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220 lines
30 KiB
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<document ID-DOI="10.1206/0003-0090.457.1.1" ID-ISSN="0003-0090" ID-Zenodo-Dep="6971356" approvalRequired="59" approvalRequired_for_taxonomicNames="59" checkinTime="1659882917898" checkinUser="felipe" docAuthor="Beck, Robin M. D., Voss, Robert S. & Jansa, Sharon A." docDate="2022" docId="03EFDD5DF6DF68CFD8B5FCAD1B33FB1D" docLanguage="en" docName="BulAmeMusNatHist.2022.457.1-350.pdf" docOrigin="Bulletin of the American Museum of Natural History 2022 (457)" docSource="https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-457/issue-1/0003-0090.457.1.1/Craniodental-Morphology-and-Phylogeny-of-Marsupials/10.1206/0003-0090.457.1.1.full" docStyle="DocumentStyle:915933466F796C9C739DF4DB6B8DCFA6.8:BulAmeMusNatHis.2011-.journal_article.1cover.type1" docStyleId="915933466F796C9C739DF4DB6B8DCFA6" docStyleName="BulAmeMusNatHis.2011-.journal_article.1cover.type1" docStyleVersion="8" docTitle="Phascolarctomorphia Aplin and Archer 1987" docType="treatment" docVersion="9" lastPageNumber="230" masterDocId="FFD6A525F63B6829DA0BFFAE1A62FFC7" masterDocTitle="Craniodental Morphology And Phylogeny Of Marsupials" masterLastPageNumber="353" masterPageNumber="1" pageNumber="228" updateTime="1659987007554" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0">
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<mods:title>Craniodental Morphology And Phylogeny Of Marsupials</mods:title>
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<mods:namePart>Beck, Robin M. D.</mods:namePart>
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<mods:affiliation>School of Science, Engineering and Environment University of Salford, U. K. & School of Biological, Earth & Environmental Sciences University of New South Wales, Australia & Division of Vertebrate Zoology (Mammalogy) American Museum of Natural History</mods:affiliation>
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<mods:namePart>Voss, Robert S.</mods:namePart>
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<mods:affiliation>Division of Vertebrate Zoology (Mammalogy) American Museum of Natural History</mods:affiliation>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Jansa, Sharon A.</mods:namePart>
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<mods:affiliation>Bell Museum and Department of Ecology, Evolution, and Behavior University of Minnesota</mods:affiliation>
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<mods:title>Bulletin of the American Museum of Natural History</mods:title>
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<mods:date>2022</mods:date>
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<mods:number>2022-06-28</mods:number>
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<mods:number>2022</mods:number>
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<mods:number>457</mods:number>
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<mods:identifier type="DOI">10.1206/0003-0090.457.1.1</mods:identifier>
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<mods:identifier type="ISSN">0003-0090</mods:identifier>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.6974221" ID-Zenodo-Dep="6974221" LSID="urn:lsid:plazi:treatment:03EFDD5DF6DF68CFD8B5FCAD1B33FB1D" httpUri="http://treatment.plazi.org/id/03EFDD5DF6DF68CFD8B5FCAD1B33FB1D" lastPageId="230" lastPageNumber="230" pageId="228" pageNumber="228">
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<subSubSection box="[702,1198,771,795]" pageId="228" pageNumber="228" type="nomenclature">
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<paragraph blockId="228.[702,1198,771,795]" box="[702,1198,771,795]" pageId="228" pageNumber="228">
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<heading box="[702,1198,771,795]" centered="true" fontSize="9" level="2" pageId="228" pageNumber="228" reason="2">
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<taxonomicName authority="Aplin and Archer, 1987" authorityName="Aplin and Archer" authorityYear="1987" box="[702,1198,771,795]" genus="Phascolarctomorphia" pageId="228" pageNumber="228" rank="genus">
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Phascolarctomorphia
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<bibRefCitation author="Aplin, K. P. & M. Archer" box="[942,1198,771,795]" pageId="228" pageNumber="228" pagination="xv - lxxii" refId="ref189347" refString="Aplin, K. P., and M. Archer. 1987. Recent advances in marsupial systematics with a new syncretic classification. In M. Archer (editor), Possums and opossums: studies in evolution: xv - lxxii. Sydney: Surrey Beatty and Sons." type="book chapter" year="1987">Aplin and Archer, 1987</bibRefCitation>
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</taxonomicName>
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</heading>
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</paragraph>
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</subSubSection>
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<subSubSection pageId="228" pageNumber="228" type="synonymic_list">
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<paragraph blockId="228.[684,1212,821,1738]" pageId="228" pageNumber="228">
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CONTENTS: †
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<taxonomicName box="[875,979,821,845]" class="Mammalia" family="Phascolarctidae" genus="Litokoala" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diprotodontia" pageId="228" pageNumber="228" phylum="Chordata" rank="genus">
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<emphasis box="[875,979,821,845]" italics="true" pageId="228" pageNumber="228">Litokoala</emphasis>
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</taxonomicName>
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, †
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<taxonomicName baseAuthorityName="Black and Archer" baseAuthorityYear="1997" box="[1017,1146,821,845]" class="Mammalia" family="Phascolarctidae" genus="Nimiokoala" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diprotodontia" pageId="228" pageNumber="228" phylum="Chordata" rank="genus">
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<emphasis box="[1017,1146,821,845]" italics="true" pageId="228" pageNumber="228">Nimiokoala</emphasis>
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</taxonomicName>
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, and
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<taxonomicName box="[684,827,854,878]" class="Mammalia" family="Phascolarctidae" genus="Phascolarctos" kingdom="Animalia" order="Diprotodontia" pageId="228" pageNumber="228" phylum="Chordata" rank="genus">
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<emphasis box="[684,827,854,878]" italics="true" pageId="228" pageNumber="228">Phascolarctos</emphasis>
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</taxonomicName>
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(fig. 44).
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</paragraph>
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</subSubSection>
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<subSubSection pageId="228" pageNumber="228" type="description">
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<paragraph blockId="228.[684,1212,821,1738]" pageId="228" pageNumber="228">
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<collectionCode box="[716,775,887,911]" pageId="228" pageNumber="228">STEM</collectionCode>
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AGE: 32.4 Mya (95%
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<collectionCode box="[1032,1091,887,910]" country="United Kingdom" httpUri="http://biocol.org/urn:lsid:biocol.org:col:13184" lsid="urn:lsid:biocol.org:col:13184" name="Hampstead Scientific Society" pageId="228" pageNumber="228" type="Herbarium">HPD</collectionCode>
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: 29.1–36.4 Mya).
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</paragraph>
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<paragraph blockId="228.[684,1212,821,1738]" pageId="228" pageNumber="228">
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<collectionCode box="[716,800,953,977]" pageId="228" pageNumber="228">CROWN</collectionCode>
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AGE: 25.3 Mya (95%
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<collectionCode box="[1038,1097,954,977]" country="United Kingdom" httpUri="http://biocol.org/urn:lsid:biocol.org:col:13184" lsid="urn:lsid:biocol.org:col:13184" name="Hampstead Scientific Society" pageId="228" pageNumber="228" type="Herbarium">HPD</collectionCode>
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: 19.6–30.3 Mya).
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</paragraph>
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<paragraph blockId="228.[684,1212,821,1738]" pageId="228" pageNumber="228">
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UNAMBIGUOUS CRANIODENTAL
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<collectionCode box="[684,746,1052,1076]" pageId="228" pageNumber="228">SYNAPOMORPHIES</collectionCode>
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: Extracranial course of mandibular nerve traverses a bony canal in the roof of the hypotympanic sinus (char. 52: 1→2; ci = 0.231); postgenoid vein emerges from the postglenoid foramen in the posteromedial corner of the glenoid fossa, medial or anteromedial to the postglenoid process (char. 77: 0→1; ci = 0.250); and additional cuspid labial to m1 protoconid present, forming a cusplike protostylid (char. 165: 0→1; ci = 0.286).
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</paragraph>
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</subSubSection>
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<subSubSection lastPageId="230" lastPageNumber="230" pageId="228" pageNumber="228" type="discussion">
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<paragraph blockId="228.[684,1212,821,1738]" lastBlockId="230.[108,637,226,1242]" lastPageId="230" lastPageNumber="230" pageId="228" pageNumber="228">
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COMMENTS:
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<taxonomicName authorityName="Owen" authorityYear="1839" box="[863,1038,1383,1407]" class="Mammalia" family="Phascolarctidae" kingdom="Animalia" order="Diprotodontia" pageId="228" pageNumber="228" phylum="Chordata" rank="family">Phascolarctidae</taxonomicName>
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is consistently recovered in our molecular, morphological and total-evidence analyses (
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<figureCitation box="[954,1076,1449,1473]" captionStart-0="FIG" captionStart-1="FIG" captionStart-2="FIG" captionStart-3="FIG" captionStart-4="FIG" captionStart-5="FIG" captionStart-6="FIG" captionStartId-0="172.[107,149,1310,1331]" captionStartId-1="173.[108,150,1286,1307]" captionStartId-2="175.[108,148,1247,1268]" captionStartId-3="176.[108,148,1354,1375]" captionStartId-4="178.[108,150,1330,1351]" captionStartId-5="179.[108,150,1262,1283]" captionStartId-6="181.[108,150,938,959]" captionTargetBox-0="[108,1182,229,1274]" captionTargetBox-1="[113,1168,231,1252]" captionTargetBox-2="[128,1155,229,1215]" captionTargetBox-3="[125,1195,236,1325]" captionTargetBox-4="[114,1186,237,1297]" captionTargetBox-5="[132,1186,227,1231]" captionTargetId-0="figure-128@172.[107,1211,224,1284]" captionTargetId-1="figure-136@173.[108,1212,224,1256]" captionTargetId-2="figure-208@175.[108,1212,224,1221]" captionTargetId-3="figure-100@176.[108,1212,224,1328]" captionTargetId-4="figure-125@178.[108,1212,224,1301]" captionTargetId-5="figure-164@179.[131,1190,224,1235]" captionTargetId-6="figure-12@181.[241,1216,224,1485]" captionTargetPageId-0="172" captionTargetPageId-1="173" captionTargetPageId-2="175" captionTargetPageId-3="176" captionTargetPageId-4="178" captionTargetPageId-5="179" captionTargetPageId-6="181" captionText-0="FIG. 27. Fifty-percent majority rule consensus of post-burn-in trees that results from Bayesian analysis of our nuclear sequence-only dataset. Black dots at nodes indicate ≥0.95 Bayesian posterior probability (“strong support”); dark gray dots indicate 0.75–0.94 Bayesian posterior probability (“moderate support”); light gray dots indicate 0.50–0.74 Bayesian posterior probability (“weak support”)." captionText-1="FIG. 28. Fifty-percent majority rule consensus of post-burn-in trees that results from undated Bayesian analysis of our mitochondrial sequence-only dataset. Black dots at nodes indicate ≥0.95 Bayesian posterior probability (“strong support”); dark gray dots indicate 0.75–0.94 Bayesian posterior probability (“moderate support”); light gray dots indicate 0.50–0.74 Bayesian posterior probability (“weak support”)." captionText-2="FIG. 29. Fifty-percent majority rule consensus of post-burn-in trees that results from undated Bayesian analysis of our combined nuclear and mitochondrial sequence dataset. Black dots at nodes indicate ≥0.95 Bayesian posterior probability (“strong support”); dark gray dots indicate 0.75–0.94 Bayesian posterior probability (“moderate support”); light gray dots indicate 0.50–0.74 Bayesian posterior probability (“weak support”)." captionText-3="FIG. 30. Strict consensus of 99,999 most parsimonious trees (each of 949 steps, consistency index =0.269, retention index = 0.815) that result from maximum parsimony analysis of our morphological dataset. Black dots at nodes indicate ≥70% bootstrap support (“strong support”); dark gray dots indicate 50%–69% bootstrap support (“moderate support”); light gray dots indicate <50 bootstrap support (“weak support”)." captionText-4="FIG. 31. Fifty-percent majority rule consensus of post-burn-in trees that results from undated Bayesian analysis of our morphological dataset. Black dots at nodes indicate ≥0.95 Bayesian posterior probability (“strong support”); dark gray dots indicate 0.75–0.94 Bayesian posterior probability (“moderate support”); light gray dots indicate 0.50–0.74 Bayesian posterior probability (“weak support”)." captionText-5="FIG. 32. Fifty-percent majority rule consensus of post-burn-in trees that results from undated Bayesian analysis of our total evidence dataset. Black dots at nodes indicate ≥0.95 Bayesian posterior probability (“strong support”); dark gray dots indicate 0.75–0.94 Bayesian posterior probability (“moderate support”); light gray dots indicate 0.50–0.74 Bayesian posterior probability (“weak support”)." captionText-6="FIG. 33. (opposite page and at right) Fifty-per- cent majority rule consensus of post-burn-in trees that results from dated Bayesian analysis (using combined tip-and-node dating and separate Independent Gamma Rates [IGR] clock models for the molecular and morpho- logical partitions) of our total evidence data- set. Black dots at nodes indicate ≥0.95 Bayesian posterior probability (“strong support”); dark gray dots indicate 0.75–0.94 Bayesian poste- rior probability (“moderate support”); light gray dots indicate 0.50–0.74 Bayesian poste- rior probability (“weak support”). Nodes with- out dots were constrained a priori so that their ages could be calibrated (see tables 6, 13). Orange bars represent 95% Highest Posterior Density (HPD) intervals on the ages of nodes. For clarity, 95% HPD intervals are not shown for the ages of fossil terminals." figureDoi-0="http://doi.org/10.5281/zenodo.6971420" figureDoi-1="http://doi.org/10.5281/zenodo.6971422" figureDoi-2="http://doi.org/10.5281/zenodo.6971424" figureDoi-3="http://doi.org/10.5281/zenodo.6971426" figureDoi-4="http://doi.org/10.5281/zenodo.6971428" figureDoi-5="http://doi.org/10.5281/zenodo.6971431" httpUri-0="https://zenodo.org/record/6971420/files/figure.png" httpUri-1="https://zenodo.org/record/6971422/files/figure.png" httpUri-2="https://zenodo.org/record/6971424/files/figure.png" httpUri-3="https://zenodo.org/record/6971426/files/figure.png" httpUri-4="https://zenodo.org/record/6971428/files/figure.png" httpUri-5="https://zenodo.org/record/6971431/files/figure.png" pageId="228" pageNumber="228">figs. 27–33</figureCitation>
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) as sister to the remaining vombatiforms, which collectively comprise Vombatomorphia (note that we consider †
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<taxonomicName authorityName="Gill" authorityYear="1872" box="[766,955,1548,1572]" class="Mammalia" family="Thylacoleonidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diprotodontia" pageId="228" pageNumber="228" phylum="Chordata" rank="family">Thylacoleonidae</taxonomicName>
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to be
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<taxonomicName authorityName="Owen" authorityYear="1866" box="[1050,1212,1548,1572]" class="Mammalia" kingdom="Animalia" order="Diprotodontia" pageId="228" pageNumber="228" phylum="Chordata" rank="order">Diprotodontia</taxonomicName>
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incertae sedis and not a member of Vombatiformes; see above).
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<bibRefCitation author="Aplin, K. P. & M. Archer" box="[920,1212,1614,1639]" pageId="228" pageNumber="228" pagination="xv - lxxii" refId="ref189347" refString="Aplin, K. P., and M. Archer. 1987. Recent advances in marsupial systematics with a new syncretic classification. In M. Archer (editor), Possums and opossums: studies in evolution: xv - lxxii. Sydney: Surrey Beatty and Sons." type="book chapter" year="1987">Aplin and Archer (1987)</bibRefCitation>
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placed
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<taxonomicName authorityName="Owen" authorityYear="1839" box="[769,944,1647,1671]" class="Mammalia" family="Phascolarctidae" kingdom="Animalia" order="Diprotodontia" pageId="228" pageNumber="228" phylum="Chordata" rank="family">Phascolarctidae</taxonomicName>
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in its own infraorder, Phascolarctomorphia (coordinate to Vombatomorphia), and it remains the only known phascolarctomorphian family; thus, the craniodental synapomorphies of
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<taxonomicName authorityName="Owen" authorityYear="1839" box="[322,491,259,283]" class="Mammalia" family="Phascolarctidae" kingdom="Animalia" order="Diprotodontia" pageId="230" pageNumber="230" phylum="Chordata" rank="family">Phascolarctidae</taxonomicName>
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apply equally to Phascolarctomorphia.
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</paragraph>
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<caption ID-DOI="http://doi.org/10.5281/zenodo.6971457" ID-Zenodo-Dep="6971457" httpUri="https://zenodo.org/record/6971457/files/figure.png" pageId="229" pageNumber="229" startId="229.[108,150,1689,1710]" targetBox="[145,1177,224,1657]" targetPageId="229">
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<paragraph blockId="229.[108,1212,1686,1740]" pageId="229" pageNumber="229">
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FIG. 44.
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<taxonomicName baseAuthorityName="Goldfuss" baseAuthorityYear="1817" box="[196,417,1687,1711]" class="Mammalia" family="Phascolarctidae" genus="Phascolarctos" kingdom="Animalia" order="Diprotodontia" pageId="229" pageNumber="229" phylum="Chordata" rank="species">
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<emphasis box="[196,417,1687,1711]" italics="true" pageId="229" pageNumber="229">Phascolarctos cinereus</emphasis>
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</taxonomicName>
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(
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<taxonomicName authorityName="Owen" authorityYear="1866" box="[430,580,1687,1711]" class="Mammalia" kingdom="Animalia" order="Diprotodontia" pageId="229" pageNumber="229" phylum="Chordata" rank="order">Diprotodontia</taxonomicName>
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,
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<taxonomicName authorityName="Owen" authorityYear="1839" box="[590,751,1687,1711]" class="Mammalia" family="Phascolarctidae" kingdom="Animalia" order="Diprotodontia" pageId="229" pageNumber="229" phylum="Chordata" rank="family">Phascolarctidae</taxonomicName>
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; based on AMNH 65611, an adult male from Queensland).
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</paragraph>
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</caption>
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<paragraph blockId="230.[108,637,226,1242]" pageId="230" pageNumber="230">
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Known phascolarctids are craniodentally distinctive (
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<bibRefCitation author="Sonntag" box="[210,375,358,382]" firstAuthor="Sonntag" pageId="230" pageNumber="230" pagination="863 - 896" refId="ref227856" refString="Sonntag, C. F. 1922. On the myology and classification of the wombat, koala, and phalangers. Proceedings of the Zoological Society of London 92 (4): 863 - 896." type="journal article" year="1922">Sonntag, 1922</bibRefCitation>
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;
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<bibRefCitation author="Archer, M." box="[390,550,358,382]" pageId="230" pageNumber="230" pagination="809 - 815" refId="ref189996" refString="Archer, M. 1984 a. On the importance of being a koala. In M. Archer and G. Clayton (editors), Vertebrate zoogeography and evolution in Australasia: 809 - 815. Perth: Hesperian Press." type="book chapter" year="1984">Archer, 1984a</bibRefCitation>
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,
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<bibRefCitation author="Archer, M." box="[565,631,358,382]" pageId="230" pageNumber="230" pagination="633 - 808" refId="ref190081" refString="Archer, M. 1984 c. The Australian marsupial radiation. In M. Archer and G. Clayton (editors), Vertebrate zoogeography and evolution in Australasia: 633 - 808. Perth: Hesperian Press." type="book chapter" year="1984">1984c</bibRefCitation>
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;
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||
<bibRefCitation author="Aplin, K. P." box="[108,251,391,415]" pageId="230" pageNumber="230" pagination="369 - 391" refId="ref189198" refString="Aplin, K. P. 1987. Basicranial anatomy of the early Miocene diprotodontian Wynyardia bassiana (Marsupialia: Wynyardiidae) and its implications for wynyardiid phylogeny and classification. In M. Archer (editor), Possums and opossums: studies in evolution: 369 - 391. Sydney: Surrey Beatty and Sons." type="book chapter" year="1987">Aplin, 1987</bibRefCitation>
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,
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<bibRefCitation author="Aplin, K. P." box="[273,329,391,415]" pageId="230" pageNumber="230" refId="ref189256" refString="Aplin, K. P. 1990. Basicranial regions of diprotodontian marsupials: anatomy, ontogeny and phylogeny. Ph. D. dissertation, School of Biological Science, University of New South Wales, Sydney." type="book" year="1990">1990</bibRefCitation>
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||
;
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||
<bibRefCitation author="Lee, A. K. & F. N. Carrick" box="[350,631,391,415]" pageId="230" pageNumber="230" pagination="1 - 31" refId="ref213179" refString="Lee, A. K., and F. N. Carrick. 1989. 31. Phascolarctidae. In D. W. Walton and B. J. Richardson (editors), Fauna of Australia, vol. 1 B. Mammalia: 1 - 31. Canberra: AGPS." type="book chapter" year="1989">Lee and Carrick, 1989</bibRefCitation>
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;
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<bibRefCitation author="Springer and Woodburne" box="[108,494,424,448]" firstAuthor="Springer" pageId="230" pageNumber="230" pagination="210 - 221" refId="ref228239" refString="Springer, M. S., and M. O. Woodburne. 1989. The distribution of some basicranial characters within the Marsupialia and a phylogeny of the Phalangeriformes. Journal of Vertebrate Paleontology 9 (2): 210 - 221." type="journal article" year="1989">Springer and Woodburne, 1989</bibRefCitation>
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||
;
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||
<bibRefCitation author="Black, K. H. & M. Archer" pageId="230" pageNumber="230" pagination="209 - 228" refId="ref193871" refString="Black, K. H., and M. Archer. 1997 b. Nimiokoala gen. nov. (Marsupialia, Phascolarctidae) from Riversleigh, northwestern Queensland, with a revision of Litokoala. Memoirs of the Queensland Museum 41 (2): 209 - 228." type="journal article" year="1997">Black and Archer, 1997b</bibRefCitation>
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;
|
||
<bibRefCitation author="Louys, J. & K. P. Aplin & R. M. D. Beck & M. Archer" box="[284,488,457,482]" pageId="230" pageNumber="230" pagination="981 - 992" refId="ref213784" refString="Louys, J., K. P. Aplin, R. M. D. Beck, and M. Archer. 2009. Cranial anatomy of Oligo-Miocene koalas (Diprotodontia: Phascolarctidae): stages in the evolution of an extreme leaf-eating specialization. Journal of Vertebrate Paleontology 29 (4): 981 - 992." type="journal article" year="2009">Louys et al., 2009</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Black, K. H. & J. Louys & G. J. Price" pageId="230" pageNumber="230" pagination="237 - 264" refId="ref194240" refString="Black, K. H., J. Louys, and G. J. Price. 2014 a. Understanding morphological variation in the extant koala as a framework for identification of species boundaries in extinct koalas (Phascolarctidae; Marsupialia). Journal of Systematic Palaeontology 12 (2): 237 - 264." type="journal article" year="2014">Black et al., 2014a</bibRefCitation>
|
||
), and monophyly of this clade is supported by three unambiguous craniodental synapomorphies in our analysis, although all show some degree of homoplasy. Perhaps the most striking of these is the extracranial course of the mandibular nerve, which traverses a bony canal in the roof of the hypotympanic sinus in all three of our phascolarctid terminals (†
|
||
<taxonomicName box="[436,536,722,746]" class="Mammalia" family="Phascolarctidae" genus="Litokoala" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diprotodontia" pageId="230" pageNumber="230" phylum="Chordata" rank="genus">
|
||
<emphasis box="[436,536,722,746]" italics="true" pageId="230" pageNumber="230">Litokoala</emphasis>
|
||
</taxonomicName>
|
||
, †
|
||
<taxonomicName baseAuthorityName="Black and Archer" baseAuthorityYear="1997" class="Mammalia" family="Phascolarctidae" genus="Nimiokoala" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diprotodontia" pageId="230" pageNumber="230" phylum="Chordata" rank="genus">
|
||
<emphasis italics="true" pageId="230" pageNumber="230">Nimiokoala</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName box="[218,360,755,779]" class="Mammalia" family="Phascolarctidae" genus="Phascolarctos" kingdom="Animalia" order="Diprotodontia" pageId="230" pageNumber="230" phylum="Chordata" rank="genus">
|
||
<emphasis box="[218,360,755,779]" italics="true" pageId="230" pageNumber="230">Phascolarctos</emphasis>
|
||
</taxonomicName>
|
||
;
|
||
<bibRefCitation author="Aplin, K. P." box="[372,498,755,779]" pageId="230" pageNumber="230" pagination="369 - 391" refId="ref189198" refString="Aplin, K. P. 1987. Basicranial anatomy of the early Miocene diprotodontian Wynyardia bassiana (Marsupialia: Wynyardiidae) and its implications for wynyardiid phylogeny and classification. In M. Archer (editor), Possums and opossums: studies in evolution: 369 - 391. Sydney: Surrey Beatty and Sons." type="book chapter" year="1987">Aplin, 1987</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Aplin, K. P." box="[508,562,755,779]" pageId="230" pageNumber="230" refId="ref189256" refString="Aplin, K. P. 1990. Basicranial regions of diprotodontian marsupials: anatomy, ontogeny and phylogeny. Ph. D. dissertation, School of Biological Science, University of New South Wales, Sydney." type="book" year="1990">1990</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Louys, J. & K. P. Aplin & R. M. D. Beck & M. Archer" pageId="230" pageNumber="230" pagination="981 - 992" refId="ref213784" refString="Louys, J., K. P. Aplin, R. M. D. Beck, and M. Archer. 2009. Cranial anatomy of Oligo-Miocene koalas (Diprotodontia: Phascolarctidae): stages in the evolution of an extreme leaf-eating specialization. Journal of Vertebrate Paleontology 29 (4): 981 - 992." type="journal article" year="2009">Louys et al., 2009</bibRefCitation>
|
||
), a feature that (as far as we are aware) occurs in no other metatherians.
|
||
<superScript attach="left" box="[466,483,818,835]" fontSize="7" pageId="230" pageNumber="230">34</superScript>
|
||
</paragraph>
|
||
<paragraph blockId="230.[108,637,226,1242]" pageId="230" pageNumber="230">
|
||
In contrast to
|
||
<bibRefCitation author="Black, K. H. & M. Archer & S. J. Hand" box="[301,519,854,878]" pageId="230" pageNumber="230" pagination="125 - 138" refId="ref194056" refString="Black, K. H., M. Archer, and S. J. Hand. 2012 a. New Tertiary koala (Marsupialia, Phascolarctidae) from Riversleigh, Australia, with a revision of phascolarctid phylogenetics, paleoecology, and paleobiodiversity. Journal of Vertebrate Paleontology 32 (1): 125 - 138." type="journal article" year="2012">Black et al. (2012a)</bibRefCitation>
|
||
, we found †
|
||
<taxonomicName baseAuthorityName="Black and Archer" baseAuthorityYear="1997" box="[121,245,887,911]" class="Mammalia" family="Phascolarctidae" genus="Nimiokoala" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diprotodontia" pageId="230" pageNumber="230" phylum="Chordata" rank="genus">
|
||
<emphasis box="[121,245,887,911]" italics="true" pageId="230" pageNumber="230">Nimiokoala</emphasis>
|
||
</taxonomicName>
|
||
, rather than †
|
||
<taxonomicName box="[403,503,887,911]" class="Mammalia" family="Phascolarctidae" genus="Litokoala" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diprotodontia" pageId="230" pageNumber="230" phylum="Chordata" rank="genus">
|
||
<emphasis box="[403,503,887,911]" italics="true" pageId="230" pageNumber="230">Litokoala</emphasis>
|
||
</taxonomicName>
|
||
, to be more closely related to
|
||
<taxonomicName box="[296,439,920,944]" class="Mammalia" family="Phascolarctidae" genus="Phascolarctos" kingdom="Animalia" order="Diprotodontia" pageId="230" pageNumber="230" phylum="Chordata" rank="genus">
|
||
<emphasis box="[296,439,920,944]" italics="true" pageId="230" pageNumber="230">Phascolarctos</emphasis>
|
||
</taxonomicName>
|
||
, with the †
|
||
<taxonomicName baseAuthorityName="Black and Archer" baseAuthorityYear="1997" class="Mammalia" family="Phascolarctidae" genus="Nimiokoala" higherTaxonomySource="GBIF" kingdom="Animalia" order="Diprotodontia" pageId="230" pageNumber="230" phylum="Chordata" rank="genus">
|
||
<emphasis italics="true" pageId="230" pageNumber="230">Nimiokoala</emphasis>
|
||
</taxonomicName>
|
||
+
|
||
<taxonomicName box="[194,337,953,977]" class="Mammalia" family="Phascolarctidae" genus="Phascolarctos" kingdom="Animalia" order="Diprotodontia" pageId="230" pageNumber="230" phylum="Chordata" rank="genus">
|
||
<emphasis box="[194,337,953,977]" italics="true" pageId="230" pageNumber="230">Phascolarctos</emphasis>
|
||
</taxonomicName>
|
||
clade supported by a single unambiguous synapomorphy (see file S
|
||
<quantity box="[549,592,986,1010]" metricMagnitude="-2" metricUnit="m" metricValue="7.62" pageId="230" pageNumber="230" unit="in" value="3.0">3 in</quantity>
|
||
the online supplement): maxillary and frontal bones in contact on medial orbital wall (char. 13 0→1; ci = 0.143). Like other vombatiform families, the oldest record of
|
||
<taxonomicName authorityName="Owen" authorityYear="1839" box="[285,456,1118,1142]" class="Mammalia" family="Phascolarctidae" kingdom="Animalia" order="Diprotodontia" pageId="230" pageNumber="230" phylum="Chordata" rank="family">Phascolarctidae</taxonomicName>
|
||
is from late Oligocene sites in
|
||
<collectingCountry box="[290,391,1151,1175]" name="Australia" pageId="230" pageNumber="230">Australia</collectingCountry>
|
||
(
|
||
<bibRefCitation author="Bassarova, M. & M. Archer" box="[410,631,1151,1176]" pageId="230" pageNumber="230" pagination="25 - 27" refId="ref192343" refString="Bassarova, M., and M. Archer. 1999. Living and extinct pseudocheirids (Marsupialia, Pseudocheiridae): Phylogenetic relationships and changes in diversity through time. Australian Mammalogy 21: 25 - 27." type="journal article" year="1999">Archer et al., 1999</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Long, J. A. & M. Archer & T. F. Flannery & S. J. Hand" box="[108,293,1184,1209]" pageId="230" pageNumber="230" refId="ref213561" refString="Long, J. A., M. Archer, T. F. Flannery, and S. J. Hand. 2002. Prehistoric mammals of Australia and New Guinea: one hundred million years of evolution, Sydney: UNSW Press." type="book" year="2002">Long et al., 2002</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Archer, M. & S. J. Hand" box="[304,565,1184,1208]" pageId="230" pageNumber="230" pagination="575 - 646" refId="ref190275" refString="Archer, M., and S. J. Hand. 2006. The Australian marsupial radiation. In J. R. Merrick, M. Archer, G. M. Hickey, and M. S. Y. Lee (editors), Evolution and biogeography of Australasian vertebrates: 575 - 646. Sydney: Auscipub Pty Ltd." type="book chapter" year="2006">Archer and Hand, 2006</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Black, K. H. & M. Archer & S. J. Hand & H. Godthelp" pageId="230" pageNumber="230" pagination="983 - 1078" refId="ref194115" refString="Black, K. H., M. Archer, S. J. Hand, and H. Godthelp. 2012 b. The rise of Australian marsupials: a synopsis of biostratigraphic, phylogenetic, palaeoecologic and palaeobiogeographic understanding. In J. A. Talent (editor), Earth and life: global biodiversity, extinction intervals and biogeographic perturbations through time: 983 - 1078. Dordrecht: Springer Verlag." type="book chapter" year="2012">Black et al., 2012b</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Black, K. H. & G. J. Price & M. Archer & S. J. Hand" box="[254,325,1218,1242]" pageId="230" pageNumber="230" pagination="1186 - 1201" refId="ref194298" refString="Black, K. H., G. J. Price, M. Archer, and S. J. Hand. 2014 b. Bearing up well? Understanding the past, present and future of Australia's koalas. Gondwana Research 25 (3): 1186 - 1201." type="journal article" year="2014">2014b</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |