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<document ID-DOI="10.1038/s41467-022-34204-w" ID-GBIF-Dataset="78018b36-e0be-4bd2-9efb-ead93f0bc73b" ID-PMC="PMC9666559" ID-PubMed="36379946" ID-Zenodo-Dep="7346844" approvedBy="tatia" checkinTime="1669112309551" checkinUser="tatiana" docAuthor="Pates, Stephen, Botting, Joseph P., Muir, Lucy A. &amp; Wolfe, Joanna M." docDate="2022" docId="03FD8143EC04FFAAFF75FA0BFF0F9E2A" docLanguage="en" docName="NatCommun.13.6969.pdf" docOrigin="Nature Communications 13 (1)" docSource="http://dx.doi.org/10.1038/s41467-022-34204-w" docTitle="Panarthropoda Nielsen 1995" docType="treatment" docVersion="3" lastPageNumber="5" masterDocId="FFC4F93BEC06FFAEFFE2FFF0FFE99904" masterDocTitle="Ordovician opabiniid-like animals and the role of the proboscis in euarthropod head evolution" masterLastPageNumber="15" masterPageNumber="1" pageNumber="3" updateTime="1669115630990" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>Ordovician opabiniid-like animals and the role of the proboscis in euarthropod head evolution</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Pates, Stephen</mods:namePart>
<mods:affiliation>Department of Zoology, University of Cambridge, Cambridge, UK.</mods:affiliation>
<mods:nameIdentifier type="email">sp587@cam.ac.uk</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Botting, Joseph P.</mods:namePart>
<mods:affiliation>Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing, China &amp; Department of Natural Sciences, Amgueddfa Cymru — National Museum Wales, Cardiff, UK.</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Muir, Lucy A.</mods:namePart>
<mods:affiliation>Department of Natural Sciences, Amgueddfa Cymru — National Museum Wales, Cardiff, UK.</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Wolfe, Joanna M.</mods:namePart>
<mods:affiliation>Museum of Comparative Zoology and Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, MA, USA</mods:affiliation>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>Nature Communications</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2022</mods:date>
<mods:detail type="pubDate">
<mods:number>2022-11-15</mods:number>
</mods:detail>
<mods:detail type="volume">
<mods:number>13</mods:number>
</mods:detail>
<mods:detail type="issue">
<mods:number>1</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>1</mods:start>
<mods:end>15</mods:end>
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<mods:location>
<mods:url>http://dx.doi.org/10.1038/s41467-022-34204-w</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">10.1038/s41467-022-34204-w</mods:identifier>
<mods:identifier type="GBIF-Dataset">78018b36-e0be-4bd2-9efb-ead93f0bc73b</mods:identifier>
<mods:identifier type="PMC">PMC9666559</mods:identifier>
<mods:identifier type="PubMed">36379946</mods:identifier>
<mods:identifier type="Zenodo-Dep">7346844</mods:identifier>
</mods:mods>
<treatment ID-DOI="http://doi.org/10.5281/zenodo.7346858" ID-Zenodo-Dep="7346858" LSID="urn:lsid:plazi:treatment:03FD8143EC04FFAAFF75FA0BFF0F9E2A" httpUri="http://treatment.plazi.org/id/03FD8143EC04FFAAFF75FA0BFF0F9E2A" lastPageId="4" lastPageNumber="5" pageId="2" pageNumber="3">
<subSubSection box="[151,416,1531,1552]" pageId="2" pageNumber="3" type="nomenclature">
<paragraph blockId="2.[106,786,1360,1581]" box="[151,416,1531,1552]" pageId="2" pageNumber="3">
Castle Bank
<taxonomicName authorityName="Nielsen" authorityYear="1995" box="[271,416,1531,1552]" kingdom="Animalia" pageId="2" pageNumber="3" rank="superPhylum" superPhylum="Panarthropoda">euarthropod A</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection box="[151,485,1560,1581]" pageId="2" pageNumber="3" type="description">
<paragraph blockId="2.[106,786,1360,1581]" box="[151,485,1560,1581]" pageId="2" pageNumber="3">
<figureCitation box="[151,267,1560,1581]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStart-3="Fig" captionStartId-0="7.[106,135,1414,1431]" captionStartId-1="8.[106,135,1620,1637]" captionStartId-2="9.[106,135,1309,1326]" captionStartId-3="10.[106,135,897,914]" captionTargetBox-0="[131,1484,137,1386]" captionTargetBox-1="[141,1460,133,1596]" captionTargetBox-2="[142,1460,130,1285]" captionTargetBox-3="[141,1459,130,870]" captionTargetId-0="graphics-300@7.[142,1460,740,1357]" captionTargetId-1="figure-175@8.[518,896,498,865]" captionTargetId-2="figure-399@9.[518,896,485,842]" captionTargetId-3="figure-1957@10.[517,895,500,871]" captionTargetPageId-0="7" captionTargetPageId-1="8" captionTargetPageId-2="9" captionTargetPageId-3="10" captionText-0="Fig. 6 | Castle Bank euarthropod A (NMW.2021.3 G.8) from the Castle Bank biota. a Overview of whole specimen. Boxes indicate areas detailed in Fig. 7. b Explanatory drawing of (a). an annulations of proboscis, cf-l left blades of caudal fan, cf-r right blades of caudal fan, df dorsal furrow in trunk,fl dorsolateral flap,ic internal canal of proboscis, ls lateral sclerite, pr proboscis, re subrectangular elements posterior to lateral sclerites, sp spine on proboscis." captionText-1="Fig. 7 | Details of anterior of Castle Bank euarthropod A (NMW.2021.3G.8). a Anterior of head including lateral sclarites, rectangular elements and spinose proboscis with annulations (b) Details of spinose proboscis and annulations; an annulation, ic internal canal of proboscis, ls-l left lateral sclerite, ls-r right lateral sclerite,mr marginalrim to carapace element,pr proboscis,re rectangular element posterior to lateral sclerites, sp spine." captionText-2="Fig. 8 | Details of Castle Bank euarthropod A (NMW.2021.3 G.8), photographed using fluorescence. a Overview of whole specimen.b Details of tail fan with spinose margin. c Details of proboscis with dorsal spines. sp spine." captionText-3="Fig. 9 | Details of posterior body and tail fan of Castle Bank euarthropod A (NMW.2021.3 G.8). cb-l left caudal blade,cb-r right caudal blade,df dorsal furrow,fl lateral flap, sp spine." figureDoi-0="http://doi.org/10.5281/zenodo.7346862" figureDoi-1="http://doi.org/10.5281/zenodo.7346866" figureDoi-2="http://doi.org/10.5281/zenodo.7346868" figureDoi-3="http://doi.org/10.5281/zenodo.7346870" httpUri-0="https://zenodo.org/record/7346862/files/figure.png" httpUri-1="https://zenodo.org/record/7346866/files/figure.png" httpUri-2="https://zenodo.org/record/7346868/files/figure.png" httpUri-3="https://zenodo.org/record/7346870/files/figure.png" pageId="2" pageNumber="3">
Figures 6
<emphasis box="[241,253,1561,1581]" italics="true" pageId="2" pageNumber="3">–</emphasis>
9
</figureCitation>
, Supplementary
<figureCitation box="[433,485,1560,1581]" captionStart="Fig" captionStartId="6.[106,135,1286,1303]" captionTargetBox="[142,1461,131,1262]" captionTargetId="figure-410@6.[518,896,507,885]" captionTargetPageId="6" captionText="Fig. 5 | Details of trunk of Mieridduryn bonniae nov. gen. et sp. (NMW.2021.3 G.7). a Overview of trunk. b Details of left dorsolateral flap and associated setal blades. c, d details of lobopodous limbs with annulations and spines. an annulations on lobopodous limbs,lf left dorsolateral flap, ll left lobopodous limb, sb setal blade, sp spines on posterior margin of lobopodous limb." figureDoi="http://doi.org/10.5281/zenodo.7346860" httpUri="https://zenodo.org/record/7346860/files/figure.png" pageId="2" pageNumber="3">Fig. 5</figureCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="2" pageNumber="3" type="materials_examined">
<paragraph blockId="2.[106,788,1617,1724]" pageId="2" pageNumber="3">
<emphasis bold="true" box="[106,423,1618,1638]" pageId="2" pageNumber="3">Material, locality, and horizon</emphasis>
.
<materialsCitation collectionCode="NMW" country="United Kingdom" location="Powys" pageId="2" pageNumber="3" specimenCode="NMW.2021.3 G.8" specimenCount="1" stateProvince="Powys">
<specimenCode box="[438,601,1617,1638]" pageId="2" pageNumber="3">NMW.2021.3 G.8</specimenCode>
, known from part and counterpart. Collected from the Darriwilian (Middle Ordovician,
<taxonomicName authority="Biozone" authorityName="Biozone" box="[106,462,1674,1695]" class="Pterobranchia" family="Dichograptidae" genus="Didymograptus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="2" pageNumber="3" phylum="Hemichordata" rank="species" species="murchisoni">
<emphasis bold="true" box="[106,370,1675,1694]" italics="true" pageId="2" pageNumber="3">Didymograptus murchisoni</emphasis>
Biozone
</taxonomicName>
)
<locationDeviation pageId="2" pageNumber="3">Gilwern Volcanic Formation at Castle Bank</locationDeviation>
,
<locationDeviation box="[232,404,1703,1724]" pageId="2" pageNumber="3">near Llandrindod</locationDeviation>
,
<collectingRegion box="[413,475,1703,1724]" pageId="2" pageNumber="3">Powys</collectingRegion>
(
<collectingCountry box="[486,519,1703,1724]" pageId="2" pageNumber="3">UK</collectingCountry>
)
<superScript attach="none" box="[525,556,1700,1712]" fontSize="5" pageId="2" pageNumber="3">
<bibRefCitation author="Botting, J. P." box="[525,539,1700,1712]" journalOrPublisher="Bull. Geosci." pageId="2" pageNumber="3" pagination="265 - 267" part="96" refId="ref9535" refString="33. Botting, J. P. Hexactins in the ' protomonaxonid' sponge Choiaella and proposal of Ascospongiae (class nov.) as a formal replacement for the Protomonaxonida. Bull. Geosci. 96, 265 - 267 (2021)." title="Hexactins in the ' protomonaxonid' sponge Choiaella and proposal of Ascospongiae (class nov.) as a formal replacement for the Protomonaxonida" type="journal article" year="2021">33</bibRefCitation>
,
<bibRefCitation author="Botting, J. P. &amp; Ma, J." box="[542,556,1700,1712]" journalOrPublisher="Palaeoworld" pageId="2" pageNumber="3" pagination="621 - 632" part="31" refId="ref9580" refString="34. Botting, J. P. &amp; Ma, J. A probable hyalonematid sponge (Hexactinellida: Amphidiscophora) from the Middle Ordovician of the Builth Inlier, Wales. Palaeoworld 31, 621 - 632 (2022)." title="A probable hyalonematid sponge (Hexactinellida: Amphidiscophora) from the Middle Ordovician of the Builth Inlier, Wales" type="journal article" year="2022">34</bibRefCitation>
</superScript>
.
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection lastPageId="3" lastPageNumber="4" pageId="2" pageNumber="3" type="description">
<paragraph blockId="2.[106,787,1760,1981]" pageId="2" pageNumber="3">
<emphasis bold="true" box="[106,224,1760,1780]" pageId="2" pageNumber="3">Description</emphasis>
. NMW.2021.3 G.8 is a complete specimen preserved compressed to givea lateral viewand measures ~3 mm alongthe dorsal margin (
<figureCitation box="[187,243,1817,1838]" captionStart="Fig" captionStartId="7.[106,135,1414,1431]" captionTargetBox="[131,1484,137,1386]" captionTargetId="graphics-300@7.[142,1460,740,1357]" captionTargetPageId="7" captionText="Fig. 6 | Castle Bank euarthropod A (NMW.2021.3 G.8) from the Castle Bank biota. a Overview of whole specimen. Boxes indicate areas detailed in Fig. 7. b Explanatory drawing of (a). an annulations of proboscis, cf-l left blades of caudal fan, cf-r right blades of caudal fan, df dorsal furrow in trunk,fl dorsolateral flap,ic internal canal of proboscis, ls lateral sclerite, pr proboscis, re subrectangular elements posterior to lateral sclerites, sp spine on proboscis." figureDoi="http://doi.org/10.5281/zenodo.7346862" httpUri="https://zenodo.org/record/7346862/files/figure.png" pageId="2" pageNumber="3">Fig. 6</figureCitation>
).
</paragraph>
<paragraph blockId="2.[106,787,1760,1981]" lastBlockId="2.[816,1498,1360,1981]" pageId="2" pageNumber="3">
The head region preserves evidence for what appear to be two lateral cephalic sclerites proximal to an annulated proboscis (
<figureCitation box="[714,781,1874,1895]" captionStart="Fig" captionStartId="7.[106,135,1414,1431]" captionTargetBox="[131,1484,137,1386]" captionTargetId="graphics-300@7.[142,1460,740,1357]" captionTargetPageId="7" captionText="Fig. 6 | Castle Bank euarthropod A (NMW.2021.3 G.8) from the Castle Bank biota. a Overview of whole specimen. Boxes indicate areas detailed in Fig. 7. b Explanatory drawing of (a). an annulations of proboscis, cf-l left blades of caudal fan, cf-r right blades of caudal fan, df dorsal furrow in trunk,fl dorsolateral flap,ic internal canal of proboscis, ls lateral sclerite, pr proboscis, re subrectangular elements posterior to lateral sclerites, sp spine on proboscis." figureDoi="http://doi.org/10.5281/zenodo.7346862" httpUri="https://zenodo.org/record/7346862/files/figure.png" pageId="2" pageNumber="3">Figs. 6</figureCitation>
,
<figureCitation box="[106,117,1903,1924]" captionStart="Fig" captionStartId="8.[106,135,1620,1637]" captionTargetBox="[141,1460,133,1596]" captionTargetId="figure-175@8.[518,896,498,865]" captionTargetPageId="8" captionText="Fig. 7 | Details of anterior of Castle Bank euarthropod A (NMW.2021.3G.8). a Anterior of head including lateral sclarites, rectangular elements and spinose proboscis with annulations (b) Details of spinose proboscis and annulations; an annulation, ic internal canal of proboscis, ls-l left lateral sclerite, ls-r right lateral sclerite,mr marginalrim to carapace element,pr proboscis,re rectangular element posterior to lateral sclerites, sp spine." figureDoi="http://doi.org/10.5281/zenodo.7346866" httpUri="https://zenodo.org/record/7346866/files/figure.png" pageId="2" pageNumber="3">7</figureCitation>
). A pair of trapezoidal sclerites &lt;1 mm in length, which are compressed and superimposed on each other, have been rotated forwards to cover the anterior of the head and proximal part of the proboscis (Supplementary
<figureCitation box="[978,1033,1360,1381]" captionStart="Fig" captionStartId="6.[106,135,1286,1303]" captionTargetBox="[142,1461,131,1262]" captionTargetId="figure-410@6.[518,896,507,885]" captionTargetPageId="6" captionText="Fig. 5 | Details of trunk of Mieridduryn bonniae nov. gen. et sp. (NMW.2021.3 G.7). a Overview of trunk. b Details of left dorsolateral flap and associated setal blades. c, d details of lobopodous limbs with annulations and spines. an annulations on lobopodous limbs,lf left dorsolateral flap, ll left lobopodous limb, sb setal blade, sp spines on posterior margin of lobopodous limb." figureDoi="http://doi.org/10.5281/zenodo.7346860" httpUri="https://zenodo.org/record/7346860/files/figure.png" pageId="2" pageNumber="3">Fig. 5</figureCitation>
). The preserved portion of the proboscis is less than 1 mm in length. The cephalic sclerites preserve a marginal rim, and the element covering the right side of the head also displays numerous triangular spines on the anterior and ventral margins (
<figureCitation box="[822,890,1474,1495]" captionStart="Fig" captionStartId="8.[106,135,1620,1637]" captionTargetBox="[141,1460,133,1596]" captionTargetId="figure-175@8.[518,896,498,865]" captionTargetPageId="8" captionText="Fig. 7 | Details of anterior of Castle Bank euarthropod A (NMW.2021.3G.8). a Anterior of head including lateral sclarites, rectangular elements and spinose proboscis with annulations (b) Details of spinose proboscis and annulations; an annulation, ic internal canal of proboscis, ls-l left lateral sclerite, ls-r right lateral sclerite,mr marginalrim to carapace element,pr proboscis,re rectangular element posterior to lateral sclerites, sp spine." figureDoi="http://doi.org/10.5281/zenodo.7346866" httpUri="https://zenodo.org/record/7346866/files/figure.png" pageId="2" pageNumber="3">Fig. 7a</figureCitation>
, Supplementary
<figureCitation box="[1055,1111,1474,1495]" captionStart="Fig" captionStartId="6.[106,135,1286,1303]" captionTargetBox="[142,1461,131,1262]" captionTargetId="figure-410@6.[518,896,507,885]" captionTargetPageId="6" captionText="Fig. 5 | Details of trunk of Mieridduryn bonniae nov. gen. et sp. (NMW.2021.3 G.7). a Overview of trunk. b Details of left dorsolateral flap and associated setal blades. c, d details of lobopodous limbs with annulations and spines. an annulations on lobopodous limbs,lf left dorsolateral flap, ll left lobopodous limb, sb setal blade, sp spines on posterior margin of lobopodous limb." figureDoi="http://doi.org/10.5281/zenodo.7346860" httpUri="https://zenodo.org/record/7346860/files/figure.png" pageId="2" pageNumber="3">Fig. 5</figureCitation>
). The proboscis, which curves ventrally and is likely incomplete distally, displays annulations and a dark linear feature that runs parallel to the long axis, interpreted as an internal canal, as well as regularly spaced (approximately one per three to four annulations) stout triangular dorsal spines (
<figureCitation box="[1241,1304,1589,1610]" captionStart="Fig" captionStartId="8.[106,135,1620,1637]" captionTargetBox="[141,1460,133,1596]" captionTargetId="figure-175@8.[518,896,498,865]" captionTargetPageId="8" captionText="Fig. 7 | Details of anterior of Castle Bank euarthropod A (NMW.2021.3G.8). a Anterior of head including lateral sclarites, rectangular elements and spinose proboscis with annulations (b) Details of spinose proboscis and annulations; an annulation, ic internal canal of proboscis, ls-l left lateral sclerite, ls-r right lateral sclerite,mr marginalrim to carapace element,pr proboscis,re rectangular element posterior to lateral sclerites, sp spine." figureDoi="http://doi.org/10.5281/zenodo.7346866" httpUri="https://zenodo.org/record/7346866/files/figure.png" pageId="2" pageNumber="3">Figs. 7</figureCitation>
,
<figureCitation box="[1316,1342,1589,1610]" captionStart="Fig" captionStartId="9.[106,135,1309,1326]" captionTargetBox="[142,1460,130,1285]" captionTargetId="figure-399@9.[518,896,485,842]" captionTargetPageId="9" captionText="Fig. 8 | Details of Castle Bank euarthropod A (NMW.2021.3 G.8), photographed using fluorescence. a Overview of whole specimen.b Details of tail fan with spinose margin. c Details of proboscis with dorsal spines. sp spine." figureDoi="http://doi.org/10.5281/zenodo.7346868" httpUri="https://zenodo.org/record/7346868/files/figure.png" pageId="2" pageNumber="3">8c</figureCitation>
). The proboscis is more similar in preservation to the trapezoidal sclerites than to the trunk.
</paragraph>
<paragraph blockId="2.[816,1498,1360,1981]" pageId="2" pageNumber="3">
The trunk, which measures ca. 2 mm along the dorsal margin, is curved, narrow and tapers slightly to the posterior. The trunk displays a wrinkled texture. At the anterior (just behind the lateral carapace elements), faint subrectangular elements sit dorsal to the body region (
<figureCitation box="[822,886,1789,1810]" captionStart="Fig" captionStartId="8.[106,135,1620,1637]" captionTargetBox="[141,1460,133,1596]" captionTargetId="figure-175@8.[518,896,498,865]" captionTargetPageId="8" captionText="Fig. 7 | Details of anterior of Castle Bank euarthropod A (NMW.2021.3G.8). a Anterior of head including lateral sclarites, rectangular elements and spinose proboscis with annulations (b) Details of spinose proboscis and annulations; an annulation, ic internal canal of proboscis, ls-l left lateral sclerite, ls-r right lateral sclerite,mr marginalrim to carapace element,pr proboscis,re rectangular element posterior to lateral sclerites, sp spine." figureDoi="http://doi.org/10.5281/zenodo.7346866" httpUri="https://zenodo.org/record/7346866/files/figure.png" pageId="2" pageNumber="3">Fig. 7a</figureCitation>
; Supplementary
<figureCitation box="[1052,1106,1789,1810]" captionStart="Fig" captionStartId="6.[106,135,1286,1303]" captionTargetBox="[142,1461,131,1262]" captionTargetId="figure-410@6.[518,896,507,885]" captionTargetPageId="6" captionText="Fig. 5 | Details of trunk of Mieridduryn bonniae nov. gen. et sp. (NMW.2021.3 G.7). a Overview of trunk. b Details of left dorsolateral flap and associated setal blades. c, d details of lobopodous limbs with annulations and spines. an annulations on lobopodous limbs,lf left dorsolateral flap, ll left lobopodous limb, sb setal blade, sp spines on posterior margin of lobopodous limb." figureDoi="http://doi.org/10.5281/zenodo.7346860" httpUri="https://zenodo.org/record/7346860/files/figure.png" pageId="2" pageNumber="3">Fig. 5</figureCitation>
). Further to the posterior, indents in the dorsal margin and faint transverse lineations indicate the position of intersegmental furrows and associated subrectangular lateral swimming flaps (
<figureCitation box="[936,1005,1874,1895]" captionStart="Fig" captionStartId="7.[106,135,1414,1431]" captionTargetBox="[131,1484,137,1386]" captionTargetId="graphics-300@7.[142,1460,740,1357]" captionTargetPageId="7" captionText="Fig. 6 | Castle Bank euarthropod A (NMW.2021.3 G.8) from the Castle Bank biota. a Overview of whole specimen. Boxes indicate areas detailed in Fig. 7. b Explanatory drawing of (a). an annulations of proboscis, cf-l left blades of caudal fan, cf-r right blades of caudal fan, df dorsal furrow in trunk,fl dorsolateral flap,ic internal canal of proboscis, ls lateral sclerite, pr proboscis, re subrectangular elements posterior to lateral sclerites, sp spine on proboscis." figureDoi="http://doi.org/10.5281/zenodo.7346862" httpUri="https://zenodo.org/record/7346862/files/figure.png" pageId="2" pageNumber="3">Figs. 6</figureCitation>
,
<figureCitation box="[1020,1033,1874,1895]" captionStart="Fig" captionStartId="10.[106,135,897,914]" captionTargetBox="[141,1459,130,870]" captionTargetId="figure-1957@10.[517,895,500,871]" captionTargetPageId="10" captionText="Fig. 9 | Details of posterior body and tail fan of Castle Bank euarthropod A (NMW.2021.3 G.8). cb-l left caudal blade,cb-r right caudal blade,df dorsal furrow,fl lateral flap, sp spine." figureDoi="http://doi.org/10.5281/zenodo.7346870" httpUri="https://zenodo.org/record/7346870/files/figure.png" pageId="2" pageNumber="3">9</figureCitation>
). Fainter lines overlain by wrinkles preserved towards the middle of the trunk are also tentatively identified as dorsal furrows and lateral flaps, and indicate the presence of at least 12 furrows and flaps (
<figureCitation box="[970,1026,1960,1981]" captionStart="Fig" captionStartId="7.[106,135,1414,1431]" captionTargetBox="[131,1484,137,1386]" captionTargetId="graphics-300@7.[142,1460,740,1357]" captionTargetPageId="7" captionText="Fig. 6 | Castle Bank euarthropod A (NMW.2021.3 G.8) from the Castle Bank biota. a Overview of whole specimen. Boxes indicate areas detailed in Fig. 7. b Explanatory drawing of (a). an annulations of proboscis, cf-l left blades of caudal fan, cf-r right blades of caudal fan, df dorsal furrow in trunk,fl dorsolateral flap,ic internal canal of proboscis, ls lateral sclerite, pr proboscis, re subrectangular elements posterior to lateral sclerites, sp spine on proboscis." figureDoi="http://doi.org/10.5281/zenodo.7346862" httpUri="https://zenodo.org/record/7346862/files/figure.png" pageId="2" pageNumber="3">Fig. 6</figureCitation>
).
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7346850" ID-Zenodo-Dep="7346850" httpUri="https://zenodo.org/record/7346850/files/figure.png" pageId="3" pageNumber="4" startId="3.[106,135,1493,1510]" targetBox="[142,1460,130,1463]" targetPageId="3">
<paragraph blockId="3.[106,784,1492,1563]" lastBlockId="3.[820,1498,1492,1564]" pageId="3" pageNumber="4">
<emphasis bold="true" pageId="3" pageNumber="4">
Fig. 2 | Details of anterior of
<taxonomicName authority="Pates &amp; Botting &amp; Muir &amp; Wolfe, 2022" authorityName="Pates &amp; Botting &amp; Muir &amp; Wolfe" authorityYear="2022" box="[357,538,1492,1510]" family="Opabiniidae" genus="Mieridduryn" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Arthropoda" rank="species" species="bonniae" status="nov. gen. et sp.">
<emphasis bold="true" box="[357,538,1492,1510]" italics="true" pageId="3" pageNumber="4">Mieridduryn bonniae</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[543,678,1493,1509]" pageId="3" pageNumber="4" rank="species">nov. gen. et sp.</taxonomicNameLabel>
(NMW.2021.3 G.7). a
</emphasis>
Anteriorof the head region including proboscis,mouth, gut, view of anterior details including spinous proboscis,mouth and gut.
<emphasis bold="true" box="[675,687,1546,1562]" pageId="3" pageNumber="4">b</emphasis>
Mouth and anterior flapswith strengtheningrays.
<emphasis bold="true" box="[1135,1145,1493,1509]" pageId="3" pageNumber="4">c</emphasis>
Explanatory drawingof (
<emphasis bold="true" box="[1350,1361,1493,1509]" pageId="3" pageNumber="4">b</emphasis>
). anannulations on proboscis, gu gut, lfleftdorsolateral flap, mo mouth,rf rightdorsolateral flap, sb setal blades, sp spines on proboscis, sr strengthening rays.
</paragraph>
</caption>
<paragraph blockId="3.[106,788,1615,1751]" pageId="3" pageNumber="4">
The body terminates in a caudalfan ~0.25 mm long, composed of triangular caudal blades with spinose lateral and posterior margins. Three blades that widen to the posterior can be observed on the near (right) side, whereas two are observed on the left side (
<figureCitation box="[111,191,1730,1751]" captionStart="Fig" captionStartId="9.[106,135,1309,1326]" captionTargetBox="[142,1460,130,1285]" captionTargetId="figure-399@9.[518,896,485,842]" captionTargetPageId="9" captionText="Fig. 8 | Details of Castle Bank euarthropod A (NMW.2021.3 G.8), photographed using fluorescence. a Overview of whole specimen.b Details of tail fan with spinose margin. c Details of proboscis with dorsal spines. sp spine." figureDoi="http://doi.org/10.5281/zenodo.7346868" httpUri="https://zenodo.org/record/7346868/files/figure.png" pageId="3" pageNumber="4">Figs. 8b</figureCitation>
,
<figureCitation box="[201,214,1730,1751]" captionStart="Fig" captionStartId="10.[106,135,897,914]" captionTargetBox="[141,1459,130,870]" captionTargetId="figure-1957@10.[517,895,500,871]" captionTargetPageId="10" captionText="Fig. 9 | Details of posterior body and tail fan of Castle Bank euarthropod A (NMW.2021.3 G.8). cb-l left caudal blade,cb-r right caudal blade,df dorsal furrow,fl lateral flap, sp spine." figureDoi="http://doi.org/10.5281/zenodo.7346870" httpUri="https://zenodo.org/record/7346870/files/figure.png" pageId="3" pageNumber="4">9</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection lastPageId="4" lastPageNumber="5" pageId="3" pageNumber="4" type="discussion">
<paragraph blockId="3.[106,787,1787,1979]" lastBlockId="3.[816,1498,1617,1981]" pageId="3" pageNumber="4">
<emphasis bold="true" box="[106,195,1787,1807]" pageId="3" pageNumber="4">Remarks</emphasis>
. The wrinkled texture of the body, change in slope on the dorsal margin and orientation of the paired head sclerites (rotated forwards), as well as the lack of internal structures (e.g. guts, nervous system) suggest that this specimen may represent a moult rather than a carcass
<bibRefCitation author="Daley, A. C. &amp; Drage, H. B." box="[194,209,1898,1910]" journalOrPublisher="Arthropod Struct. Dev." pageId="3" pageNumber="4" pagination="71 - 96" part="45" refId="ref9796" refString="39. Daley, A. C. &amp; Drage, H. B. The fossil record of ecdysis, and trends in the moulting behaviour of trilobites. Arthropod Struct. Dev. 45, 71 - 96 (2016)." title="The fossil record of ecdysis, and trends in the moulting behaviour of trilobites" type="journal article" year="2016">
<superScript attach="left" box="[194,209,1898,1910]" fontSize="5" pageId="3" pageNumber="4">39</superScript>
</bibRefCitation>
, though this is not conclusive. The presence of an internal cavity does not refute this, as similar structures have been reported from isolated radiodont appendages (e.g. ref. 15 Fig. 13). If this specimen does represent a moult, then it is possible that it bore only a single cephalic sclerite. The appearance of a paired sclerite may have been caused by folding, shearing and breakage of a single, larger, sclerite during the moulting process. The smaller
<emphasis box="[1335,1340,1704,1724]" italics="true" pageId="3" pageNumber="4">‘</emphasis>
rectangular elements
<emphasis box="[877,882,1732,1752]" italics="true" pageId="3" pageNumber="4">’</emphasis>
posterior to this apparently paired sclerite may represent additional broken fragments.
</paragraph>
<paragraph blockId="3.[816,1498,1617,1981]" pageId="3" pageNumber="4">
If the presence of multiple head sclerites in NMW.2021.3 G.8 (rather than one in NMW.2021.3 G.7) is confirmed, then this difference and the distinct morphology of the dorsal spines on the proboscis together suggest that NMW.2021.3 G.7 may represent a distinct species. Other potentially diagnostic features of NMW.2021.3 G.8, such as the spinose caudal fan, are not preserved in NMW.2021.3 G.7, but are present in opabiniids
<taxonomicName box="[1030,1120,1961,1980]" family="Opabiniidae" genus="Opabinia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[1030,1120,1961,1980]" italics="true" pageId="3" pageNumber="4">Opabinia</emphasis>
</taxonomicName>
and
<emphasis bold="true" box="[1168,1256,1962,1981]" italics="true" pageId="3" pageNumber="4">Utaurora</emphasis>
<bibRefCitation author="Pates, S. &amp; Wolfe, J. M. &amp; Lerosey-Aubril, R. &amp; Daley, A. C. &amp; Ortega- Hernandez, J." box="[1256,1271,1957,1969]" journalOrPublisher="Proc. R. Soc. B" pageId="3" pageNumber="4" pagination="20212093" part="289" refId="ref9671" refString="36. Pates, S., Wolfe, J. M., Lerosey-Aubril, R., Daley, A. C. &amp; Ortega- Hernandez, J. New opabiniid diversifies the weirdest wonders of the euarthropod stem group. Proc. R. Soc. B 289, 20212093 (2022)." title="New opabiniid diversifies the weirdest wonders of the euarthropod stem group" type="journal article" year="2022">
<superScript attach="left" box="[1256,1271,1957,1969]" fontSize="5" pageId="3" pageNumber="4">36</superScript>
</bibRefCitation>
.
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7346854" ID-Zenodo-Dep="7346854" httpUri="https://zenodo.org/record/7346854/files/figure.png" pageId="4" pageNumber="5" startId="4.[106,135,1049,1066]" targetBox="[143,1460,134,1025]" targetPageId="4">
<paragraph blockId="4.[106,1498,1049,1093]" pageId="4" pageNumber="5">
<emphasis bold="true" box="[106,1184,1049,1067]" pageId="4" pageNumber="5">
Fig. 3 | Details of proboscis of
<taxonomicName authority="Pates &amp; Botting &amp; Muir &amp; Wolfe, 2022" authorityName="Pates &amp; Botting &amp; Muir &amp; Wolfe" authorityYear="2022" box="[371,553,1049,1067]" family="Opabiniidae" genus="Mieridduryn" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="5" phylum="Arthropoda" rank="species" species="bonniae" status="nov. gen. et sp.">
<emphasis bold="true" box="[371,553,1049,1067]" italics="true" pageId="4" pageNumber="5">Mieridduryn bonniae</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[557,692,1050,1066]" pageId="4" pageNumber="5" rank="species">nov. gen. et sp.</taxonomicNameLabel>
(NMW.2021.3 G.7) under different lighting conditions. a
</emphasis>
S8 microscope,cross-polarised light, stitched images, with contrast increased.
<emphasis bold="true" box="[451,463,1076,1092]" pageId="4" pageNumber="5">b</emphasis>
M125 microscope under high angle light. an annulation, sp spine.
</paragraph>
</caption>
<paragraph blockId="4.[106,788,1160,1838]" pageId="4" pageNumber="5">
Alternatively, NMW.2021.3 G.8 could alsobe regarded asan earlier ontogenetic stage of
<taxonomicName authorityName="Pates &amp; Botting &amp; Muir &amp; Wolfe" authorityYear="2022" box="[324,528,1190,1209]" family="Opabiniidae" genus="Mieridduryn" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="5" phylum="Arthropoda" rank="species" species="bonniae">
<emphasis bold="true" box="[324,528,1190,1209]" italics="true" pageId="4" pageNumber="5">Mieridduryn bonniae</emphasis>
</taxonomicName>
than NMW.2021.3 G.7. At ~3 mm, euarthropod A falls within the size range of some larger crustacean larvae (e.g. decapods, remipedes)
<bibRefCitation author="Martin, J. W. &amp; Olesen, J. &amp; Hoeg, J. T." box="[539,555,1243,1255]" journalOrPublisher="Johns Hopkins University Press" pageId="4" pageNumber="5" refId="ref9840" refString="40. Martin, J. W., Olesen, J. &amp; Hoeg, J. T. Atlas of Crustacean Larvae. (Johns Hopkins University Press, 2014)." title="Atlas of Crustacean Larvae" type="book" year="2014">
<superScript attach="right" box="[539,555,1243,1255]" fontSize="5" pageId="4" pageNumber="5">40</superScript>
</bibRefCitation>
; thus, postembryonic morphological changes such as differing spine form on the proboscis and numberofheadsclerites couldexplain the observeddifferences in the two specimens. Within radiodonts the number of head sclerites is stable at the family level, and does not change during ontogeny (in at least one species,
<taxonomicName authorityName="Cong, Ma, Hou, Edgecombe &amp; Strausfeld" authorityYear="2014" box="[292,515,1390,1409]" genus="Lyrarapax" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="5" rank="species" species="unguispinus">
<emphasis bold="true" box="[292,515,1390,1409]" italics="true" pageId="4" pageNumber="5">Lyrarapax unguispinus</emphasis>
</taxonomicName>
<bibRefCitation author="Liu, J. &amp; et al." box="[515,528,1386,1398]" journalOrPublisher="Natl Sci. Rev." pageId="4" pageNumber="5" pagination="863 - 869" part="5" refId="ref9874" refString="41. Liu, J. et al. Origin of raptorial feeding in juvenile euarthropods revealed by a Cambrian radiodontan. Natl Sci. Rev. 5, 863 - 869 (2018)." title="Origin of raptorial feeding in juvenile euarthropods revealed by a Cambrian radiodontan" type="journal article" year="2018">
<superScript attach="left" box="[515,528,1386,1398]" fontSize="5" pageId="4" pageNumber="5">41</superScript>
</bibRefCitation>
), but the number of carapace elements does change during ontogeny in other euarthropods. For example, in the fossil phosphatocopine
<taxonomicName authorityName="Huo, Shu &amp; Fu" authorityYear="1983" box="[537,670,1447,1466]" class="Ostracoda" genus="Dabashanella" higherTaxonomySource="CoL,GBIF" kingdom="Animalia" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[537,670,1447,1466]" italics="true" pageId="4" pageNumber="5">Dabashanella</emphasis>
</taxonomicName>
and the living ostracod
<taxonomicName authorityName="Hornibrook" authorityYear="1949" box="[259,344,1476,1495]" class="Ostracoda" family="Punciidae" genus="Manawa" kingdom="Animalia" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[259,344,1476,1495]" italics="true" pageId="4" pageNumber="5">Manawa</emphasis>
</taxonomicName>
, a univalved carapace develops into a bivalved one
<superScript attach="left" box="[237,269,1500,1512]" fontSize="5" pageId="4" pageNumber="5">
<bibRefCitation author="Maas, A. &amp; Waloszek, D. &amp; Muller, K." box="[237,251,1500,1512]" journalOrPublisher="Foss. Strat." pageId="4" pageNumber="5" pagination="1 - 238" part="49" refId="ref9910" refString="42. Maas, A., Waloszek, D. &amp; Muller, K. Morphology, ontogeny and phylogeny of the Phosphatocopina (Crustacea) from the Upper Cambrian ' Orsten' of Sweden. Foss. Strat. 49, 1 - 238 (2003)." title="Morphology, ontogeny and phylogeny of the Phosphatocopina (Crustacea) from the Upper Cambrian ' Orsten' of Sweden" type="journal article" year="2003">42</bibRefCitation>
,
<bibRefCitation author="Wakayama, N." box="[255,269,1500,1512]" journalOrPublisher="J. Zool." pageId="4" pageNumber="5" pagination="406 - 413" part="273" refId="ref9959" refString="43. Wakayama, N. Embryonic development clarifies polyphyly in ostracod crustaceans. J. Zool. 273, 406 - 413 (2007)." title="Embryonic development clarifies polyphyly in ostracod crustaceans" type="journal article" year="2007">43</bibRefCitation>
</superScript>
. Metamorphosis is expected to be ancestral for panarthropods
<bibRefCitation author="Wolfe, J. M." box="[252,267,1529,1541]" journalOrPublisher="Integr. Comp. Biol." pageId="4" pageNumber="5" pagination="499 - 509" part="57" refId="ref9986" refString="44. Wolfe, J. M. Metamorphosis Is Ancestral for Crown Euarthropods, and Evolved in the Cambrian or Earlier. Integr. Comp. Biol. 57, 499 - 509 (2017)." title="Metamorphosis Is Ancestral for Crown Euarthropods, and Evolved in the Cambrian or Earlier" type="journal article" year="2017">
<superScript attach="left" box="[252,267,1529,1541]" fontSize="5" pageId="4" pageNumber="5">44</superScript>
</bibRefCitation>
and has been observed in some members of the upper stem group (e.g. the megacheiran
<taxonomicName authorityName="Walcott" authorityYear="1912" box="[506,617,1561,1580]" family="Leanchoiliidae" genus="Leanchoilia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[506,617,1561,1580]" italics="true" pageId="4" pageNumber="5">Leanchoilia</emphasis>
</taxonomicName>
<bibRefCitation author="Liu, Y. &amp; et al." box="[617,631,1557,1569]" journalOrPublisher="PNAS" pageId="4" pageNumber="5" pagination="5542 - 5546" part="113" refId="ref10024" refString="45. Liu, Y. et al. Three-dimensionally preserved minute larva of a greatappendage arthropod from the early Cambrian Chengjiang biota. PNAS 113, 5542 - 5546 (2016)." title="Three-dimensionally preserved minute larva of a greatappendage arthropod from the early Cambrian Chengjiang biota" type="journal article" year="2016">
<superScript attach="left" box="[617,631,1557,1569]" fontSize="5" pageId="4" pageNumber="5">45</superScript>
</bibRefCitation>
, but not all, see ref. 46); hence a comparatively minor morphological change during ontogeny for NMW.2021.3 G.7 and NMW.2021.3 G.8 cannot be ruled out with the current data. Thus,we leave the smaller specimen in open nomenclature, and consider both possibilities (a distinct species or an earlier ontogenetic stage of
<taxonomicName authority="Pates &amp; Botting &amp; Muir &amp; Wolfe, 2022" authorityName="Pates &amp; Botting &amp; Muir &amp; Wolfe" authorityYear="2022" box="[375,493,1704,1723]" family="Opabiniidae" genus="Mieridduryn" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus" status="nov. gen.">
<emphasis bold="true" box="[375,493,1704,1723]" italics="true" pageId="4" pageNumber="5">Mieridduryn</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[496,588,1703,1724]" pageId="4" pageNumber="5" rank="genus">nov. gen.</taxonomicNameLabel>
) for this specimen in our phylogenetic analyses (section below).
</paragraph>
<paragraph blockId="4.[106,788,1160,1838]" pageId="4" pageNumber="5">
Further discussion of the relationship of
<taxonomicName authority="Pates &amp; Botting &amp; Muir &amp; Wolfe, 2022" authorityName="Pates &amp; Botting &amp; Muir &amp; Wolfe" authorityYear="2022" box="[568,685,1761,1780]" family="Opabiniidae" genus="Mieridduryn" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus" status="nov. gen.">
<emphasis bold="true" box="[568,685,1761,1780]" italics="true" pageId="4" pageNumber="5">Mieridduryn</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[694,787,1760,1781]" pageId="4" pageNumber="5" rank="genus">nov. gen.</taxonomicNameLabel>
and euarthropod A to opabiniids is provided in Supplementary Information.
</paragraph>
</subSubSection>
</treatment>
</document>
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