biotas in space and time</article-title>
<source>Integrative and Comparative Biology</source>
<year>2003</year>
<volume>843</volume>
<fpage>104</fpage>
<lpage>113</lpage>
<comment>
<ext-link ext-link-type="uri" xlink:href="https://doi.org/10.1093/icb/43.1.104">https://doi.org/10.1093/icb/43.1.104</ext-link>
</comment>
</nlm-citation>
</ref>
</ref-list>
</back>
</article>