419 lines
58 KiB
XML
419 lines
58 KiB
XML
<document ID-DOI="http://doi.org/10.5281/zenodo.6229967" ID-GBIF-Dataset="da622693-9fe4-4554-b304-43dd49278f94" ID-GBIF-Taxon="120677360" ID-Zenodo-Dep="6229967" checkinTime="1462977918346" checkinUser="jeremy" docAuthor="Long, John A., Trinajstic, Kate, Young, Gavin C. & Senden, Tim" docDate="2008" docId="126A8965FF9F761AFF5EBA4EFD7EFA75" docLanguage="en" docName="nature06966.pdf" docOrigin="Nature 453" docStyle="DocumentStyle{}" docTitle="Materpiscis attenboroughi Long, Trinajstic, Young & Senden, 2008, sp. nov" docType="treatment" docVersion="7" lastPageId="2" lastPageNumber="652" masterDocId="EE53F11DFF9F7618FFBCBE39FFD7FFA4" masterDocTitle="Live birth in the Devonian period" masterLastPageNumber="653" masterPageNumber="650" pageId="0" pageNumber="650" updateTime="1645564576954" updateUser="ExternalLinkService">
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<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:titleInfo>
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<mods:title>Live birth in the Devonian period</mods:title>
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</mods:titleInfo>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Long, John A.</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Trinajstic, Kate</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Young, Gavin C.</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Senden, Tim</mods:namePart>
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</mods:name>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:relatedItem type="host">
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<mods:titleInfo>
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<mods:title>Nature</mods:title>
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</mods:titleInfo>
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<mods:part>
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<mods:date>2008</mods:date>
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<mods:detail type="volume">
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<mods:number>453</mods:number>
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</mods:detail>
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<mods:extent unit="page">
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<mods:start>650</mods:start>
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<mods:end>653</mods:end>
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</mods:extent>
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</mods:part>
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</mods:relatedItem>
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<mods:classification>journal article</mods:classification>
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<mods:identifier type="DOI">10.1038/nature06966</mods:identifier>
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<mods:identifier type="GBIF-Dataset">da622693-9fe4-4554-b304-43dd49278f94</mods:identifier>
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<mods:identifier type="Zenodo-Dep">270098</mods:identifier>
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<mods:identifier type="ZooBank">F4178D99-A31D-460B-ABE6-52097CB7F8AD</mods:identifier>
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</mods:mods>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.6229967" ID-GBIF-Taxon="120677360" ID-Zenodo-Dep="6229967" LSID="urn:lsid:plazi:treatment:126A8965FF9F761AFF5EBA4EFD7EFA75" httpUri="http://treatment.plazi.org/id/126A8965FF9F761AFF5EBA4EFD7EFA75" lastPageId="2" lastPageNumber="652" pageId="0" pageNumber="650">
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<subSubSection box="[226,633,1142,1164]" pageId="0" pageNumber="650" type="nomenclature">
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<paragraph blockId="0.[93,766,512,1919]" box="[226,633,1142,1164]" pageId="0" pageNumber="650">
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<taxonomicName authority="Long, Trinajstic, Young & Senden, 2008" authorityName="Long, Trinajstic, Young & Senden" authorityYear="2008" box="[226,476,1142,1164]" class="Placodermi" family="Ptyctodontidae" genus="Materpiscis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ptyctodontida" pageId="0" pageNumber="650" phylum="Chordata" rank="species" species="attenboroughi" status="sp. nov.">
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<emphasis box="[226,476,1142,1164]" italics="true" pageId="0" pageNumber="650">Materpiscis attenboroughi</emphasis>
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</taxonomicName>
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gen. et
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<taxonomicNameLabel box="[554,628,1143,1164]" pageId="0" pageNumber="650">sp. nov</taxonomicNameLabel>
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.
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</paragraph>
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</subSubSection>
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<subSubSection pageId="0" pageNumber="650" type="etymology">
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<paragraph blockId="0.[93,766,512,1919]" pageId="0" pageNumber="650">
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<emphasis bold="true" box="[93,210,1193,1214]" pageId="0" pageNumber="650">Etymology.</emphasis>
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Generic name from the Latin meaning ‘mother fish’; species name in honour of Sir David Attenborough, who first drew attention to the Gogo fish sites in his
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<number box="[468,515,1249,1270]" pageId="0" pageNumber="650" value="1979.0">1979</number>
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series
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<emphasis box="[582,708,1248,1270]" italics="true" pageId="0" pageNumber="650">Life on Earth</emphasis>
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.
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</paragraph>
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</subSubSection>
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<subSubSection pageId="0" pageNumber="650" type="materials_examined">
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<paragraph blockId="0.[93,766,512,1919]" pageId="0" pageNumber="650">
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<materialsCitationGroup pageId="0" pageNumber="650">
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<collectingRegionGroup pageId="0" pageNumber="650">
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<materialsCitation ID-GBIF-Occurrence="1265947869" collectionCode="Western Australian Museum, Perth" country="Australia" location="Stromatoporoid camp locality, Gogo Station, near Fitzroy Crossing" pageId="0" pageNumber="650" specimenCode="WAM 0 7.12.1" specimenCount="1" stateProvince="Western Australia" typeStatus="holotype">
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<emphasis bold="true" box="[93,195,1278,1299]" pageId="0" pageNumber="650">
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<typeStatus box="[93,191,1278,1299]" pageId="0" pageNumber="650">Holotype</typeStatus>
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.
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</emphasis>
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<specimenCode box="[201,340,1278,1299]" pageId="0" pageNumber="650">
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<specimenCode box="[201,283,1278,1299]" pageId="0" pageNumber="650">WAM 0</specimenCode>
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<number box="[284,328,1278,1299]" pageId="0" pageNumber="650" value="7.12">7.12</number>
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.
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<number box="[331,340,1278,1299]" pageId="0" pageNumber="650" value="1.0">1</number>
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</specimenCode>
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(
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<httpUri box="[353,721,1277,1300]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34981" pageId="0" pageNumber="650">
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<collectionCode box="[353,714,1277,1300]" pageId="0" pageNumber="650">Western Australian Museum, Perth</collectionCode>
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)
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</httpUri>
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.
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<emphasis bold="true" box="[93,272,1306,1328]" pageId="0" pageNumber="650">Age and locality.</emphasis>
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From the
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<location LSID="urn:lsid:plazi:treatment:126A8965FF9F761AFF5EBA4EFD7EFA75:9F1C6EA8FF9F7618FE38BB23FE58FAE9" country="Australia" name="Stromatoporoid camp locality, Gogo Station, near Fitzroy Crossing" pageId="0" pageNumber="650" stateProvince="Western Australia">Stromatoporoid camp locality, Gogo Station, near Fitzroy Crossing</location>
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,
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<collectingRegion box="[412,593,1334,1357]" pageId="0" pageNumber="650">Western Australia</collectingRegion>
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(Late Devonian, early Frasnian)
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</materialsCitation>
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.
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</collectingRegionGroup>
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</materialsCitationGroup>
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</paragraph>
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</subSubSection>
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<subSubSection pageId="0" pageNumber="650" type="diagnosis">
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<paragraph blockId="0.[93,766,512,1919]" pageId="0" pageNumber="650">
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<emphasis bold="true" box="[93,201,1391,1413]" pageId="0" pageNumber="650">Diagnosis.</emphasis>
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<botanicCollectionCode box="[206,222,1392,1413]" pageId="0" pageNumber="650">A</botanicCollectionCode>
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small aspinothoracid ptyctodontid fish having an ante- riorly pointed nuchal plate that participates in the posterior margin of the skull roof, broad roughly triangular-shaped preorbitals that meet mesially; the marginal plate has a large postorbital region with parallel rows of tubercles adorning it; the submarginal is strap-like and strongly curved mesially; robust triturating tooth plates that meet only at anterior tips, superognathals with moderately high anterior dorsal process. The body is scaleless.
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</paragraph>
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</subSubSection>
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<subSubSection lastPageId="2" lastPageNumber="652" pageId="0" pageNumber="650" type="description">
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<paragraph blockId="0.[93,766,512,1919]" lastBlockId="0.[802,1476,510,1919]" pageId="0" pageNumber="650">
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<emphasis bold="true" box="[93,221,1616,1638]" pageId="0" pageNumber="650">Description.</emphasis>
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<taxonomicName authority="Long, Trinajstic, Young & Senden, 2008" authorityName="Long, Trinajstic, Young & Senden" authorityYear="2008" box="[226,335,1617,1638]" class="Placodermi" family="Ptyctodontidae" genus="Materpiscis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ptyctodontida" pageId="0" pageNumber="650" phylum="Chordata" rank="genus" status="gen. nov.">
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<emphasis box="[226,335,1617,1638]" italics="true" pageId="0" pageNumber="650">Materpiscis</emphasis>
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</taxonomicName>
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<taxonomicNameLabel box="[341,431,1617,1638]" pageId="0" pageNumber="650" rank="genus">gen. nov.</taxonomicNameLabel>
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(
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<figureCitation box="[442,526,1616,1639]" captionStart-0="Figure 1" captionStart-1="Figure 2" captionStart-2="Figure 3" captionStartId-0="1.[112,170,1137,1156]" captionStartId-1="1.[1056,1114,1439,1458]" captionStartId-2="2.[93,151,798,817]" captionTargetBox-0="[273,774,416,1105]" captionTargetBox-1="[113,920,1444,1995]" captionTargetBox-2="[122,735,134,777]" captionTargetId-0="figure@1.[271,677,414,838]" captionTargetId-1="figure@1.[111,922,1439,1997]" captionTargetId-2="figure@2.[122,736,405,777]" captionTargetPageId-0="1" captionTargetPageId-1="1" captionTargetPageId-2="2" captionText-0="Figure 1 | Holotype of Materpiscis attenboroughi gen. et sp. nov. (WAM 07.12.1). a, Body with embryo. The boxed area is magnified in panel c. b, Embryo at early preparation. c, d, Umbilical cord and embryo at later preparation. e, Adult skull roof in dorsal view. f, Adult left marginal. h, Dermal armour restored in lateral view. i, Left adult tooth plates in mesial view. g, Austroptyctodus gardineri (WAM 86.9.662) showing embryos. ADL, anterior dorsolateral plate; AL, anterior lateral plate; Ce, central; df. bas, dorsal fin basal plate of adult; haem, haemal arch; Ifg, inferognathal; MD, median dorsal plate; Mg, marginal; neur, neural arch; Nu, nuchal; PNu, paranuchal plate; PRO, preorbital; PTO, postorbital; Sgn, supragnathal; SM, submarginal plate; umb, umbilical cord." captionText-1="Figure 2 | Holotype of Materpiscis attenboroughi gen. et sp. nov. (WAM 07.12.1). a – e, SEM images of umbilical cord. a, Umbilicus section. b, Image showing a vascular canal (v). c, image showing pit (p). d, Image showing possible appendiculae (app) and pit (p). e, Image showing outer epithelium surface (ep) and stalk wall. f, High-resolution computer tomogram (HRCT) showing embryonic bones. g, HRCT tomogram showing supragnathals in occlusion. h, HRCT showing umbilicus (white arrowheads). Abbreviations as for Fig. 1 except for: ep, epithelium; sym, symphysis." captionText-2="Figure 3 | Materpiscis gen. nov. a, Diagram showing position of embryo and yolk sac within the mother. b, Artist’s reconstruction of Materpiscis gen. nov. giving birth (by B. Choo)." httpUri-0="https://zenodo.org/record/270100/files/figure.png" httpUri-1="https://zenodo.org/record/270101/files/figure.png" httpUri-2="https://zenodo.org/record/270102/files/figure.png" pageId="0" pageNumber="650">
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Figs
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<number box="[489,526,1617,1639]" pageId="0" pageNumber="650" value="2.0" valueMax="3.0" valueMin="1.0">1–3</number>
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</figureCitation>
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) is readily distinguished from all other known ptyctodontids by the combination of lacking a spinal plate with anteriorly pointed nuchal plate (
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<figureCitation box="[611,670,1672,1695]" captionStart="Figure 1" captionStartId="1.[112,170,1137,1156]" captionTargetBox="[273,774,416,1105]" captionTargetId="figure@1.[271,677,414,838]" captionTargetPageId="1" captionText="Figure 1 | Holotype of Materpiscis attenboroughi gen. et sp. nov. (WAM 07.12.1). a, Body with embryo. The boxed area is magnified in panel c. b, Embryo at early preparation. c, d, Umbilical cord and embryo at later preparation. e, Adult skull roof in dorsal view. f, Adult left marginal. h, Dermal armour restored in lateral view. i, Left adult tooth plates in mesial view. g, Austroptyctodus gardineri (WAM 86.9.662) showing embryos. ADL, anterior dorsolateral plate; AL, anterior lateral plate; Ce, central; df. bas, dorsal fin basal plate of adult; haem, haemal arch; Ifg, inferognathal; MD, median dorsal plate; Mg, marginal; neur, neural arch; Nu, nuchal; PNu, paranuchal plate; PRO, preorbital; PTO, postorbital; Sgn, supragnathal; SM, submarginal plate; umb, umbilical cord." httpUri="https://zenodo.org/record/270100/files/figure.png" pageId="0" pageNumber="650">
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||
Fig.
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||
<number box="[660,670,1673,1695]" pageId="0" pageNumber="650" value="1.0">1</number>
|
||
</figureCitation>
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||
e), and a much-expanded postorbital region on the marginal plate (
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<figureCitation box="[684,741,1700,1723]" captionStart="Figure 1" captionStartId="1.[112,170,1137,1156]" captionTargetBox="[273,774,416,1105]" captionTargetId="figure@1.[271,677,414,838]" captionTargetPageId="1" captionText="Figure 1 | Holotype of Materpiscis attenboroughi gen. et sp. nov. (WAM 07.12.1). a, Body with embryo. The boxed area is magnified in panel c. b, Embryo at early preparation. c, d, Umbilical cord and embryo at later preparation. e, Adult skull roof in dorsal view. f, Adult left marginal. h, Dermal armour restored in lateral view. i, Left adult tooth plates in mesial view. g, Austroptyctodus gardineri (WAM 86.9.662) showing embryos. ADL, anterior dorsolateral plate; AL, anterior lateral plate; Ce, central; df. bas, dorsal fin basal plate of adult; haem, haemal arch; Ifg, inferognathal; MD, median dorsal plate; Mg, marginal; neur, neural arch; Nu, nuchal; PNu, paranuchal plate; PRO, preorbital; PTO, postorbital; Sgn, supragnathal; SM, submarginal plate; umb, umbilical cord." httpUri="https://zenodo.org/record/270100/files/figure.png" pageId="0" pageNumber="650">
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Fig.
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<number box="[731,741,1700,1723]" pageId="0" pageNumber="650" value="1.0">1</number>
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||
</figureCitation>
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||
f). The nuchal plate participates in the posterior margin of the skull roof and the centrals are unusually broad (
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<figureCitation box="[494,553,1756,1779]" captionStart="Figure 1" captionStartId="1.[112,170,1137,1156]" captionTargetBox="[273,774,416,1105]" captionTargetId="figure@1.[271,677,414,838]" captionTargetPageId="1" captionText="Figure 1 | Holotype of Materpiscis attenboroughi gen. et sp. nov. (WAM 07.12.1). a, Body with embryo. The boxed area is magnified in panel c. b, Embryo at early preparation. c, d, Umbilical cord and embryo at later preparation. e, Adult skull roof in dorsal view. f, Adult left marginal. h, Dermal armour restored in lateral view. i, Left adult tooth plates in mesial view. g, Austroptyctodus gardineri (WAM 86.9.662) showing embryos. ADL, anterior dorsolateral plate; AL, anterior lateral plate; Ce, central; df. bas, dorsal fin basal plate of adult; haem, haemal arch; Ifg, inferognathal; MD, median dorsal plate; Mg, marginal; neur, neural arch; Nu, nuchal; PNu, paranuchal plate; PRO, preorbital; PTO, postorbital; Sgn, supragnathal; SM, submarginal plate; umb, umbilical cord." httpUri="https://zenodo.org/record/270100/files/figure.png" pageId="0" pageNumber="650">
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Fig.
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<number box="[543,553,1757,1779]" pageId="0" pageNumber="650" value="1.0">1</number>
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</figureCitation>
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e). The skull roof is unusual in having prominent raised ridges for the main sensory-line canals (
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<figureCitation box="[168,225,1812,1835]" captionStart="Figure 1" captionStartId="1.[112,170,1137,1156]" captionTargetBox="[273,774,416,1105]" captionTargetId="figure@1.[271,677,414,838]" captionTargetPageId="1" captionText="Figure 1 | Holotype of Materpiscis attenboroughi gen. et sp. nov. (WAM 07.12.1). a, Body with embryo. The boxed area is magnified in panel c. b, Embryo at early preparation. c, d, Umbilical cord and embryo at later preparation. e, Adult skull roof in dorsal view. f, Adult left marginal. h, Dermal armour restored in lateral view. i, Left adult tooth plates in mesial view. g, Austroptyctodus gardineri (WAM 86.9.662) showing embryos. ADL, anterior dorsolateral plate; AL, anterior lateral plate; Ce, central; df. bas, dorsal fin basal plate of adult; haem, haemal arch; Ifg, inferognathal; MD, median dorsal plate; Mg, marginal; neur, neural arch; Nu, nuchal; PNu, paranuchal plate; PRO, preorbital; PTO, postorbital; Sgn, supragnathal; SM, submarginal plate; umb, umbilical cord." httpUri="https://zenodo.org/record/270100/files/figure.png" pageId="0" pageNumber="650">
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Fig.
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<number box="[215,225,1813,1835]" pageId="0" pageNumber="650" value="1.0">1</number>
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</figureCitation>
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||
e). The tooth plates can be moved against each other as in life. This shows that only the anterior-most biting edges of the jaws met in full occlusion (
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<figureCitation box="[409,467,1868,1891]" captionStart="Figure 1" captionStartId="1.[112,170,1137,1156]" captionTargetBox="[273,774,416,1105]" captionTargetId="figure@1.[271,677,414,838]" captionTargetPageId="1" captionText="Figure 1 | Holotype of Materpiscis attenboroughi gen. et sp. nov. (WAM 07.12.1). a, Body with embryo. The boxed area is magnified in panel c. b, Embryo at early preparation. c, d, Umbilical cord and embryo at later preparation. e, Adult skull roof in dorsal view. f, Adult left marginal. h, Dermal armour restored in lateral view. i, Left adult tooth plates in mesial view. g, Austroptyctodus gardineri (WAM 86.9.662) showing embryos. ADL, anterior dorsolateral plate; AL, anterior lateral plate; Ce, central; df. bas, dorsal fin basal plate of adult; haem, haemal arch; Ifg, inferognathal; MD, median dorsal plate; Mg, marginal; neur, neural arch; Nu, nuchal; PNu, paranuchal plate; PRO, preorbital; PTO, postorbital; Sgn, supragnathal; SM, submarginal plate; umb, umbilical cord." httpUri="https://zenodo.org/record/270100/files/figure.png" pageId="0" pageNumber="650">
|
||
Fig.
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||
<date box="[456,475,1868,1891]" pageId="0" pageNumber="650">
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||
<number box="[456,467,1868,1891]" pageId="0" pageNumber="650" value="1.0">1</number>
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||
i
|
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</date>
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||
</figureCitation>
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||
). The trunk shield plates do not differ significantly from those of many other aspinothoracid ptyctodontids, in particular
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<taxonomicName authority="Long, 1997" authorityName="Long" authorityYear="1997" box="[1079,1233,512,533]" class="Placodermi" family="Ptyctodontidae" genus="Austroptyctodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ptyctodontida" pageId="0" pageNumber="650" phylum="Chordata" rank="genus">
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<emphasis box="[1079,1233,512,533]" italics="true" pageId="0" pageNumber="650">Austroptyctodus</emphasis>
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</taxonomicName>
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<bibRefCitation author="Long, J. A." box="[1234,1242,510,522]" pageId="0" pageNumber="650" refString="7. Long, J. A. Ptyctodontid fishes (Vertebrata, Placodermi) from the Late Devonian Gogo Formation, Western Australia, with a revision of the European genus Ctenurella Ørvig, 1960. Geodiversitas 19, 515–555 (1997)." type="journal article" year="1997">
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||
<number box="[1234,1242,510,522]" pageId="0" pageNumber="650" value="7.0">7</number>
|
||
</bibRefCitation>
|
||
, except for proportions and ornamentation, the median dorsal having a low median crest. A restoration of the dermal armour is shown in
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<figureCitation box="[1284,1340,568,591]" captionStart="Figure 1" captionStartId="1.[112,170,1137,1156]" captionTargetBox="[273,774,416,1105]" captionTargetId="figure@1.[271,677,414,838]" captionTargetPageId="1" captionText="Figure 1 | Holotype of Materpiscis attenboroughi gen. et sp. nov. (WAM 07.12.1). a, Body with embryo. The boxed area is magnified in panel c. b, Embryo at early preparation. c, d, Umbilical cord and embryo at later preparation. e, Adult skull roof in dorsal view. f, Adult left marginal. h, Dermal armour restored in lateral view. i, Left adult tooth plates in mesial view. g, Austroptyctodus gardineri (WAM 86.9.662) showing embryos. ADL, anterior dorsolateral plate; AL, anterior lateral plate; Ce, central; df. bas, dorsal fin basal plate of adult; haem, haemal arch; Ifg, inferognathal; MD, median dorsal plate; Mg, marginal; neur, neural arch; Nu, nuchal; PNu, paranuchal plate; PRO, preorbital; PTO, postorbital; Sgn, supragnathal; SM, submarginal plate; umb, umbilical cord." httpUri="https://zenodo.org/record/270100/files/figure.png" pageId="0" pageNumber="650">
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Fig.
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||
<number box="[1330,1340,568,591]" pageId="0" pageNumber="650" value="1.0">1</number>
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</figureCitation>
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h.
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<botanicCollectionCode box="[1371,1387,569,590]" pageId="0" pageNumber="650">A</botanicCollectionCode>
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||
detailed description of the dermal and axial skeleton, braincase and dentition of
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<taxonomicName authority="Long, Trinajstic, Young & Senden, 2008" authorityName="Long, Trinajstic, Young & Senden" authorityYear="2008" box="[828,937,626,647]" class="Placodermi" family="Ptyctodontidae" genus="Materpiscis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ptyctodontida" pageId="0" pageNumber="650" phylum="Chordata" rank="genus" status="gen. nov.">
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<emphasis box="[828,937,626,647]" italics="true" pageId="0" pageNumber="650">Materpiscis</emphasis>
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</taxonomicName>
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<taxonomicNameLabel box="[943,1035,626,647]" pageId="0" pageNumber="650" rank="genus">gen. nov.</taxonomicNameLabel>
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, and its embryonic skeletal morphology, will be published elsewhere.
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</paragraph>
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<paragraph blockId="0.[802,1476,510,1919]" pageId="0" pageNumber="650">
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The Gogo Formation has yielded the earliest phosphatized gnathostome muscle tissues and nerve fibres
|
||
<bibRefCitation author=" Trinajstic, K., Marshall, E., Long, J. & Bifield, K." box="[1238,1246,708,721]" pageId="0" pageNumber="650" refString="9. Trinajstic, K., Marshall, E., Long, J. & Bifield, K. Exceptional preservation of nerve and muscle tissues in Devonian placoderm fish and their phylogenetic implications. Biol. Lett. 3, 197–200 (2007)." type="journal article" year="2007">
|
||
<number box="[1238,1246,708,721]" pageId="0" pageNumber="650" value="9.0">9</number>
|
||
</bibRefCitation>
|
||
, and the new specimen shows additional soft-tissue preservation never before recorded in any fossil. The new specimen is exceptional in revealing a small partial skeleton located within the upper body cavity of a pregnant, adult female ptyctodontid placoderm (
|
||
<figureCitation box="[1209,1268,823,846]" captionStart="Figure 1" captionStartId="1.[112,170,1137,1156]" captionTargetBox="[273,774,416,1105]" captionTargetId="figure@1.[271,677,414,838]" captionTargetPageId="1" captionText="Figure 1 | Holotype of Materpiscis attenboroughi gen. et sp. nov. (WAM 07.12.1). a, Body with embryo. The boxed area is magnified in panel c. b, Embryo at early preparation. c, d, Umbilical cord and embryo at later preparation. e, Adult skull roof in dorsal view. f, Adult left marginal. h, Dermal armour restored in lateral view. i, Left adult tooth plates in mesial view. g, Austroptyctodus gardineri (WAM 86.9.662) showing embryos. ADL, anterior dorsolateral plate; AL, anterior lateral plate; Ce, central; df. bas, dorsal fin basal plate of adult; haem, haemal arch; Ifg, inferognathal; MD, median dorsal plate; Mg, marginal; neur, neural arch; Nu, nuchal; PNu, paranuchal plate; PRO, preorbital; PTO, postorbital; Sgn, supragnathal; SM, submarginal plate; umb, umbilical cord." httpUri="https://zenodo.org/record/270100/files/figure.png" pageId="0" pageNumber="650">
|
||
Fig.
|
||
<number box="[1259,1268,823,846]" pageId="0" pageNumber="650" value="1.0">1</number>
|
||
</figureCitation>
|
||
a–d). The relatively complete adult skeleton includes much of the tail (
|
||
<figureCitation box="[1339,1398,851,874]" captionStart="Figure 1" captionStartId="1.[112,170,1137,1156]" captionTargetBox="[273,774,416,1105]" captionTargetId="figure@1.[271,677,414,838]" captionTargetPageId="1" captionText="Figure 1 | Holotype of Materpiscis attenboroughi gen. et sp. nov. (WAM 07.12.1). a, Body with embryo. The boxed area is magnified in panel c. b, Embryo at early preparation. c, d, Umbilical cord and embryo at later preparation. e, Adult skull roof in dorsal view. f, Adult left marginal. h, Dermal armour restored in lateral view. i, Left adult tooth plates in mesial view. g, Austroptyctodus gardineri (WAM 86.9.662) showing embryos. ADL, anterior dorsolateral plate; AL, anterior lateral plate; Ce, central; df. bas, dorsal fin basal plate of adult; haem, haemal arch; Ifg, inferognathal; MD, median dorsal plate; Mg, marginal; neur, neural arch; Nu, nuchal; PNu, paranuchal plate; PRO, preorbital; PTO, postorbital; Sgn, supragnathal; SM, submarginal plate; umb, umbilical cord." httpUri="https://zenodo.org/record/270100/files/figure.png" pageId="0" pageNumber="650">
|
||
Fig.
|
||
<number box="[1388,1398,852,874]" pageId="0" pageNumber="650" value="1.0">1</number>
|
||
</figureCitation>
|
||
a). The single, partially ossified juvenile skeleton inside the body cavity of the adult clearly belongs to the same species based on characteristic tooth-plate and marginal plate morphology (
|
||
<figureCitation box="[1257,1314,936,959]" captionStart="Figure 1" captionStartId="1.[112,170,1137,1156]" captionTargetBox="[273,774,416,1105]" captionTargetId="figure@1.[271,677,414,838]" captionTargetPageId="1" captionText="Figure 1 | Holotype of Materpiscis attenboroughi gen. et sp. nov. (WAM 07.12.1). a, Body with embryo. The boxed area is magnified in panel c. b, Embryo at early preparation. c, d, Umbilical cord and embryo at later preparation. e, Adult skull roof in dorsal view. f, Adult left marginal. h, Dermal armour restored in lateral view. i, Left adult tooth plates in mesial view. g, Austroptyctodus gardineri (WAM 86.9.662) showing embryos. ADL, anterior dorsolateral plate; AL, anterior lateral plate; Ce, central; df. bas, dorsal fin basal plate of adult; haem, haemal arch; Ifg, inferognathal; MD, median dorsal plate; Mg, marginal; neur, neural arch; Nu, nuchal; PNu, paranuchal plate; PRO, preorbital; PTO, postorbital; Sgn, supragnathal; SM, submarginal plate; umb, umbilical cord." httpUri="https://zenodo.org/record/270100/files/figure.png" pageId="0" pageNumber="650">
|
||
Fig.
|
||
<number box="[1304,1314,937,958]" pageId="0" pageNumber="650" value="1.0">1</number>
|
||
</figureCitation>
|
||
c, d). The small internal individual must be an embryo, rather than an ingested prey item, because the delicate bones show no breakage or etching from stomach acids, and the dentition is of similar gross morphology to that of the adult. The location of the preserved embryo close to the vertebral column suggests that it is in the uterus rather than the gut, a view supported by the presence of mineralized soft tissue forming a placental connection between embryo and mother. Furthermore, a second specimen from Gogo, previously described as
|
||
<taxonomicName authority="Long, 1997" authorityName="Long" authorityYear="1997" box="[1320,1474,1162,1183]" class="Placodermi" family="Ptyctodontidae" genus="Austroptyctodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ptyctodontida" pageId="0" pageNumber="650" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1320,1474,1162,1183]" italics="true" pageId="0" pageNumber="650">Austroptyctodus</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicName authority="Long, 1997" authorityName="Long" authorityYear="1997" box="[802,890,1190,1211]" class="Placodermi" family="Ptyctodontidae" genus="Austroptyctodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ptyctodontida" pageId="0" pageNumber="650" phylum="Chordata" rank="species" species="gardineri">
|
||
<emphasis box="[802,890,1190,1211]" italics="true" pageId="0" pageNumber="650">gardineri</emphasis>
|
||
</taxonomicName>
|
||
<bibRefCitation author="Long, J. A." box="[890,897,1188,1200]" pageId="0" pageNumber="650" refString="7. Long, J. A. Ptyctodontid fishes (Vertebrata, Placodermi) from the Late Devonian Gogo Formation, Western Australia, with a revision of the European genus Ctenurella Ørvig, 1960. Geodiversitas 19, 515–555 (1997)." type="journal article" year="1997">
|
||
<emphasis box="[890,897,1188,1200]" italics="true" pageId="0" pageNumber="650">
|
||
<number box="[890,897,1188,1200]" pageId="0" pageNumber="650" value="7.0">7</number>
|
||
</emphasis>
|
||
</bibRefCitation>
|
||
, has been re-examined and we have identified three small embryos in the same position posterior to the adult anterior lateral plate as in
|
||
<taxonomicName authority="Long, Trinajstic, Young & Senden, 2008" authorityName="Long, Trinajstic, Young & Senden" authorityYear="2008" box="[925,1034,1247,1268]" class="Placodermi" family="Ptyctodontidae" genus="Materpiscis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ptyctodontida" pageId="0" pageNumber="650" phylum="Chordata" rank="genus" status="gen. nov.">
|
||
<emphasis box="[925,1034,1247,1268]" italics="true" pageId="0" pageNumber="650">Materpiscis</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel box="[1047,1144,1247,1268]" pageId="0" pageNumber="650" rank="genus">gen. nov.</taxonomicNameLabel>
|
||
(
|
||
<figureCitation box="[1162,1222,1246,1269]" captionStart="Figure 1" captionStartId="1.[112,170,1137,1156]" captionTargetBox="[273,774,416,1105]" captionTargetId="figure@1.[271,677,414,838]" captionTargetPageId="1" captionText="Figure 1 | Holotype of Materpiscis attenboroughi gen. et sp. nov. (WAM 07.12.1). a, Body with embryo. The boxed area is magnified in panel c. b, Embryo at early preparation. c, d, Umbilical cord and embryo at later preparation. e, Adult skull roof in dorsal view. f, Adult left marginal. h, Dermal armour restored in lateral view. i, Left adult tooth plates in mesial view. g, Austroptyctodus gardineri (WAM 86.9.662) showing embryos. ADL, anterior dorsolateral plate; AL, anterior lateral plate; Ce, central; df. bas, dorsal fin basal plate of adult; haem, haemal arch; Ifg, inferognathal; MD, median dorsal plate; Mg, marginal; neur, neural arch; Nu, nuchal; PNu, paranuchal plate; PRO, preorbital; PTO, postorbital; Sgn, supragnathal; SM, submarginal plate; umb, umbilical cord." httpUri="https://zenodo.org/record/270100/files/figure.png" pageId="0" pageNumber="650">
|
||
Fig.
|
||
<quantity box="[1212,1237,1247,1269]" metricMagnitude="-3" metricUnit="kg" metricValue="1.0" pageId="0" pageNumber="650" unit="g" value="1.0">1g</quantity>
|
||
</figureCitation>
|
||
). Cases demonstrating viviparity in the fossil record are extremely rare, the best being Mesozoic ichthyosaurs
|
||
<bibRefCitation box="[1038,1072,1301,1314]" pageId="0" pageNumber="650">10,11</bibRefCitation>
|
||
and mosasauroids
|
||
<bibRefCitation author="Caldwell, M. W.& Lee, M. S. Y." box="[1279,1294,1301,1314]" pageId="0" pageNumber="650" refString="12. Caldwell, M. W.& Lee, M. S. Y. Live birth in marine lizards (mosasauroids). Proc. R. Soc. Lond. B 268, 2397–2401 (2001)." type="journal article" year="2001">
|
||
<number box="[1279,1294,1301,1314]" pageId="0" pageNumber="650" value="12.0">12</number>
|
||
</bibRefCitation>
|
||
. Presumed fetal specimens of a holocephalan fish from the Lower Carboniferous of Montana
|
||
<date box="[893,965,1357,1381]" pageId="0" pageNumber="650" value="2013-05">
|
||
<bibRefCitation author="Lund, R." box="[893,908,1357,1370]" pageId="0" pageNumber="650" refString="13. Lund, R. Viviparity and interuterine feeding in a new holocephalan fish from the Lower Carboniferous of Montana. Science 209, 697–699 (1980)." type="journal article" year="1980">
|
||
<number box="[893,908,1357,1370]" pageId="0" pageNumber="650" value="13.0">13</number>
|
||
</bibRefCitation>
|
||
may
|
||
</date>
|
||
extend the record of viviparity back into the Palaeozoic era, but were not found in or near the body cavity of the mother. Our new example extends the definite record of vertebrate viviparity back by some
|
||
<number box="[1150,1186,1445,1466]" pageId="0" pageNumber="650" value="200.0">200</number>
|
||
million years.
|
||
</paragraph>
|
||
<paragraph blockId="0.[802,1476,510,1919]" pageId="0" pageNumber="650">
|
||
The embryos of
|
||
<taxonomicName authority="Long, Trinajstic, Young & Senden, 2008" authorityName="Long, Trinajstic, Young & Senden" authorityYear="2008" box="[1004,1113,1474,1495]" class="Placodermi" family="Ptyctodontidae" genus="Materpiscis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ptyctodontida" pageId="0" pageNumber="650" phylum="Chordata" rank="genus" status="gen. nov.">
|
||
<emphasis box="[1004,1113,1474,1495]" italics="true" pageId="0" pageNumber="650">Materpiscis</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel box="[1124,1220,1474,1495]" pageId="0" pageNumber="650" rank="genus">gen. nov.</taxonomicNameLabel>
|
||
and
|
||
<taxonomicName authority="Long, 1997" authorityName="Long" authorityYear="1997" box="[1279,1433,1473,1494]" class="Placodermi" family="Ptyctodontidae" genus="Austroptyctodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ptyctodontida" pageId="0" pageNumber="650" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1279,1433,1473,1494]" italics="true" pageId="0" pageNumber="650">Austroptyctodus</emphasis>
|
||
</taxonomicName>
|
||
are about
|
||
<number box="[868,893,1501,1524]" pageId="0" pageNumber="650" value="25.0">25</number>
|
||
% of the adult size, based on proportions of the anterior lateral plate. The embryonic bones are remarkably thin and fragile, yet perfectly preserved, with the upper and lower dentition still in occlusion in
|
||
<taxonomicName authority="Long, Trinajstic, Young & Senden, 2008" authorityName="Long, Trinajstic, Young & Senden" authorityYear="2008" box="[926,1035,1587,1608]" class="Placodermi" family="Ptyctodontidae" genus="Materpiscis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ptyctodontida" pageId="0" pageNumber="650" phylum="Chordata" rank="genus" status="gen. nov.">
|
||
<emphasis box="[926,1035,1587,1608]" italics="true" pageId="0" pageNumber="650">Materpiscis</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel box="[1039,1129,1587,1608]" pageId="0" pageNumber="650" rank="genus">gen. nov.</taxonomicNameLabel>
|
||
(
|
||
<figureCitation box="[1139,1193,1586,1609]" captionStart="Figure 1" captionStartId="1.[112,170,1137,1156]" captionTargetBox="[273,774,416,1105]" captionTargetId="figure@1.[271,677,414,838]" captionTargetPageId="1" captionText="Figure 1 | Holotype of Materpiscis attenboroughi gen. et sp. nov. (WAM 07.12.1). a, Body with embryo. The boxed area is magnified in panel c. b, Embryo at early preparation. c, d, Umbilical cord and embryo at later preparation. e, Adult skull roof in dorsal view. f, Adult left marginal. h, Dermal armour restored in lateral view. i, Left adult tooth plates in mesial view. g, Austroptyctodus gardineri (WAM 86.9.662) showing embryos. ADL, anterior dorsolateral plate; AL, anterior lateral plate; Ce, central; df. bas, dorsal fin basal plate of adult; haem, haemal arch; Ifg, inferognathal; MD, median dorsal plate; Mg, marginal; neur, neural arch; Nu, nuchal; PNu, paranuchal plate; PRO, preorbital; PTO, postorbital; Sgn, supragnathal; SM, submarginal plate; umb, umbilical cord." httpUri="https://zenodo.org/record/270100/files/figure.png" pageId="0" pageNumber="650">
|
||
Figs
|
||
<number box="[1185,1193,1586,1609]" pageId="0" pageNumber="650" value="1.0">1</number>
|
||
</figureCitation>
|
||
b–d and
|
||
<quantity box="[1281,1307,1586,1608]" metricMagnitude="-3" metricUnit="kg" metricValue="2.0" pageId="0" pageNumber="650" unit="g" value="2.0">2g</quantity>
|
||
). The skull bones of this specimen are oriented posteriorly, indicating that the embryo was curled. Only some dermal plates are represented in the embryos. Absence of certain bones at an early developmental stage is also recorded in tiny (
|
||
<quantity box="[994,1103,1699,1722]" metricMagnitude="-2" metricUnit="m" metricValue="1.5" metricValueMax="1.6" metricValueMin="1.4" pageId="0" pageNumber="650" unit="mm" value="15.0" valueMax="16.0" valueMin="14.0">14–16 mm</quantity>
|
||
) juvenile skeletons of the antiarch placoderm
|
||
<taxonomicName box="[913,1016,1727,1748]" class="Insecta" family="Tortricidae" genus="Asterolepis" higherTaxonomySource="CoL" kingdom="Animalia" order="Lepidoptera" pageId="0" pageNumber="650" phylum="Arthropoda" rank="genus">
|
||
<emphasis box="[913,1016,1727,1748]" italics="true" pageId="0" pageNumber="650">Asterolepis</emphasis>
|
||
</taxonomicName>
|
||
<bibRefCitation author="Upeniece, I." box="[1015,1030,1725,1738]" pageId="0" pageNumber="650" refString="14. Upeniece, I. The unique fossil assemblage from the Lode Quarry (Upper Devonian, Latvia). Mitt. Mus. Natkd. Berl. Geowiss. 4, 101–119 (2001)." type="journal article" year="2001">
|
||
<number box="[1015,1030,1725,1738]" pageId="0" pageNumber="650" value="14.0">14</number>
|
||
</bibRefCitation>
|
||
. Identified embryonic bones comprise paired preorbitals, a paranuchal, anterior dorsolateral, anterior lateral and a marginal plate, and both sets of tooth plates in articulation (
|
||
<figureCitation box="[809,872,1811,1834]" captionStart="Figure 1" captionStartId="1.[112,170,1137,1156]" captionTargetBox="[273,774,416,1105]" captionTargetId="figure@1.[271,677,414,838]" captionTargetPageId="1" captionText="Figure 1 | Holotype of Materpiscis attenboroughi gen. et sp. nov. (WAM 07.12.1). a, Body with embryo. The boxed area is magnified in panel c. b, Embryo at early preparation. c, d, Umbilical cord and embryo at later preparation. e, Adult skull roof in dorsal view. f, Adult left marginal. h, Dermal armour restored in lateral view. i, Left adult tooth plates in mesial view. g, Austroptyctodus gardineri (WAM 86.9.662) showing embryos. ADL, anterior dorsolateral plate; AL, anterior lateral plate; Ce, central; df. bas, dorsal fin basal plate of adult; haem, haemal arch; Ifg, inferognathal; MD, median dorsal plate; Mg, marginal; neur, neural arch; Nu, nuchal; PNu, paranuchal plate; PRO, preorbital; PTO, postorbital; Sgn, supragnathal; SM, submarginal plate; umb, umbilical cord." httpUri="https://zenodo.org/record/270100/files/figure.png" pageId="0" pageNumber="650">
|
||
Figs
|
||
<number box="[863,872,1811,1834]" pageId="0" pageNumber="650" value="1.0">1</number>
|
||
</figureCitation>
|
||
b–d, g and
|
||
<quantity box="[1004,1029,1812,1833]" metricMagnitude="-3" metricUnit="kg" metricValue="2.0" pageId="0" pageNumber="650" unit="g" value="2.0">2g</quantity>
|
||
, f). The supragnathals of
|
||
<taxonomicName authority="Long, Trinajstic, Young & Senden, 2008" authorityName="Long, Trinajstic, Young & Senden" authorityYear="2008" box="[1312,1421,1813,1834]" class="Placodermi" family="Ptyctodontidae" genus="Materpiscis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ptyctodontida" pageId="0" pageNumber="650" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1312,1421,1813,1834]" italics="true" pageId="0" pageNumber="650">Materpiscis</emphasis>
|
||
</taxonomicName>
|
||
gen. nov. meet at a distinct medially directed symphysis (sym,
|
||
<figureCitation box="[1392,1449,1840,1863]" captionStart="Figure 2" captionStartId="1.[1056,1114,1439,1458]" captionTargetBox="[113,920,1444,1995]" captionTargetId="figure@1.[111,922,1439,1997]" captionTargetPageId="1" captionText="Figure 2 | Holotype of Materpiscis attenboroughi gen. et sp. nov. (WAM 07.12.1). a – e, SEM images of umbilical cord. a, Umbilicus section. b, Image showing a vascular canal (v). c, image showing pit (p). d, Image showing possible appendiculae (app) and pit (p). e, Image showing outer epithelium surface (ep) and stalk wall. f, High-resolution computer tomogram (HRCT) showing embryonic bones. g, HRCT tomogram showing supragnathals in occlusion. h, HRCT showing umbilicus (white arrowheads). Abbreviations as for Fig. 1 except for: ep, epithelium; sym, symphysis." httpUri="https://zenodo.org/record/270101/files/figure.png" pageId="0" pageNumber="650">
|
||
Fig.
|
||
<quantity box="[1436,1462,1840,1862]" metricMagnitude="-3" metricUnit="kg" metricValue="2.0" pageId="0" pageNumber="650" unit="g" value="2.0">2g</quantity>
|
||
</figureCitation>
|
||
), lending increased structural support for biting ability immediately after birth.
|
||
</paragraph>
|
||
<footnote pageId="0" pageNumber="650">
|
||
<paragraph blockId="0.[93,1475,1939,2002]" pageId="0" pageNumber="650">1Museum Victoria,Melbourne, PO Box 666,Melbourne 3001, Australia.2Research School of Earth Sciences, The Australian National University, Canberra 0 200, Australia. 3School of Geosciences,Monash University,Clayton,Victoria 3800, Australia.4School of Earth and Geographical Sciences,The University of Western Australia,Perth 6009, Western Australia, Australia. 5Department of Applied Mathematics, Research School of Physical Sciences and Engineering, The Australian National University, Canberra 0 200, Australia.</paragraph>
|
||
</footnote>
|
||
<caption httpUri="https://zenodo.org/record/270099/files/figure.png" pageId="1" pageNumber="651" targetBox="[273,774,416,1105]" targetPageId="1">
|
||
<paragraph blockId="1.[112,783,1135,1282]" pageId="1" pageNumber="651">
|
||
<caption httpUri="https://zenodo.org/record/270100/files/figure.png" pageId="1" pageNumber="651" targetBox="[273,774,416,1105]" targetPageId="1">
|
||
<emphasis bold="true" box="[112,185,1137,1156]" pageId="1" pageNumber="651">Figure 1</emphasis>
|
||
<emphasis bold="true" box="[191,195,1135,1157]" pageId="1" pageNumber="651">|</emphasis>
|
||
Holotype of
|
||
<taxonomicName authority="Long, Trinajstic, Young & Senden, 2008" authorityName="Long, Trinajstic, Young & Senden" authorityYear="2008" box="[314,549,1137,1156]" class="Placodermi" family="Ptyctodontidae" genus="Materpiscis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ptyctodontida" pageId="1" pageNumber="651" phylum="Chordata" rank="species" species="attenboroughi">
|
||
<emphasis bold="true" box="[314,549,1137,1156]" italics="true" pageId="1" pageNumber="651">Materpiscis attenboroughi</emphasis>
|
||
</taxonomicName>
|
||
gen. et sp. nov. (WAM 07.12.1). a, Body with embryo. The boxed area is magnified in panel
|
||
</caption>
|
||
<emphasis bold="true" pageId="1" pageNumber="651">c. b, Embryo at early preparation. c, d, Umbilical cord and embryo at later preparation. e, Adult skull roof in dorsal view. f, Adult left marginal.</emphasis>
|
||
<emphasis bold="true" pageId="1" pageNumber="651">
|
||
h, Dermal armour restored in lateral view. i, Left adult tooth plates in mesial view. g,
|
||
<taxonomicName authority="Long, 1997" authorityName="Long" authorityYear="1997" box="[185,415,1263,1282]" class="Placodermi" family="Ptyctodontidae" genus="Austroptyctodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ptyctodontida" pageId="1" pageNumber="651" phylum="Chordata" rank="species" species="gardineri">
|
||
<emphasis bold="true" box="[185,415,1263,1282]" italics="true" pageId="1" pageNumber="651">Austroptyctodus gardineri</emphasis>
|
||
</taxonomicName>
|
||
(WAM 86.9.662) showing embryos.
|
||
</emphasis>
|
||
</paragraph>
|
||
<paragraph blockId="1.[821,1493,1137,1257]" pageId="1" pageNumber="651">
|
||
<emphasis bold="true" pageId="1" pageNumber="651">ADL, anterior dorsolateral plate; AL, anterior lateral plate; Ce, central; df.bas, dorsal fin basal plate of adult; haem, haemal arch; Ifg, inferognathal; MD, median dorsal plate; Mg, marginal; neur, neural arch; Nu, nuchal; PNu, paranuchal plate; PRO, preorbital; PTO, postorbital;</emphasis>
|
||
<emphasis bold="true" box="[821,1395,1238,1257]" pageId="1" pageNumber="651">Sgn, supragnathal; SM, submarginal plate; umb, umbilical cord.</emphasis>
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="1.[112,784,1323,1402]" lastBlockId="1.[821,1493,1323,1402]" pageId="1" pageNumber="651">
|
||
A thin tube of mineralized white material with slight helical twisting (umb,
|
||
<figureCitation box="[217,269,1351,1374]" captionStart="Figure 1" captionStartId="1.[112,170,1137,1156]" captionTargetBox="[273,774,416,1105]" captionTargetId="figure@1.[271,677,414,838]" captionTargetPageId="1" captionText="Figure 1 | Holotype of Materpiscis attenboroughi gen. et sp. nov. (WAM 07.12.1). a, Body with embryo. The boxed area is magnified in panel c. b, Embryo at early preparation. c, d, Umbilical cord and embryo at later preparation. e, Adult skull roof in dorsal view. f, Adult left marginal. h, Dermal armour restored in lateral view. i, Left adult tooth plates in mesial view. g, Austroptyctodus gardineri (WAM 86.9.662) showing embryos. ADL, anterior dorsolateral plate; AL, anterior lateral plate; Ce, central; df. bas, dorsal fin basal plate of adult; haem, haemal arch; Ifg, inferognathal; MD, median dorsal plate; Mg, marginal; neur, neural arch; Nu, nuchal; PNu, paranuchal plate; PRO, preorbital; PTO, postorbital; Sgn, supragnathal; SM, submarginal plate; umb, umbilical cord." httpUri="https://zenodo.org/record/270100/files/figure.png" pageId="1" pageNumber="651">
|
||
Fig.
|
||
<number box="[259,269,1352,1373]" pageId="1" pageNumber="651" value="1.0">1</number>
|
||
</figureCitation>
|
||
c, d) connects the
|
||
<taxonomicName authority="Long, Trinajstic, Young & Senden, 2008" authorityName="Long, Trinajstic, Young & Senden" authorityYear="2008" box="[452,561,1352,1373]" class="Placodermi" family="Ptyctodontidae" genus="Materpiscis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ptyctodontida" pageId="1" pageNumber="651" phylum="Chordata" rank="genus" status="gen. nov.">
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||
<emphasis box="[452,561,1352,1373]" italics="true" pageId="1" pageNumber="651">Materpiscis</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel box="[568,659,1352,1373]" pageId="1" pageNumber="651" rank="genus">gen. nov.</taxonomicNameLabel>
|
||
embryo to a region within the upper body cavity of the female, marked by an ovoid mass of coarsely crystalline calcite, possibly the position of the yolk sac (
|
||
<figureCitation box="[953,1009,1351,1374]" captionStart="Figure 1" captionStartId="1.[112,170,1137,1156]" captionTargetBox="[273,774,416,1105]" captionTargetId="figure@1.[271,677,414,838]" captionTargetPageId="1" captionText="Figure 1 | Holotype of Materpiscis attenboroughi gen. et sp. nov. (WAM 07.12.1). a, Body with embryo. The boxed area is magnified in panel c. b, Embryo at early preparation. c, d, Umbilical cord and embryo at later preparation. e, Adult skull roof in dorsal view. f, Adult left marginal. h, Dermal armour restored in lateral view. i, Left adult tooth plates in mesial view. g, Austroptyctodus gardineri (WAM 86.9.662) showing embryos. ADL, anterior dorsolateral plate; AL, anterior lateral plate; Ce, central; df. bas, dorsal fin basal plate of adult; haem, haemal arch; Ifg, inferognathal; MD, median dorsal plate; Mg, marginal; neur, neural arch; Nu, nuchal; PNu, paranuchal plate; PRO, preorbital; PTO, postorbital; Sgn, supragnathal; SM, submarginal plate; umb, umbilical cord." httpUri="https://zenodo.org/record/270100/files/figure.png" pageId="1" pageNumber="651">
|
||
Fig.
|
||
<number box="[999,1009,1352,1373]" pageId="1" pageNumber="651" value="1.0">1</number>
|
||
</figureCitation>
|
||
c, d). In modern sharks, the umbilical cord wall comprises multiple small vessels, capillaries and a dense spongy
|
||
</paragraph>
|
||
<caption pageId="1" pageNumber="651">
|
||
<paragraph blockId="1.[1056,1491,1437,1761]" pageId="1" pageNumber="651">
|
||
<caption httpUri="https://zenodo.org/record/270101/files/figure.png" pageId="1" pageNumber="651" targetBox="[113,920,1444,1995]" targetPageId="1">
|
||
<emphasis bold="true" box="[1056,1132,1439,1458]" pageId="1" pageNumber="651">Figure 2</emphasis>
|
||
<emphasis bold="true" box="[1138,1142,1437,1459]" pageId="1" pageNumber="651">|</emphasis>
|
||
Holotype of
|
||
<taxonomicName authority="Long, Trinajstic, Young & Senden, 2008" authorityName="Long, Trinajstic, Young & Senden" authorityYear="2008" box="[1257,1491,1439,1458]" class="Placodermi" family="Ptyctodontidae" genus="Materpiscis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ptyctodontida" pageId="1" pageNumber="651" phylum="Chordata" rank="species" species="attenboroughi">
|
||
<emphasis bold="true" box="[1257,1491,1439,1458]" italics="true" pageId="1" pageNumber="651">
|
||
<taxonomicName authority="Long, Trinajstic, Young & Senden, 2008" authorityName="Long, Trinajstic, Young & Senden" authorityYear="2008" box="[1257,1360,1439,1458]" class="Placodermi" family="Ptyctodontidae" genus="Materpiscis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ptyctodontida" pageId="1" pageNumber="651" phylum="Chordata" rank="genus">Materpiscis</taxonomicName>
|
||
attenboroughi
|
||
</emphasis>
|
||
</taxonomicName>
|
||
gen. et sp. nov. (WAM 07.12.1). a–e, SEM images of umbilical cord. a, Umbilicus section. b, Image showing a vascular canal (v). c, image showing pit (p). d, Image showing possible
|
||
</caption>
|
||
<emphasis bold="true" pageId="1" pageNumber="651">appendiculae (app) and pit (p). e, Image showing outer epithelium surface (ep) and stalk wall. f, High-resolution computer tomogram (HRCT) showing embryonic bones. g, HRCT tomogram showing supragnathals in occlusion. h, HRCT showing umbilicus (white arrowheads).</emphasis>
|
||
<emphasis bold="true" box="[1056,1425,1717,1736]" pageId="1" pageNumber="651">
|
||
Abbreviations as for
|
||
<figureCitation box="[1243,1293,1717,1736]" captionStart="Figure 1" captionStartId="1.[112,170,1137,1156]" captionTargetBox="[273,774,416,1105]" captionTargetId="figure@1.[271,677,414,838]" captionTargetPageId="1" captionText="Figure 1 | Holotype of Materpiscis attenboroughi gen. et sp. nov. (WAM 07.12.1). a, Body with embryo. The boxed area is magnified in panel c. b, Embryo at early preparation. c, d, Umbilical cord and embryo at later preparation. e, Adult skull roof in dorsal view. f, Adult left marginal. h, Dermal armour restored in lateral view. i, Left adult tooth plates in mesial view. g, Austroptyctodus gardineri (WAM 86.9.662) showing embryos. ADL, anterior dorsolateral plate; AL, anterior lateral plate; Ce, central; df. bas, dorsal fin basal plate of adult; haem, haemal arch; Ifg, inferognathal; MD, median dorsal plate; Mg, marginal; neur, neural arch; Nu, nuchal; PNu, paranuchal plate; PRO, preorbital; PTO, postorbital; Sgn, supragnathal; SM, submarginal plate; umb, umbilical cord." httpUri="https://zenodo.org/record/270100/files/figure.png" pageId="1" pageNumber="651">Fig. 1</figureCitation>
|
||
except for: ep,
|
||
</emphasis>
|
||
<emphasis bold="true" box="[1056,1311,1742,1761]" pageId="1" pageNumber="651">epithelium; sym, symphysis.</emphasis>
|
||
</paragraph>
|
||
</caption>
|
||
<caption httpUri="https://zenodo.org/record/270102/files/figure.png" pageId="2" pageNumber="652" targetBox="[122,735,134,777]" targetPageId="2">
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||
<paragraph blockId="2.[93,765,796,868]" pageId="2" pageNumber="652">
|
||
<emphasis bold="true" box="[93,168,798,817]" pageId="2" pageNumber="652">Figure 3</emphasis>
|
||
<emphasis bold="true" box="[175,179,796,818]" pageId="2" pageNumber="652">|</emphasis>
|
||
<taxonomicName authority="Long, Trinajstic, Young & Senden, 2008" authorityName="Long, Trinajstic, Young & Senden" authorityYear="2008" box="[184,288,798,817]" class="Placodermi" family="Ptyctodontidae" genus="Materpiscis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ptyctodontida" pageId="2" pageNumber="652" phylum="Chordata" rank="genus" status="gen. nov.">
|
||
<emphasis bold="true" box="[184,288,798,817]" italics="true" pageId="2" pageNumber="652">Materpiscis</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel box="[291,373,798,817]" pageId="2" pageNumber="652" rank="genus">gen. nov.</taxonomicNameLabel>
|
||
a, Diagram showing position of embryo and yolk sac within the mother. b, Artist’s reconstruction of
|
||
<taxonomicName authority="Long, Trinajstic, Young & Senden, 2008" authorityName="Long, Trinajstic, Young & Senden" authorityYear="2008" box="[581,682,823,842]" class="Placodermi" family="Ptyctodontidae" genus="Materpiscis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ptyctodontida" pageId="2" pageNumber="652" phylum="Chordata" rank="genus" status="gen. nov.">
|
||
<emphasis bold="true" box="[581,682,823,842]" italics="true" pageId="2" pageNumber="652">Materpiscis</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel box="[685,765,824,843]" pageId="2" pageNumber="652" rank="genus">gen. nov.</taxonomicNameLabel>
|
||
giving birth (by B. Choo).
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="2.[93,766,906,1489]" pageId="2" pageNumber="652">
|
||
connective tissue
|
||
<bibRefCitation author="Wourms, J." box="[260,275,906,919]" pageId="2" pageNumber="652" refString="15. Wourms, J. Maximization of evolutionary trends for placental viviparity in the spadenose shark, Scoliodon laticaudus. Env. Biol. Fish. 38, 269–294 (1993)." type="journal article" year="1993">
|
||
<number box="[260,275,906,919]" pageId="2" pageNumber="652" value="15.0">15</number>
|
||
</bibRefCitation>
|
||
. The fossilized tube has similar external shape and position as the umbilical cord in chondrichthyans. High-resolution scanning electron microscopy
|
||
<botanicCollectionCodeGroup box="[402,468,964,986]" pageId="2" pageNumber="652">
|
||
(
|
||
<collectionCode box="[408,461,964,986]" pageId="2" pageNumber="652">SEM</collectionCode>
|
||
)
|
||
</botanicCollectionCodeGroup>
|
||
imaging and high-resolution computer tomography scanning of this specimen revealed that the cord loops underneath the adult dorsal fin basal plate (
|
||
<figureCitation box="[649,707,1019,1042]" captionStart="Figure 2" captionStartId="1.[1056,1114,1439,1458]" captionTargetBox="[113,920,1444,1995]" captionTargetId="figure@1.[111,922,1439,1997]" captionTargetPageId="1" captionText="Figure 2 | Holotype of Materpiscis attenboroughi gen. et sp. nov. (WAM 07.12.1). a – e, SEM images of umbilical cord. a, Umbilicus section. b, Image showing a vascular canal (v). c, image showing pit (p). d, Image showing possible appendiculae (app) and pit (p). e, Image showing outer epithelium surface (ep) and stalk wall. f, High-resolution computer tomogram (HRCT) showing embryonic bones. g, HRCT tomogram showing supragnathals in occlusion. h, HRCT showing umbilicus (white arrowheads). Abbreviations as for Fig. 1 except for: ep, epithelium; sym, symphysis." httpUri="https://zenodo.org/record/270101/files/figure.png" pageId="2" pageNumber="652">
|
||
Fig.
|
||
<number box="[695,707,1019,1042]" pageId="2" pageNumber="652" value="2.0">2</number>
|
||
</figureCitation>
|
||
h and Supplementary Information). Its surface is covered with a thin layer of epithelium (
|
||
<figureCitation box="[237,293,1075,1098]" captionStart="Figure 2" captionStartId="1.[1056,1114,1439,1458]" captionTargetBox="[113,920,1444,1995]" captionTargetId="figure@1.[111,922,1439,1997]" captionTargetPageId="1" captionText="Figure 2 | Holotype of Materpiscis attenboroughi gen. et sp. nov. (WAM 07.12.1). a – e, SEM images of umbilical cord. a, Umbilicus section. b, Image showing a vascular canal (v). c, image showing pit (p). d, Image showing possible appendiculae (app) and pit (p). e, Image showing outer epithelium surface (ep) and stalk wall. f, High-resolution computer tomogram (HRCT) showing embryonic bones. g, HRCT tomogram showing supragnathals in occlusion. h, HRCT showing umbilicus (white arrowheads). Abbreviations as for Fig. 1 except for: ep, epithelium; sym, symphysis." httpUri="https://zenodo.org/record/270101/files/figure.png" pageId="2" pageNumber="652">
|
||
Fig.
|
||
<number box="[280,293,1076,1098]" pageId="2" pageNumber="652" value="2.0">2</number>
|
||
</figureCitation>
|
||
e) and has a series of pits of varying sizes (
|
||
<figureCitation box="[694,750,1075,1098]" captionStart="Figure 2" captionStartId="1.[1056,1114,1439,1458]" captionTargetBox="[113,920,1444,1995]" captionTargetId="figure@1.[111,922,1439,1997]" captionTargetPageId="1" captionText="Figure 2 | Holotype of Materpiscis attenboroughi gen. et sp. nov. (WAM 07.12.1). a – e, SEM images of umbilical cord. a, Umbilicus section. b, Image showing a vascular canal (v). c, image showing pit (p). d, Image showing possible appendiculae (app) and pit (p). e, Image showing outer epithelium surface (ep) and stalk wall. f, High-resolution computer tomogram (HRCT) showing embryonic bones. g, HRCT tomogram showing supragnathals in occlusion. h, HRCT showing umbilicus (white arrowheads). Abbreviations as for Fig. 1 except for: ep, epithelium; sym, symphysis." httpUri="https://zenodo.org/record/270101/files/figure.png" pageId="2" pageNumber="652">
|
||
Fig.
|
||
<number box="[737,750,1076,1097]" pageId="2" pageNumber="652" value="2.0">2</number>
|
||
</figureCitation>
|
||
c, d), with the wall of the umbilical cord made up of a series of vessels and a dense matrix of tissue. The path of one major vessel could be determined running through the cord (
|
||
<figureCitation box="[483,542,1159,1182]" captionStart="Figure 2" captionStartId="1.[1056,1114,1439,1458]" captionTargetBox="[113,920,1444,1995]" captionTargetId="figure@1.[111,922,1439,1997]" captionTargetPageId="1" captionText="Figure 2 | Holotype of Materpiscis attenboroughi gen. et sp. nov. (WAM 07.12.1). a – e, SEM images of umbilical cord. a, Umbilicus section. b, Image showing a vascular canal (v). c, image showing pit (p). d, Image showing possible appendiculae (app) and pit (p). e, Image showing outer epithelium surface (ep) and stalk wall. f, High-resolution computer tomogram (HRCT) showing embryonic bones. g, HRCT tomogram showing supragnathals in occlusion. h, HRCT showing umbilicus (white arrowheads). Abbreviations as for Fig. 1 except for: ep, epithelium; sym, symphysis." httpUri="https://zenodo.org/record/270101/files/figure.png" pageId="2" pageNumber="652">
|
||
Fig.
|
||
<number box="[528,542,1159,1182]" pageId="2" pageNumber="652" value="2.0">2</number>
|
||
</figureCitation>
|
||
b). Branching off from the umbilical cord are the remains of filamentous processes equivalent to the appendiculae in extant sharks (app,
|
||
<figureCitation box="[560,618,1215,1238]" captionStart="Figure 2" captionStartId="1.[1056,1114,1439,1458]" captionTargetBox="[113,920,1444,1995]" captionTargetId="figure@1.[111,922,1439,1997]" captionTargetPageId="1" captionText="Figure 2 | Holotype of Materpiscis attenboroughi gen. et sp. nov. (WAM 07.12.1). a – e, SEM images of umbilical cord. a, Umbilicus section. b, Image showing a vascular canal (v). c, image showing pit (p). d, Image showing possible appendiculae (app) and pit (p). e, Image showing outer epithelium surface (ep) and stalk wall. f, High-resolution computer tomogram (HRCT) showing embryonic bones. g, HRCT tomogram showing supragnathals in occlusion. h, HRCT showing umbilicus (white arrowheads). Abbreviations as for Fig. 1 except for: ep, epithelium; sym, symphysis." httpUri="https://zenodo.org/record/270101/files/figure.png" pageId="2" pageNumber="652">
|
||
Fig.
|
||
<number box="[604,618,1215,1238]" pageId="2" pageNumber="652" value="2.0">2</number>
|
||
</figureCitation>
|
||
d). Thus, ptyctodonts apparently evolved beyond the first live-bearing stage (oviviparity, in which the mother retains her eggs inside her body, providing only protection, with the yolk sac providing all nutrition) to matrotrophy. The relative size and position of the embryo and yolk sac in relation to the mother fish is shown in
|
||
<figureCitation box="[597,657,1354,1377]" captionStart="Figure 3" captionStartId="2.[93,151,798,817]" captionTargetBox="[122,735,134,777]" captionTargetId="figure@2.[122,736,405,777]" captionTargetPageId="2" captionText="Figure 3 | Materpiscis gen. nov. a, Diagram showing position of embryo and yolk sac within the mother. b, Artist’s reconstruction of Materpiscis gen. nov. giving birth (by B. Choo)." httpUri="https://zenodo.org/record/270102/files/figure.png" pageId="2" pageNumber="652">
|
||
Fig.
|
||
<number box="[645,657,1355,1376]" pageId="2" pageNumber="652" value="3.0">3</number>
|
||
</figureCitation>
|
||
a, with an attempted restoration of the fish giving birth shown in
|
||
<figureCitation box="[632,686,1382,1405]" captionStart="Figure 3" captionStartId="2.[93,151,798,817]" captionTargetBox="[122,735,134,777]" captionTargetId="figure@2.[122,736,405,777]" captionTargetPageId="2" captionText="Figure 3 | Materpiscis gen. nov. a, Diagram showing position of embryo and yolk sac within the mother. b, Artist’s reconstruction of Materpiscis gen. nov. giving birth (by B. Choo)." httpUri="https://zenodo.org/record/270102/files/figure.png" pageId="2" pageNumber="652">
|
||
Fig.
|
||
<number box="[674,686,1382,1405]" pageId="2" pageNumber="652" value="3.0">3</number>
|
||
</figureCitation>
|
||
b, based on the observation that several viviparous sharks give birth tail-first to their young. Further discussion of the significance of viviparity and evolution can be found in the Supplementary Information.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |