296 lines
36 KiB
XML
296 lines
36 KiB
XML
<document ID-DOI="10.1098/rsbl.2020.0735" ID-GBIF-Dataset="6f2deb98-cbd4-4ef1-b0c0-7c65e774bd74" ID-PMC="PMC7775975" ID-PubMed="33321065" ID-Zenodo-Dep="4383218" checkinTime="1608588430264" checkinUser="plazi" docAuthor="Zaher, Hussam & Smith, Krister T." docDate="2020" docId="038587B1FFDCFFE84D75EA72FB54F7FB" docLanguage="en" docName="biologyLetters.2020.0735.pdf" docOrigin="Biology Letters 16 (12)" docSource="http://dx.doi.org/10.1098/rsbl.2020.0735" docTitle="Messelopython freyi Zaher & Smith 2020, gen. et sp. nov." docType="treatment" docVersion="9" lastPageNumber="4" masterDocId="FFBCFFC9FFDDFFEB4D07EC00FFEEFFC7" masterDocTitle="Pythons in the Eocene of Europe reveal a much older divergence of the group in sympatry with boas" masterLastPageNumber="6" masterPageNumber="1" pageNumber="2" updateTime="1668126595692" 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>Pythons in the Eocene of Europe reveal a much older divergence of the group in sympatry with boas</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>Zaher, Hussam</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>Smith, Krister T.</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>Biology Letters</mods:title>
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</mods:titleInfo>
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<mods:part>
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<mods:date>2020</mods:date>
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<mods:detail type="pubDate">
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<mods:number>2020-12-16</mods:number>
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</mods:detail>
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<mods:detail type="volume">
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<mods:number>16</mods:number>
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</mods:detail>
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<mods:detail type="issue">
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<mods:number>12</mods:number>
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</mods:detail>
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<mods:extent unit="page">
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<mods:start>1</mods:start>
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<mods:end>6</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:location>
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<mods:url>http://dx.doi.org/10.1098/rsbl.2020.0735</mods:url>
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</mods:location>
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<mods:classification>journal article</mods:classification>
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<mods:identifier type="DOI">10.1098/rsbl.2020.0735</mods:identifier>
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<mods:identifier type="GBIF-Dataset">6f2deb98-cbd4-4ef1-b0c0-7c65e774bd74</mods:identifier>
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<mods:identifier type="PMC">PMC7775975</mods:identifier>
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<mods:identifier type="PubMed">33321065</mods:identifier>
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<mods:identifier type="Zenodo-Dep">4383218</mods:identifier>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.6485876" ID-GBIF-Taxon="176232740" ID-Zenodo-Dep="6485876" LSID="urn:lsid:zoobank.org:act:84AF3F8E-1B68-4807-A5A3-31D95A9ADE84" httpUri="http://treatment.plazi.org/id/038587B1FFDCFFE84D75EA72FB54F7FB" lastPageId="3" lastPageNumber="4" pageId="1" pageNumber="2">
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<subSubSection pageId="1" pageNumber="2" type="nomenclature">
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<paragraph blockId="1.[113,756,1534,2120]" box="[114,480,1650,1671]" pageId="1" pageNumber="2">
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<taxonomicName authority="Zaher & Smith, 2020" authorityName="Zaher & Smith" authorityYear="2020" box="[114,308,1650,1671]" class="Reptilia" family="Pythonidae" genus="Messelopython" kingdom="Animalia" order="Squamata" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="freyi" status="gen. et sp. nov.">
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<emphasis box="[114,308,1650,1671]" italics="true" pageId="1" pageNumber="2">Messelopython freyi</emphasis>
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</taxonomicName>
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<taxonomicNameLabel box="[316,480,1650,1671]" pageId="1" pageNumber="2" rank="species">gen. et sp. nov.</taxonomicNameLabel>
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</paragraph>
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<paragraph blockId="1.[113,756,1534,2120]" pageId="1" pageNumber="2">
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<uri pageId="1" pageNumber="2">
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urn:lsid:zoobank.org:act:
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<uuid pageId="1" pageNumber="2">E8C2F2DA-1434-4580-8D30A5EDA71A2CAC</uuid>
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<uri pageId="1" pageNumber="2">
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urn:lsid:zoobank.org:act:
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</subSubSection>
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<subSubSection pageId="1" pageNumber="2" type="materials_examined">
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<paragraph blockId="1.[113,756,1534,2120]" pageId="1" pageNumber="2">
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<materialsCitation ID-GBIF-Occurrence="3001070305" pageId="1" pageNumber="2" specimenCode="SMNK-PAL 461" specimenCount="1" typeStatus="holotype">
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<materialsCitation ID-GBIF-Occurrence="3001070306" pageId="1" pageNumber="2" specimenCode="SMNK-PAL 461" specimenCount="1" typeStatus="holotype">
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<emphasis bold="true" box="[151,252,1810,1831]" pageId="1" pageNumber="2">
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<typeStatus box="[151,252,1810,1831]" pageId="1" pageNumber="2">Holotype</typeStatus>
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</emphasis>
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:
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<specimenCode box="[275,455,1810,1832]" collectionCode="SMNK-PAL" pageId="1" pageNumber="0">SMNK-PAL 461</specimenCode>
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, nearly complete skeleton with partial skull
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</materialsCitation>
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(
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<figureCitation box="[312,429,1842,1864]" captionStart="Figure 1" captionStartId="2.[113,171,1572,1596]" captionTargetBox="[151,1426,113,1539]" captionTargetId="figure-156@2.[151,1437,113,1539]" captionTargetPageId="2" captionText="Figure 1. Hypodigm of Messelopython freyi gen. et sp. nov. (a) and (b) Photograph and drawing of skull of holotype, SMNK-PAL 461a. (c) Photograph of holotype. (d) and (e) Photograph and drawing of skull of paratype, SMF-ME 2784a. (f) Drawing of left splenial and angular in medial view in counterpart of paratype. (g) Drawing of right ectopterygoid in dorsal view in main part of paratype. (h) and (i) Photograph and drawing of skull of paratype, HLMD-Be 165. Abbreviations: an, angular; boc, basioccipital; bpt-pr, basipterygoid process; bs, basisphenoid; d, dentary; ec, ectopterygoid; fr, frontal; mx, maxilla; n, nasal; ot, otooccipital; pa, parietal; par, prearticular part of compound bone; pl, palatine; pf, postfrontal; prf, prefrontal; pro, prootic; pt, pterygoid; q, quadrate; sa, surangular part of compound bone; smx, septomaxilla; so, supraorbital; soc, supraoccipital; spl, splenial; st, supratemporal." figureDoi="http://doi.org/10.5281/zenodo.4383220" httpUri="https://zenodo.org/record/4383220/files/figure.png" pageId="1" pageNumber="2">
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figure 1
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<emphasis box="[397,429,1843,1864]" italics="true" pageId="1" pageNumber="2">a–c</emphasis>
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</figureCitation>
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)
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</materialsCitation>
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</paragraph>
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<paragraph blockId="1.[113,756,1534,2120]" pageId="1" pageNumber="2">
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<materialsCitation ID-GBIF-Occurrence="3001070304" pageId="1" pageNumber="2" specimenCode="SMF-ME 710" specimenCount="1" specimenCount-female="1" typeStatus="paratype">
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<emphasis bold="true" box="[151,255,1874,1895]" pageId="1" pageNumber="2">
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<typeStatus box="[151,255,1874,1895]" pageId="1" pageNumber="2">Paratypes</typeStatus>
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</emphasis>
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:
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<specimenCode box="[269,407,1874,1895]" pageId="1" pageNumber="2">SMF-ME 710</specimenCode>
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, nearly complete skeleton with a badly crushed skull (
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<figureCitation box="[358,455,1906,1928]" captionStart="Figure 1" captionStartId="2.[113,171,1572,1596]" captionTargetBox="[151,1426,113,1539]" captionTargetId="figure-156@2.[151,1437,113,1539]" captionTargetPageId="2" captionText="Figure 1. Hypodigm of Messelopython freyi gen. et sp. nov. (a) and (b) Photograph and drawing of skull of holotype, SMNK-PAL 461a. (c) Photograph of holotype. (d) and (e) Photograph and drawing of skull of paratype, SMF-ME 2784a. (f) Drawing of left splenial and angular in medial view in counterpart of paratype. (g) Drawing of right ectopterygoid in dorsal view in main part of paratype. (h) and (i) Photograph and drawing of skull of paratype, HLMD-Be 165. Abbreviations: an, angular; boc, basioccipital; bpt-pr, basipterygoid process; bs, basisphenoid; d, dentary; ec, ectopterygoid; fr, frontal; mx, maxilla; n, nasal; ot, otooccipital; pa, parietal; par, prearticular part of compound bone; pl, palatine; pf, postfrontal; prf, prefrontal; pro, prootic; pt, pterygoid; q, quadrate; sa, surangular part of compound bone; smx, septomaxilla; so, supraorbital; soc, supraoccipital; spl, splenial; st, supratemporal." figureDoi="http://doi.org/10.5281/zenodo.4383220" httpUri="https://zenodo.org/record/4383220/files/figure.png" pageId="1" pageNumber="2">
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figure
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<specimenCount box="[437,455,1906,1927]" pageId="1" pageNumber="2" type="female">
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1
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<emphasis box="[449,455,1906,1927]" italics="true" pageId="1" pageNumber="2">f</emphasis>
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</specimenCount>
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</figureCitation>
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);
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</materialsCitation>
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<materialsCitation ID-GBIF-Occurrence="3001070301" pageId="1" pageNumber="2" specimenCode="SMF-ME 2784" specimenCount="1">
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<specimenCode box="[487,645,1906,1927]" pageId="1" pageNumber="2">SMF-ME 2784</specimenCode>
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, a nearly complete skeleton with a skull (
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<figureCitation box="[449,578,1938,1960]" captionStart="Figure 1" captionStartId="2.[113,171,1572,1596]" captionTargetBox="[151,1426,113,1539]" captionTargetId="figure-156@2.[151,1437,113,1539]" captionTargetPageId="2" captionText="Figure 1. Hypodigm of Messelopython freyi gen. et sp. nov. (a) and (b) Photograph and drawing of skull of holotype, SMNK-PAL 461a. (c) Photograph of holotype. (d) and (e) Photograph and drawing of skull of paratype, SMF-ME 2784a. (f) Drawing of left splenial and angular in medial view in counterpart of paratype. (g) Drawing of right ectopterygoid in dorsal view in main part of paratype. (h) and (i) Photograph and drawing of skull of paratype, HLMD-Be 165. Abbreviations: an, angular; boc, basioccipital; bpt-pr, basipterygoid process; bs, basisphenoid; d, dentary; ec, ectopterygoid; fr, frontal; mx, maxilla; n, nasal; ot, otooccipital; pa, parietal; par, prearticular part of compound bone; pl, palatine; pf, postfrontal; prf, prefrontal; pro, prootic; pt, pterygoid; q, quadrate; sa, surangular part of compound bone; smx, septomaxilla; so, supraorbital; soc, supraoccipital; spl, splenial; st, supratemporal." figureDoi="http://doi.org/10.5281/zenodo.4383220" httpUri="https://zenodo.org/record/4383220/files/figure.png" pageId="1" pageNumber="2">
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figure 1
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<emphasis box="[532,578,1938,1959]" italics="true" pageId="1" pageNumber="2">d,e,g</emphasis>
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</figureCitation>
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);
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</materialsCitation>
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<materialsCitation ID-GBIF-Occurrence="3001070302" pageId="1" pageNumber="2" specimenCode="HLMD-Be 165" specimenCount="1">
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<specimenCode box="[598,752,1938,1960]" pageId="1" pageNumber="2">HLMD-Be 165</specimenCode>
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, a nearly complete skeleton with a partial skull (
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<figureCitation box="[614,721,1970,1992]" captionStart="Figure 1" captionStartId="2.[113,171,1572,1596]" captionTargetBox="[151,1426,113,1539]" captionTargetId="figure-156@2.[151,1437,113,1539]" captionTargetPageId="2" captionText="Figure 1. Hypodigm of Messelopython freyi gen. et sp. nov. (a) and (b) Photograph and drawing of skull of holotype, SMNK-PAL 461a. (c) Photograph of holotype. (d) and (e) Photograph and drawing of skull of paratype, SMF-ME 2784a. (f) Drawing of left splenial and angular in medial view in counterpart of paratype. (g) Drawing of right ectopterygoid in dorsal view in main part of paratype. (h) and (i) Photograph and drawing of skull of paratype, HLMD-Be 165. Abbreviations: an, angular; boc, basioccipital; bpt-pr, basipterygoid process; bs, basisphenoid; d, dentary; ec, ectopterygoid; fr, frontal; mx, maxilla; n, nasal; ot, otooccipital; pa, parietal; par, prearticular part of compound bone; pl, palatine; pf, postfrontal; prf, prefrontal; pro, prootic; pt, pterygoid; q, quadrate; sa, surangular part of compound bone; smx, septomaxilla; so, supraorbital; soc, supraoccipital; spl, splenial; st, supratemporal." figureDoi="http://doi.org/10.5281/zenodo.4383220" httpUri="https://zenodo.org/record/4383220/files/figure.png" pageId="1" pageNumber="2">
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||
<figureCitation box="[614,697,1970,1992]" captionStart="Figure 1" captionStartId="2.[113,171,1572,1596]" captionTargetBox="[151,1426,113,1539]" captionTargetId="figure-156@2.[151,1437,113,1539]" captionTargetPageId="2" captionText="Figure 1. Hypodigm of Messelopython freyi gen. et sp. nov. (a) and (b) Photograph and drawing of skull of holotype, SMNK-PAL 461a. (c) Photograph of holotype. (d) and (e) Photograph and drawing of skull of paratype, SMF-ME 2784a. (f) Drawing of left splenial and angular in medial view in counterpart of paratype. (g) Drawing of right ectopterygoid in dorsal view in main part of paratype. (h) and (i) Photograph and drawing of skull of paratype, HLMD-Be 165. Abbreviations: an, angular; boc, basioccipital; bpt-pr, basipterygoid process; bs, basisphenoid; d, dentary; ec, ectopterygoid; fr, frontal; mx, maxilla; n, nasal; ot, otooccipital; pa, parietal; par, prearticular part of compound bone; pl, palatine; pf, postfrontal; prf, prefrontal; pro, prootic; pt, pterygoid; q, quadrate; sa, surangular part of compound bone; smx, septomaxilla; so, supraorbital; soc, supraoccipital; spl, splenial; st, supratemporal." figureDoi="http://doi.org/10.5281/zenodo.4383220" httpUri="https://zenodo.org/record/4383220/files/figure.png" pageId="1" pageNumber="2">figure 1</figureCitation>
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<emphasis box="[697,721,1970,1991]" italics="true" pageId="1" pageNumber="2">h,i</emphasis>
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</figureCitation>
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).
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</materialsCitation>
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</paragraph>
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<paragraph blockId="1.[113,756,1534,2120]" box="[151,555,2002,2024]" pageId="1" pageNumber="2">
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<emphasis bold="true" box="[151,357,2002,2024]" pageId="1" pageNumber="2">Referred specimen</emphasis>
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: HLMD-Me 10583
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</paragraph>
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</subSubSection>
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<subSubSection pageId="1" pageNumber="2" type="diagnosis">
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<paragraph blockId="1.[113,756,1534,2120]" lastBlockId="1.[831,1475,114,456]" pageId="1" pageNumber="2">
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<emphasis bold="true" box="[151,260,2034,2056]" pageId="1" pageNumber="2">Diagnosis</emphasis>
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: Medium-sized pythonoid (electronic supplementary material, S6) sharing with
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<taxonomicName box="[488,608,2066,2088]" class="Reptilia" family="Pythonidae" kingdom="Animalia" order="Squamata" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Pythonidae</taxonomicName>
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the following synapomorphies: a toothed premaxilla with no midline diastema; palatine foramen present in palatine; and mid-sagittal crests on parietal and basisphenoid. Autapomorphies: 6 premaxillary teeth; sigmoid lateral margin of maxilla; large, crescentic supraorbital bone nearly equal in length to the frontal; ectopterygoid overlap of pterygoid reduced.
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</paragraph>
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</subSubSection>
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<subSubSection pageId="1" pageNumber="2" type="materials_examined">
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<paragraph blockId="1.[831,1475,114,456]" pageId="1" pageNumber="2">
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<materialsCitation ID-GBIF-Occurrence="3001070303" country="Germany" location="Messel Pit," pageId="1" pageNumber="2" specimenCount="1" typeStatus="holotype">
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<emphasis bold="true" box="[869,1151,274,296]" pageId="1" pageNumber="2">
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<typeStatus box="[869,924,274,295]" pageId="1" pageNumber="2">Type</typeStatus>
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locality and horizon
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</emphasis>
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:
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<location LSID="urn:lsid:plazi:treatment:038587B1FFDCFFE84D75EA72FB54F7FB:8EF3607CFFDCFFEA498AED12FB11FEEF" box="[1165,1279,274,296]" country="Germany" name="Messel Pit," pageId="1" pageNumber="2">Messel Pit,</location>
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<collectingCountry box="[1286,1387,274,295]" name="Germany" pageId="1" pageNumber="2">Germany</collectingCountry>
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;
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<geologicalTimeScale pageId="1" pageNumber="2">Middle Messel Formation, latest early to earliest middle Eocene</geologicalTimeScale>
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</materialsCitation>
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[
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<bibRefCitation author="Lenz OK & Wilde V & Mertz DF & Riegel W." box="[1435,1464,305,327]" journalOrPublisher="Int. J. Earth Sci." pageId="1" pageNumber="2" pagination="873 - 889" part="104" refId="ref5002" refString="35. Lenz OK, Wilde V, Mertz DF, Riegel W. 2015 New palynology-based astronomical and revised 40 Ar / 39 Ar ages for the Eocene maar lake of Messel (Germany). Int. J. Earth Sci. 104, 873 - 889. (doi: 10. 1007 / s 00531 - 014 - 1126 - 2)" title="New palynology-based astronomical and revised 40 Ar / 39 Ar ages for the Eocene maar lake of Messel (Germany)" type="journal article" year="2015">35</bibRefCitation>
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].
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</paragraph>
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</subSubSection>
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<subSubSection pageId="1" pageNumber="2" type="etymology">
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<paragraph blockId="1.[831,1475,114,456]" pageId="1" pageNumber="2">
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<emphasis bold="true" box="[869,986,338,359]" pageId="1" pageNumber="2">Etymology</emphasis>
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: ‘Messel’, after the
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<typeStatus box="[1222,1268,338,359]" pageId="1" pageNumber="2">type</typeStatus>
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locality [
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<bibRefCitation author="Smith KT & Schaal SFK & Habersetzer J." box="[1380,1408,337,359]" journalOrPublisher="Stuttgart, Germany: Schweizerbart" pageId="1" pageNumber="2" refId="ref4567" refString="25. Smith KT, Schaal SFK, Habersetzer J. 2018 Messel: an ancient greenhouse ecosystem, p. 355. Stuttgart, Germany: Schweizerbart." title="Messel: an ancient greenhouse ecosystem" type="book" year="2018">25</bibRefCitation>
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], and ‘python’, after its close living relatives. The specific epithet honours Eberhard ‘Dino’ Frey, for his detailed studies of fossil and extant reptiles.
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</paragraph>
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</subSubSection>
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<subSubSection lastPageId="3" lastPageNumber="4" pageId="1" pageNumber="2" type="discussion">
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||
<paragraph blockId="1.[831,1474,541,2120]" box="[831,961,541,579]" pageId="1" pageNumber="2">
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||
<heading bold="true" box="[831,961,541,579]" fontSize="16" level="1" pageId="1" pageNumber="2" reason="0">
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||
<emphasis bold="true" box="[831,961,541,579]" pageId="1" pageNumber="2">4. Results</emphasis>
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</heading>
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||
</paragraph>
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<paragraph blockId="1.[831,1474,541,2120]" pageId="1" pageNumber="2">
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The premaxilla of
|
||
<taxonomicName authorityName="Zaher & Smith" authorityYear="2020" box="[1030,1109,594,616]" class="Reptilia" family="Pythonidae" genus="Messelopython" kingdom="Animalia" order="Squamata" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="freyi">
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<emphasis box="[1030,1109,594,616]" italics="true" pageId="1" pageNumber="2">M. freyi</emphasis>
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</taxonomicName>
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(
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<figureCitation box="[1126,1209,594,616]" captionStart="Figure 1" captionStartId="2.[113,171,1572,1596]" captionTargetBox="[151,1426,113,1539]" captionTargetId="figure-156@2.[151,1437,113,1539]" captionTargetPageId="2" captionText="Figure 1. Hypodigm of Messelopython freyi gen. et sp. nov. (a) and (b) Photograph and drawing of skull of holotype, SMNK-PAL 461a. (c) Photograph of holotype. (d) and (e) Photograph and drawing of skull of paratype, SMF-ME 2784a. (f) Drawing of left splenial and angular in medial view in counterpart of paratype. (g) Drawing of right ectopterygoid in dorsal view in main part of paratype. (h) and (i) Photograph and drawing of skull of paratype, HLMD-Be 165. Abbreviations: an, angular; boc, basioccipital; bpt-pr, basipterygoid process; bs, basisphenoid; d, dentary; ec, ectopterygoid; fr, frontal; mx, maxilla; n, nasal; ot, otooccipital; pa, parietal; par, prearticular part of compound bone; pl, palatine; pf, postfrontal; prf, prefrontal; pro, prootic; pt, pterygoid; q, quadrate; sa, surangular part of compound bone; smx, septomaxilla; so, supraorbital; soc, supraoccipital; spl, splenial; st, supratemporal." figureDoi="http://doi.org/10.5281/zenodo.4383220" httpUri="https://zenodo.org/record/4383220/files/figure.png" pageId="1" pageNumber="2">figure 1</figureCitation>
|
||
) possesses a short nasal and a pair of long vomerine processes. It is toothed, with six tooth positions, and lacks a median tooth or midline diastema. The long, paired nasals taper anteriorly. The prefrontals lack a lateral foot-process but possess an ascending lamina; these laminae do not meet on the midline. The paired frontals are long and have a concave lateral margin, corresponding to the supraorbital bone, for which an articulation facet is developed (see explanation on homologies of circumorbital bones in electronic supplementary material, characters 73, 74, 77 and 170). The orbital margins of the frontals are upturned. They articulate with the parietal on an anteriorly concave suture. The postfrontal has a bifurcated dorsal articulation on the parietal corner, and it does not reach the frontal. It possesses a long articulation with the supraorbital, which extends some distance ventrally. The large size and crescentric shape of the supraorbital distinguishes
|
||
<taxonomicName authorityName="Zaher & Smith" authorityYear="2020" box="[951,1033,1138,1160]" class="Reptilia" family="Pythonidae" genus="Messelopython" kingdom="Animalia" order="Squamata" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="freyi">
|
||
<emphasis box="[951,1033,1138,1160]" italics="true" pageId="1" pageNumber="2">M. freyi</emphasis>
|
||
</taxonomicName>
|
||
from all extant Pythonoidea (electronic supplementary material,
|
||
<figureCitation box="[1098,1194,1170,1192]" captionStart="Figure 1" captionStartId="2.[113,171,1572,1596]" captionTargetBox="[151,1426,113,1539]" captionTargetId="figure-156@2.[151,1437,113,1539]" captionTargetPageId="2" captionText="Figure 1. Hypodigm of Messelopython freyi gen. et sp. nov. (a) and (b) Photograph and drawing of skull of holotype, SMNK-PAL 461a. (c) Photograph of holotype. (d) and (e) Photograph and drawing of skull of paratype, SMF-ME 2784a. (f) Drawing of left splenial and angular in medial view in counterpart of paratype. (g) Drawing of right ectopterygoid in dorsal view in main part of paratype. (h) and (i) Photograph and drawing of skull of paratype, HLMD-Be 165. Abbreviations: an, angular; boc, basioccipital; bpt-pr, basipterygoid process; bs, basisphenoid; d, dentary; ec, ectopterygoid; fr, frontal; mx, maxilla; n, nasal; ot, otooccipital; pa, parietal; par, prearticular part of compound bone; pl, palatine; pf, postfrontal; prf, prefrontal; pro, prootic; pt, pterygoid; q, quadrate; sa, surangular part of compound bone; smx, septomaxilla; so, supraorbital; soc, supraoccipital; spl, splenial; st, supratemporal." figureDoi="http://doi.org/10.5281/zenodo.4383220" httpUri="https://zenodo.org/record/4383220/files/figure.png" pageId="1" pageNumber="2">figure S1</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="1.[831,1474,541,2120]" pageId="1" pageNumber="2">The elongate parietal possesses a low, mid-sagittal crest over nearly three-quarters of its length. It is somewhat inflated anteriorly. The posterior margin of the parietal contacts the small, oval supraoccipital in an anteriorly directed V-shaped suture, as in most pythonids. The ophidiosphenoid is broad and the foramen for the maxillary branch of the trigeminal nerve is open anteriorly in the prootic. A low, mid-sagittal crest is present on the parabasisphenoid.</paragraph>
|
||
<paragraph blockId="1.[831,1474,541,2120]" pageId="1" pageNumber="2">The supratemporal extends beyond the posterior end of the braincase. The quadrate has a triangular, plate-like dorsal portion, whereby the cephalic condyle is at an oblique angle to the mandibular condyle.</paragraph>
|
||
<paragraph blockId="1.[831,1474,541,2120]" pageId="1" pageNumber="2">
|
||
The maxilla has a weakly sigmoid lateral margin and a single labial foramen. As in most pythons [
|
||
<bibRefCitation author="Scanlon JD" box="[1299,1326,1617,1639]" journalOrPublisher="Mem. Ass. Australas. Palaeont." pageId="1" pageNumber="2" pagination="1 - 35" part="25" refId="ref4227" refString="18. Scanlon JD. 2001 Montypythonoides: the Miocene snake Morelia riversleighensis (Smith & Plane, 1985) and the geographical origin of pythons. Mem. Ass. Australas. Palaeont. 25, 1 - 35. (doi: 10.1080 / 03115510108619211)" title="Montypythonoides: the Miocene snake Morelia riversleighensis (Smith & Plane, 1985) and the geographical origin of pythons" type="journal article" year="2001">18</bibRefCitation>
|
||
], it retains an anterior foramen located just on the medial side of the bone, and two dorsal foramina on the palatine process for the superior alveolar nerve. The palatine process is broad and rounded, rather than posteriorly directed and club-like as in Booidea. The ectopterygoid expands in width anteriorly, where its distal tip is bifurcated. Posteriorly, it possesses a short articulation on the pterygoid, but no pronounced posterior process as in
|
||
<taxonomicName box="[1068,1191,1874,1896]" class="Reptilia" family="Pythonidae" kingdom="Animalia" order="Squamata" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Pythonidae</taxonomicName>
|
||
. The toothed palatine is pierced by the palatine nerve. Details of the articulation with the pterygoid are unknown. Both palatine and pterygoid were toothed, although an exact tooth count cannot be given. The pterygoid terminates posteriorly just short of the mandible.
|
||
</paragraph>
|
||
<paragraph blockId="1.[831,1474,541,2120]" lastBlockId="2.[113,757,1836,2114]" lastPageId="2" lastPageNumber="3" pageId="1" pageNumber="2">The dentary has one mental foramen on the right side of SMF-ME 2784 and two on the left. The surangular notch is not particularly deep. The splenial has a posteroventral knob, which would have inserted into a concavity on the angular. The compound bone possesses a weak but distinct ventrolateral crest. The surangular crest rises relatively straight and gradually, whereas the prearticular crest is strongly convex. The anterior surangular foramen is small.</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.4383220" ID-Zenodo-Dep="4383220" httpUri="https://zenodo.org/record/4383220/files/figure.png" pageId="2" pageNumber="3" startId="2.[113,171,1572,1596]" targetBox="[151,1426,113,1539]" targetPageId="2">
|
||
<paragraph blockId="2.[113,1475,1572,1756]" pageId="2" pageNumber="3">
|
||
Figure 1. Hypodigm of
|
||
<taxonomicName authority="Zaher & Smith, 2020" authorityName="Zaher & Smith" authorityYear="2020" box="[316,475,1572,1596]" class="Reptilia" family="Pythonidae" genus="Messelopython" kingdom="Animalia" order="Squamata" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="freyi" status="gen. et sp. nov.">
|
||
<emphasis bold="true" box="[316,475,1572,1596]" italics="true" pageId="2" pageNumber="3">Messelopython freyi</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel box="[482,613,1572,1596]" pageId="2" pageNumber="3" rank="species">gen. et sp. nov.</taxonomicNameLabel>
|
||
(
|
||
<emphasis bold="true" box="[625,635,1572,1596]" italics="true" pageId="2" pageNumber="3">a</emphasis>
|
||
) and (
|
||
<emphasis bold="true" box="[693,703,1572,1596]" italics="true" pageId="2" pageNumber="3">b</emphasis>
|
||
) Photograph and drawing of skull of holotype, SMNK-PAL 461a. (
|
||
<emphasis bold="true" box="[1252,1259,1572,1596]" italics="true" pageId="2" pageNumber="3">c</emphasis>
|
||
) Photograph of holotype. (
|
||
<emphasis bold="true" box="[120,130,1604,1628]" italics="true" pageId="2" pageNumber="3">d</emphasis>
|
||
) and (
|
||
<emphasis bold="true" box="[194,203,1604,1628]" italics="true" pageId="2" pageNumber="3">e</emphasis>
|
||
) Photograph and drawing of skull of paratype, SMF-ME 2784a. (
|
||
<emphasis bold="true" box="[766,772,1604,1628]" italics="true" pageId="2" pageNumber="3">f</emphasis>
|
||
) Drawing of left splenial and angular in medial view in counterpart of paratype. (
|
||
<emphasis bold="true" box="[121,131,1636,1660]" italics="true" pageId="2" pageNumber="3">g</emphasis>
|
||
) Drawing of right ectopterygoid in dorsal view in main part of paratype. (
|
||
<emphasis bold="true" box="[757,767,1636,1660]" italics="true" pageId="2" pageNumber="3">h</emphasis>
|
||
) and (
|
||
<emphasis bold="true" box="[826,831,1636,1660]" italics="true" pageId="2" pageNumber="3">i</emphasis>
|
||
) Photograph and drawing of skull of paratype, HLMD-Be 165. Abbreviations: an, angular; boc, basioccipital; bpt-pr, basipterygoid process; bs, basisphenoid; d, dentary; ec, ectopterygoid; fr, frontal; mx, maxilla; n, nasal; ot, otooccipital; pa, parietal; par, prearticular part of compound bone; pl, palatine; pf, postfrontal; prf, prefrontal; pro, prootic; pt, pterygoid; q, quadrate; sa, surangular part of compound bone; smx, septomaxilla; so, supraorbital; soc, supraoccipital; spl, splenial; st, supratemporal.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="2.[113,757,1836,2114]" pageId="2" pageNumber="3">The maxilla possesses approximately 18 tooth spaces. The teeth are long and needle-shaped, diminishing from front to back, but anterior teeth are not notably enlarged.</paragraph>
|
||
<paragraph blockId="2.[831,1474,1836,2114]" lastBlockId="3.[113,756,1830,2108]" lastPageId="3" lastPageNumber="4" pageId="2" pageNumber="3">
|
||
There appear to be somewhere between 205 and 210 trunk vertebrae in the
|
||
<typeStatus box="[1073,1169,1868,1890]" pageId="2" pageNumber="3">holotype</typeStatus>
|
||
. HLMD-Be 165 has approximately 209 trunk vertebrae followed by at least two visible cloacals. SMF-ME 2784b preserves the tail, with at least two cloacal vertebrae with bifurcated lymphapophyses followed by approximately 67 vertebrae and tail tip. This yields an approximate total vertebral count of 275. The trunk vertebrae are short and high. Hypapophyses are present only on anterior trunk vertebrae. Mid-trunk vertebrae have an anteroposteriorly long neural spine whose base extends onto the zygosphenal tectum (electronic supplementary material,
|
||
<figureCitation box="[283,390,1894,1916]" captionStart="Figure 2" captionStartId="3.[113,171,1649,1673]" captionTargetBox="[267,1322,118,1610]" captionTargetId="figure-151@3.[304,1145,121,979]" captionTargetPageId="3" captionText="Figure 2. (a) Phylogenetic relationships of Messelopython freyi gen. et sp. nov. Support values for nodes are Jackknife/Bremer, values of 100% (Jackknife) or greater than 20 (Bremer) are indicated with an asterisk. (b) Palaeogeographic map of continental distributions at 48 Mya based on Gplates model [38] with location (palaeocoordinates for 48 Mya) of Palaeogene records of the total clades of Boidae, Loxocemidae and Pythonidae. Large arrows indicate hypothesized dispersal directions for the total clade of Pythonidae." figureDoi="http://doi.org/10.5281/zenodo.4383222" httpUri="https://zenodo.org/record/4383222/files/figure.png" pageId="3" pageNumber="4">figure S2</figureCitation>
|
||
). Short accessory processes are present. Broadly rounded zygantral mounds (see [36,37]) are present even in the posterior trunk. Haemapophyses are present in the tail.
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.4383222" ID-Zenodo-Dep="4383222" httpUri="https://zenodo.org/record/4383222/files/figure.png" pageId="3" pageNumber="4" startId="3.[113,171,1649,1673]" targetBox="[267,1322,118,1610]" targetPageId="3">
|
||
<paragraph blockId="3.[113,1474,1649,1769]" pageId="3" pageNumber="4">
|
||
Figure 2. (
|
||
<emphasis bold="true" box="[208,218,1649,1673]" italics="true" pageId="3" pageNumber="4">a</emphasis>
|
||
) Phylogenetic relationships of
|
||
<taxonomicName authority="Zaher & Smith, 2020" authorityName="Zaher & Smith" authorityYear="2020" box="[474,633,1649,1673]" class="Reptilia" family="Pythonidae" genus="Messelopython" kingdom="Animalia" order="Squamata" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="freyi" status="gen. et sp. nov.">
|
||
<emphasis bold="true" box="[474,633,1649,1673]" italics="true" pageId="3" pageNumber="4">Messelopython freyi</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel box="[639,769,1649,1673]" pageId="3" pageNumber="4" rank="species">gen. et sp. nov.</taxonomicNameLabel>
|
||
Support values for nodes are Jackknife/Bremer, values of 100% (Jackknife) or greater than 20 (Bremer) are indicated with an asterisk. (
|
||
<emphasis bold="true" box="[547,557,1681,1705]" italics="true" pageId="3" pageNumber="4">b</emphasis>
|
||
) Palaeogeographic map of continental distributions at 48 Mya based on Gplates model [
|
||
<bibRefCitation author="Muller RD" box="[1326,1349,1681,1705]" journalOrPublisher="Geochem. Geophys. Geosyst." pageId="3" pageNumber="4" pagination="2243 - 2261" part="19" refId="ref5182" refString="38. Muller RD et al. 2018 GPlates: building a virtual Earth through deep time. Geochem. Geophys. Geosyst. 19, 2243 - 2261. (doi: 10.1029 / 2018 GC 007584)" title="GPlates: building a virtual Earth through deep time" type="journal article" year="2018">38</bibRefCitation>
|
||
] with location (palaeocoordinates for 48 Mya) of Palaeogene records of the total clades of
|
||
<taxonomicName authorityName="Gray" authorityYear="1825" box="[769,826,1713,1737]" class="Reptilia" family="Boidae" kingdom="Animalia" order="Squamata" pageId="3" pageNumber="4" phylum="Chordata" rank="family">Boidae</taxonomicName>
|
||
,
|
||
<taxonomicName box="[838,944,1713,1737]" class="Reptilia" family="Loxocemidae" kingdom="Animalia" order="Squamata" pageId="3" pageNumber="4" phylum="Chordata" rank="family">Loxocemidae</taxonomicName>
|
||
and
|
||
<taxonomicName box="[992,1086,1713,1737]" class="Reptilia" family="Pythonidae" kingdom="Animalia" order="Squamata" pageId="3" pageNumber="4" phylum="Chordata" rank="family">Pythonidae</taxonomicName>
|
||
. Large arrows indicate hypothesized dispersal directions for the total clade of
|
||
<taxonomicName box="[389,483,1745,1769]" class="Reptilia" family="Pythonidae" kingdom="Animalia" order="Squamata" pageId="3" pageNumber="4" phylum="Chordata" rank="family">Pythonidae</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="3.[113,756,1830,2108]" pageId="3" pageNumber="4">
|
||
<pageTitle pageId="3" pageNumber="4">
|
||
Our phylogenetic analysis strongly supports the position of
|
||
<emphasis box="[144,339,2054,2075]" italics="true" pageId="3" pageNumber="4">Messelopython freyi</emphasis>
|
||
on the stem of Pythonidae (figure
|
||
<pageNumber ambiguity="1" box="[727,739,2055,2076]" fuzzyness="0" pageId="3" pageNumber="4" score="1.0" value="2">2</pageNumber>
|
||
<emphasis box="[739,749,2055,2076]" italics="true" pageId="3" pageNumber="4">a</emphasis>
|
||
; electronic supplementary material).
|
||
</pageTitle>
|
||
</paragraph>
|
||
<paragraph blockId="3.[831,1475,1830,2108]" pageId="3" pageNumber="4">
|
||
Excluding
|
||
<taxonomicName authorityName="Zaher & Smith" authorityYear="2020" box="[984,1175,1830,1851]" class="Reptilia" family="Pythonidae" genus="Messelopython" kingdom="Animalia" order="Squamata" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="freyi">
|
||
<emphasis box="[984,1175,1830,1851]" italics="true" pageId="3" pageNumber="4">Messelopython freyi</emphasis>
|
||
</taxonomicName>
|
||
, BioGeoBEARS results support a Gondwanan origin of Alethinophidia and dispersal of stem pythonids over
|
||
<collectingCountry box="[1042,1148,1894,1916]" name="Antarctica" pageId="3" pageNumber="4">Antarctica</collectingCountry>
|
||
to
|
||
<collectingCountry box="[1176,1270,1894,1916]" name="Australia" pageId="3" pageNumber="4">Australia</collectingCountry>
|
||
and Asia. Including
|
||
<taxonomicName authorityName="Zaher & Smith" authorityYear="2020" box="[831,906,1926,1948]" class="Reptilia" family="Pythonidae" genus="Messelopython" kingdom="Animalia" order="Squamata" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="freyi">
|
||
<emphasis box="[831,906,1926,1948]" italics="true" pageId="3" pageNumber="4">M. freyi</emphasis>
|
||
</taxonomicName>
|
||
, both DEC and DEC + J support a Laurasian, and more specifically European, origin of Pythonoidea and
|
||
<taxonomicName box="[1356,1474,1958,1980]" class="Reptilia" family="Pythonidae" kingdom="Animalia" order="Squamata" pageId="3" pageNumber="4" phylum="Chordata" rank="family">Pythonidae</taxonomicName>
|
||
(electronic supplementary material, figures S4–S9), with the dispersal of the total clade of
|
||
<taxonomicName box="[1116,1234,2022,2044]" class="Reptilia" family="Pythonidae" kingdom="Animalia" order="Squamata" pageId="3" pageNumber="4" phylum="Chordata" rank="family">Pythonidae</taxonomicName>
|
||
to Africa and Oceania (
|
||
<figureCitation box="[838,930,2054,2076]" captionStart="Figure 2" captionStartId="3.[113,171,1649,1673]" captionTargetBox="[267,1322,118,1610]" captionTargetId="figure-151@3.[304,1145,121,979]" captionTargetPageId="3" captionText="Figure 2. (a) Phylogenetic relationships of Messelopython freyi gen. et sp. nov. Support values for nodes are Jackknife/Bremer, values of 100% (Jackknife) or greater than 20 (Bremer) are indicated with an asterisk. (b) Palaeogeographic map of continental distributions at 48 Mya based on Gplates model [38] with location (palaeocoordinates for 48 Mya) of Palaeogene records of the total clades of Boidae, Loxocemidae and Pythonidae. Large arrows indicate hypothesized dispersal directions for the total clade of Pythonidae." figureDoi="http://doi.org/10.5281/zenodo.4383222" httpUri="https://zenodo.org/record/4383222/files/figure.png" pageId="3" pageNumber="4">
|
||
figure 2
|
||
<emphasis box="[919,930,2054,2075]" italics="true" pageId="3" pageNumber="4">b</emphasis>
|
||
</figureCitation>
|
||
); analyses assuming an African range for our
|
||
<emphasis box="[1405,1474,2054,2075]" italics="true" pageId="3" pageNumber="4">Python</emphasis>
|
||
terminal do not alter this conclusion.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |