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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. &amp; Jansa, Sharon A." docDate="2022" docId="03EFDD5DF6E968FAD925FCAD1E05FC16" 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="Perameloidea Gray 1825" docType="treatment" docVersion="9" lastPageNumber="211" masterDocId="FFD6A525F63B6829DA0BFFAE1A62FFC7" masterDocTitle="Craniodental Morphology And Phylogeny Of Marsupials" masterLastPageNumber="353" masterPageNumber="1" pageNumber="210" 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:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Beck, Robin M. D.</mods:namePart>
<mods:affiliation>School of Science, Engineering and Environment University of Salford, U. K. &amp; School of Biological, Earth &amp; Environmental Sciences University of New South Wales, Australia &amp; 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>Voss, Robert S.</mods:namePart>
<mods:affiliation>Division of Vertebrate Zoology (Mammalogy) American Museum of Natural History</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Jansa, Sharon A.</mods:namePart>
<mods:affiliation>Bell Museum and Department of Ecology, Evolution, and Behavior University of Minnesota</mods:affiliation>
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<mods:date>2022</mods:date>
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<mods:number>2022-06-28</mods:number>
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<paragraph blockId="210.[814,1085,771,795]" box="[814,1085,771,795]" pageId="210" pageNumber="210">
<heading box="[814,1085,771,795]" centered="true" fontSize="9" level="2" pageId="210" pageNumber="210" reason="2">
<taxonomicName authority="Gray, 1825" authorityName="Gray" authorityYear="1825" box="[814,1085,771,795]" pageId="210" pageNumber="210" rank="superFamily" superFamily="Perameloidea">
Perameloidea
<bibRefCitation author="Gray, J. E." box="[968,1085,771,795]" pageId="210" pageNumber="210" pagination="337 - 344" refId="ref207550" refString="Gray, J. E. 1825. An outline of an attempt at the disposition of the Mammalia into tribes and families with a list of the genera apparently appertaining to each tribe. Annals of Philosophy 10 (2): 337 - 344." type="journal article" year="1825">Gray, 1825</bibRefCitation>
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection pageId="210" pageNumber="210" type="synonymic_list">
<paragraph blockId="210.[684,1214,821,1738]" pageId="210" pageNumber="210">
CONTENTS:
<taxonomicName authorityName="Gill" authorityYear="1872" box="[846,1016,821,845]" class="Mammalia" family="Chaeropodidae" kingdom="Animalia" order="Peramelemorphia" pageId="210" pageNumber="210" phylum="Chordata" rank="family">Chaeropodidae</taxonomicName>
,
<taxonomicName authorityName="Gray" authorityYear="1825" box="[1026,1162,821,845]" class="Mammalia" family="Peramelidae" kingdom="Animalia" order="Peramelemorphia" pageId="210" pageNumber="210" phylum="Chordata" rank="family">Peramelidae</taxonomicName>
, and
<taxonomicName authorityName="Bensley" authorityYear="1903" box="[684,852,854,878]" class="Mammalia" family="Thylacomyidae" kingdom="Animalia" order="Peramelemorphia" pageId="210" pageNumber="210" phylum="Chordata" rank="family">Thylacomyidae</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection pageId="210" pageNumber="210" type="description">
<paragraph blockId="210.[684,1214,821,1738]" pageId="210" pageNumber="210">
<collectionCode box="[716,775,887,911]" pageId="210" pageNumber="210">STEM</collectionCode>
AGE: 26.1 Mya (95%
<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="210" pageNumber="210" type="Herbarium">HPD</collectionCode>
: 20.731.8 Mya).
</paragraph>
<paragraph blockId="210.[684,1214,821,1738]" pageId="210" pageNumber="210">
<collectionCode box="[716,800,953,977]" pageId="210" pageNumber="210">CROWN</collectionCode>
AGE: 20.0 Mya (95%
<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="210" pageNumber="210" type="Herbarium">HPD</collectionCode>
: 15.324.5 Mya).
</paragraph>
<paragraph blockId="210.[684,1214,821,1738]" pageId="210" pageNumber="210">
UNAMBIGUOUS CRANIODENTAL
<collectionCode box="[684,748,1052,1076]" pageId="210" pageNumber="210">SYNAPOMORPHIES</collectionCode>
: Nasals truncated anterior to lacrimals (char. 3: 0→1; ci = 0.333); lacrimal exposure on orbital rim forming a distinct crest (char. 8: 0→2; ci = 0.118); frontal and squamosal in contact on lateral aspect of braincase (char. 26: 0→1; ci = 0.071); palatine fenestrae present (char. 38: 0→1; ci = 0.071); posterior epitympanic sinus of squamosal present, enclosed laterally (char. 84: 0→1; ci = 0.071); and entocristid indistinct or absent (char. 176: 0→1; ci = 0.077).
</paragraph>
</subSubSection>
<subSubSection lastPageId="211" lastPageNumber="211" pageId="210" pageNumber="210" type="discussion">
<paragraph blockId="210.[684,1214,821,1738]" pageId="210" pageNumber="210">
COMMENTS: We follow
<bibRefCitation author="Van Dyck and Strahan's" firstAuthor="Van Dyck" pageId="210" pageNumber="210" refId="ref231893" refString="Van Dyck, S., and R. Strahan (editors). 2008. The mammals of Australia, 3 rd ed. Sydney: New Holland Publishers." type="book" year="2008">
<collectingRegion box="[977,1020,1383,1406]" country="Turkey" name="Van" pageId="210" pageNumber="210">Van</collectingRegion>
Dyck and Strahans (2008)
</bibRefCitation>
use of Perameloidea to refer only to the peramelemorphian crown clade, as did
<bibRefCitation author="Kear, B. P. &amp; K. P. Aplin &amp; M. Westerman" pageId="210" pageNumber="210" pagination="37537" refId="ref210965" refString="Kear, B. P., K. P. Aplin, and M. Westerman. 2016. Bandicoot fossils and DNA elucidate lineage antiquity amongst xeric-adapted Australasian marsupials. Scientific Reports 6: 37537." type="journal article" year="2016">Kear et al. (2016)</bibRefCitation>
. By contrast, other recent studies (e.g.,
<bibRefCitation author="Chamberlain, P. M. &amp; K. J. Travouillon &amp; M. Archer &amp; S. J. Hand" box="[747,1026,1515,1539]" pageId="210" pageNumber="210" pagination="197 - 206" refId="ref197024" refString="Chamberlain, P. M., K. J. Travouillon, M. Archer, and S. J. Hand. 2015. Kutjamarcoot brevirostrum gen. et sp. nov., a new short-snouted, early Miocene bandicoot (Marsupialia: Peramelemorphia) from the Kutjamarpu Local Fauna (Wipajiri Formation) in South Australia. Alcheringa: an Australasian Journal of Palaeontology 40 (2): 197 - 206." type="journal article" year="2015">Chamberlain et al., 2015</bibRefCitation>
;
<bibRefCitation author="Warburton and Travouillon" firstAuthor="Warburton" pageId="210" pageNumber="210" pagination="151 - 181" refId="ref232688" refString="Warburton, N. M., and K. J. Travouillon. 2016. The biology and palaeontology of the Peramelemorphia: a review of current knowledge and future research directions. Australian Journal of Zoology 64 (3): 151 - 181." type="journal article" year="2016">Warburton and Travouillon, 2016</bibRefCitation>
) have used Perameloidea to refer to the crown clade plus all taxa more closely related to it than to the fossil genus †
<taxonomicName baseAuthorityName="Muirhead and Filan" baseAuthorityYear="1995" box="[1135,1205,1614,1638]" class="Mammalia" family="Yaralidae" genus="Yarala" higherTaxonomySource="GBIF" kingdom="Animalia" order="Peramelemorphia" pageId="210" pageNumber="210" phylum="Chordata" rank="genus">
<emphasis box="[1135,1205,1614,1638]" italics="true" pageId="210" pageNumber="210">Yarala</emphasis>
</taxonomicName>
, which has been placed in the family †
<taxonomicName box="[1109,1212,1647,1671]" class="Mammalia" family="Yaralidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Peramelemorphia" pageId="210" pageNumber="210" phylum="Chordata" rank="family">Yaralidae</taxonomicName>
and superfamily †Yaraloidea (
<bibRefCitation author="Muirhead, J." box="[1025,1208,1680,1705]" pageId="210" pageNumber="210" pagination="512 - 523" refId="ref217438" refString="Muirhead, J. 2000. Yaraloidea (Marsupialia, Peramelemorphia), a new superfamily of marsupial and a description and analysis of the cranium of the Miocene of Yarala burchfieldi. Journal of Paleontology 74 (3): 512 - 523." type="journal article" year="2000">Muirhead, 2000</bibRefCitation>
;
<bibRefCitation author="Warburton and Travouillon" box="[684,1057,1714,1738]" firstAuthor="Warburton" pageId="210" pageNumber="210" pagination="151 - 181" refId="ref232688" refString="Warburton, N. M., and K. J. Travouillon. 2016. The biology and palaeontology of the Peramelemorphia: a review of current knowledge and future research directions. Australian Journal of Zoology 64 (3): 151 - 181." type="journal article" year="2016">Warburton and Travouillon, 2016</bibRefCitation>
).
</paragraph>
<paragraph blockId="211.[108,638,226,1638]" pageId="211" pageNumber="211">
Within Perameloidea, all our molecular (figs. 2729) and total-evidence (figs. 32,
<figureCitation box="[600,628,259,283]" captionStart="FIG" captionStartId="181.[108,150,938,959]" captionTargetId="figure-12@181.[241,1216,224,1485]" captionTargetPageId="181" captionText="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 0.750.94 Bayesian poste- rior probability (“moderate support”); light gray dots indicate 0.500.74 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." pageId="211" pageNumber="211">33</figureCitation>
) analyses support
<taxonomicName authorityName="Gray" authorityYear="1825" box="[323,465,292,316]" class="Mammalia" family="Peramelidae" kingdom="Animalia" order="Peramelemorphia" pageId="211" pageNumber="211" phylum="Chordata" rank="family">Peramelidae</taxonomicName>
and
<taxonomicName authorityName="Reid" authorityYear="1837" box="[536,636,292,315]" class="Mammalia" family="Thylacomyidae" genus="Macrotis" kingdom="Animalia" order="Peramelemorphia" pageId="211" pageNumber="211" phylum="Chordata" rank="genus">
<emphasis box="[536,636,292,315]" italics="true" pageId="211" pageNumber="211">Macrotis</emphasis>
</taxonomicName>
(
<taxonomicName authorityName="Bensley" authorityYear="1903" box="[116,299,325,349]" class="Mammalia" family="Thylacomyidae" kingdom="Animalia" order="Peramelemorphia" pageId="211" pageNumber="211" phylum="Chordata" rank="family">Thylacomyidae</taxonomicName>
) as sister taxa, with
<taxonomicName authorityName="Ogilby" authorityYear="1838" class="Mammalia" family="Chaeropodidae" genus="Chaeropus" kingdom="Animalia" order="Peramelemorphia" pageId="211" pageNumber="211" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="211" pageNumber="211">Chaeropus</emphasis>
</taxonomicName>
(
<taxonomicName authorityName="Gill" authorityYear="1872" box="[191,374,358,382]" class="Mammalia" family="Chaeropodidae" kingdom="Animalia" order="Peramelemorphia" pageId="211" pageNumber="211" phylum="Chordata" rank="family">Chaeropodidae</taxonomicName>
) the first perameloid family to diverge. This topology is congruent with most other recent molecular and totalevidence phylogenetic analyses (
<bibRefCitation author="Meredith, R. W. &amp; M. Westerman &amp; M. S. Springer" pageId="211" pageNumber="211" pagination="1 - 20" refId="ref216532" refString="Meredith, R. W., M. Westerman, and M. S. Springer. 2008 a. A timescale and phylogeny for &quot; bandicoots &quot; (Peramelemorphia: Marsupialia) based on sequences for five nuclear genes. Molecular Phylogenetics and Evolution 47 (1): 1 - 20." type="journal article" year="2008">Meredith et al., 2008a</bibRefCitation>
;
<bibRefCitation author="Westerman" box="[279,564,490,514]" etAl="et al." firstAuthor="Westerman" pageId="211" pageNumber="211" pagination="97 - 108" refId="ref233182" refString="Westerman, M., et al. 2012. Phylogenetic relationships of living and recently extinct bandicoots based on nuclear and mitochondrial DNA sequences. Molecular Phylogenetics and Evolution 62 (1): 97 - 108." type="journal article" year="2012">Westerman et al., 2012</bibRefCitation>
;
<bibRefCitation author="Kear, B. P. &amp; K. P. Aplin &amp; M. Westerman" pageId="211" pageNumber="211" pagination="37537" refId="ref210965" refString="Kear, B. P., K. P. Aplin, and M. Westerman. 2016. Bandicoot fossils and DNA elucidate lineage antiquity amongst xeric-adapted Australasian marsupials. Scientific Reports 6: 37537." type="journal article" year="2016">Kear et al., 2016</bibRefCitation>
;
<bibRefCitation author="Travouillon and Phillips" box="[259,623,523,547]" firstAuthor="Travouillon" pageId="211" pageNumber="211" pagination="224 - 256" refId="ref230320" refString="Travouillon, K. J., and M. J. Phillips. 2018. Total evidence analysis of the phylogenetic relationships of bandicoots and bilbies (Marsupialia: Peramelemorphia): reassessment of two species and description of a new species. Zootaxa 4378 (2): 224 - 256." type="journal article" year="2018">Travouillon and Phillips, 2018</bibRefCitation>
). However, the dated total-evidence analysis of
<bibRefCitation author="Travouillon and Phillips" box="[140,497,589,613]" firstAuthor="Travouillon" pageId="211" pageNumber="211" pagination="224 - 256" refId="ref230320" refString="Travouillon, K. J., and M. J. Phillips. 2018. Total evidence analysis of the phylogenetic relationships of bandicoots and bilbies (Marsupialia: Peramelemorphia): reassessment of two species and description of a new species. Zootaxa 4378 (2): 224 - 256." type="journal article" year="2018">Travouillon and Phillips (2018</bibRefCitation>
: figs. 1E, 2) placed
<taxonomicName authorityName="Ogilby" authorityYear="1838" box="[190,308,622,646]" class="Mammalia" family="Chaeropodidae" genus="Chaeropus" kingdom="Animalia" order="Peramelemorphia" pageId="211" pageNumber="211" phylum="Chordata" rank="genus">
<emphasis box="[190,308,622,646]" italics="true" pageId="211" pageNumber="211">Chaeropus</emphasis>
</taxonomicName>
sister to
<taxonomicName authorityName="Reid" authorityYear="1837" box="[419,514,623,646]" class="Mammalia" family="Thylacomyidae" genus="Macrotis" kingdom="Animalia" order="Peramelemorphia" pageId="211" pageNumber="211" phylum="Chordata" rank="genus">
<emphasis box="[419,514,623,646]" italics="true" pageId="211" pageNumber="211">Macrotis</emphasis>
</taxonomicName>
, the dated total-evidence analyses of
<bibRefCitation author="Travouillon" etAl="et al." firstAuthor="Travouillon" pageId="211" pageNumber="211" pagination="109 - 125" refId="ref231075" refString="Travouillon, K. J., R. M. D. Beck, and J. A. Case. 2021. Upper Oligocene - lower-Middle Miocene peramelemorphians from the Etadunna, Namba and Wipajiri formations of South Australia. Alcheringa: an Australasian Journal of Palaeontology 45 (1): 109 - 125." type="journal article" year="2021">Travouillon et al. (2021)</bibRefCitation>
placed
<taxonomicName authorityName="Bensley" authorityYear="1903" box="[273,446,688,712]" class="Mammalia" family="Thylacomyidae" kingdom="Animalia" order="Peramelemorphia" pageId="211" pageNumber="211" phylum="Chordata" rank="family">Thylacomyidae</taxonomicName>
outside
<taxonomicName authorityName="Ogilby" authorityYear="1838" class="Mammalia" family="Chaeropodidae" genus="Chaeropus" kingdom="Animalia" order="Peramelemorphia" pageId="211" pageNumber="211" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="211" pageNumber="211">Chaeropus</emphasis>
</taxonomicName>
+
<taxonomicName authorityName="Gray" authorityYear="1825" box="[183,326,722,746]" class="Mammalia" family="Peramelidae" kingdom="Animalia" order="Peramelemorphia" pageId="211" pageNumber="211" phylum="Chordata" rank="family">Peramelidae</taxonomicName>
,
<superScript attach="left" box="[331,349,719,736]" fontSize="7" pageId="211" pageNumber="211">31</superScript>
and the molecular phylogenetic analyses of
<bibRefCitation author="Travouillon" box="[355,636,755,779]" etAl="et al." firstAuthor="Travouillon" pageId="211" pageNumber="211" pagination="1 - 56" refId="ref232735" refString="Warburton, N. M., K. J. Travouillon, and A. B. Camens. 2019. Skeletal atlas of the thylacine (Thylacinus cynocephalus). Palaeontologia Electronica 22.2.29 A: 1 - 56." type="book chapter" year="2019">Travouillon et al. (2019)</bibRefCitation>
placed
<taxonomicName authorityName="Ogilby" authorityYear="1838" box="[196,316,788,812]" class="Mammalia" family="Chaeropodidae" genus="Chaeropus" kingdom="Animalia" order="Peramelemorphia" pageId="211" pageNumber="211" phylum="Chordata" rank="genus">
<emphasis box="[196,316,788,812]" italics="true" pageId="211" pageNumber="211">Chaeropus</emphasis>
</taxonomicName>
either sister to
<taxonomicName authorityName="Desmarest" authorityYear="1817" box="[519,608,788,812]" class="Mammalia" family="Peramelidae" genus="Isoodon" kingdom="Animalia" order="Peramelemorphia" pageId="211" pageNumber="211" phylum="Chordata" rank="genus">
<emphasis box="[519,608,788,812]" italics="true" pageId="211" pageNumber="211">Isoodon</emphasis>
</taxonomicName>
+
<taxonomicName authorityName="E. Geoffroy Saint-Hilaire" authorityYear="1804" box="[108,219,821,845]" class="Mammalia" family="Peramelidae" genus="Perameles" kingdom="Animalia" order="Peramelemorphia" pageId="211" pageNumber="211" phylum="Chordata" rank="genus">
<emphasis box="[108,219,821,845]" italics="true" pageId="211" pageNumber="211">Perameles</emphasis>
</taxonomicName>
+
<taxonomicName box="[257,371,822,845]" class="Mammalia" family="Peramelidae" genus="Peroryctes" kingdom="Animalia" order="Peramelemorphia" pageId="211" pageNumber="211" phylum="Chordata" rank="genus">
<emphasis box="[257,371,822,845]" italics="true" pageId="211" pageNumber="211">Peroryctes</emphasis>
</taxonomicName>
(in an undated maximum likelihood analysis;
<bibRefCitation author="Travouillon" etAl="et al." firstAuthor="Travouillon" pageId="211" pageNumber="211" pagination="1 - 56" refId="ref232735" refString="Warburton, N. M., K. J. Travouillon, and A. B. Camens. 2019. Skeletal atlas of the thylacine (Thylacinus cynocephalus). Palaeontologia Electronica 22.2.29 A: 1 - 56." type="book chapter" year="2019">Travouillon et al., 2019</bibRefCitation>
: fig. 15) or sister to
<taxonomicName authorityName="Desmarest" authorityYear="1817" box="[403,490,887,911]" class="Mammalia" family="Peramelidae" genus="Isoodon" kingdom="Animalia" order="Peramelemorphia" pageId="211" pageNumber="211" phylum="Chordata" rank="genus">
<emphasis box="[403,490,887,911]" italics="true" pageId="211" pageNumber="211">Isoodon</emphasis>
</taxonomicName>
(in a Bayesian node-dating analysis;
<bibRefCitation author="Travouillon" etAl="et al." firstAuthor="Travouillon" pageId="211" pageNumber="211" pagination="1 - 56" refId="ref232735" refString="Warburton, N. M., K. J. Travouillon, and A. B. Camens. 2019. Skeletal atlas of the thylacine (Thylacinus cynocephalus). Palaeontologia Electronica 22.2.29 A: 1 - 56." type="book chapter" year="2019">Travouillon et al., 2019</bibRefCitation>
: fig. 16).
</paragraph>
<paragraph blockId="211.[108,638,226,1638]" pageId="211" pageNumber="211">
We identified a number of unambiguous cranidodental synapomorphies as characterizing Perameloidea, although most of these are homoplastic. Perhaps the most notable of these is truncation of the nasals anterior to the lacrimals, a trait that is present in all known perameloids (and arose homoplastically in
<taxonomicName baseAuthorityName="Gadow" baseAuthorityYear="1892" box="[108,224,1218,1241]" class="Mammalia" family="Notoryctidae" genus="Notoryctes" kingdom="Animalia" order="Notoryctemorphia" pageId="211" pageNumber="211" phylum="Chordata" rank="genus">
<emphasis box="[108,224,1218,1241]" italics="true" pageId="211" pageNumber="211">Notoryctes</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Beck &amp; Voss &amp; Jansa" authorityYear="2022" box="[286,382,1218,1241]" class="Mammalia" family="Potoroidae" genus="Potorous" kingdom="Animalia" order="Diprotodontia" pageId="211" pageNumber="211" phylum="Chordata" rank="genus">
<emphasis box="[286,382,1218,1241]" italics="true" pageId="211" pageNumber="211">Potorous</emphasis>
</taxonomicName>
) but that is absent in nonperameloid peramelemorphians, all of which are “short-snouted” (
<bibRefCitation author="Travouillon" etAl="et al." firstAuthor="Travouillon" pageId="211" pageNumber="211" pagination="1528 - 1546" refId="ref230485" refString="Travouillon, K. J., Y. Gurovich, R. M. D. Beck, and J. Muirhead. 2010. An exceptionally well-preserved short-snouted bandicoot (Marsupialia; Peramelemorphia) from Riversleigh's Oligo-Miocene deposits, northwestern Queensland, Australia. Journal of Vertebrate Paleontology 30 (5): 1528 - 1546." type="journal article" year="2010">Travouillon et al., 2010</bibRefCitation>
,
<bibRefCitation author="Travouillon" box="[185,256,1317,1341]" firstAuthor="Travouillon" pageId="211" pageNumber="211" pagination="153 - 168" refId="ref230601" refString="Travouillon, K. J., Y. Gurovich, M. Archer, S. J. Hand, and J. Muirhead. 2013 b. The genus Galadi: three new bandicoots (Marsupialia, Peramelemorphia) from Riversleigh's Miocene deposits, Northwestern Queensland, Australia. Journal of Vertebrate Paleontology 33 (1): 153 - 168." type="journal article" year="2013" yearSuffix="b">2013b</bibRefCitation>
,
<bibRefCitation author="Travouillon" box="[275,347,1317,1341]" firstAuthor="Travouillon" pageId="211" pageNumber="211" pagination="141 - 167" refId="ref230854" refString="Travouillon, K. J., M. Archer, S. J. Hand, and J. Muirhead. 2015 b. Sexually dimorphic bandicoots (Marsupialia: Peramelemorphia) from the Oligo-Miocene of Australia, first cranial ontogeny for fossil bandicoots and new species descriptions. Journal of Mammalian Evolution 22: 141 - 167." type="journal article" year="2015" yearSuffix="b">2015b</bibRefCitation>
;
<bibRefCitation author="Gurovich, Y. &amp; K. J. Travouillon &amp; R. M. D. Beck &amp; J. Muirhead &amp; M. Archer" box="[367,631,1317,1341]" pageId="211" pageNumber="211" pagination="265 - 290" refId="ref208146" refString="Gurovich, Y., K. J. Travouillon, R. M. D. Beck, J. Muirhead, and M. Archer. 2014. Biogeographical implications of a new mouse-sized fossil bandicoot (Marsupialia: Peramelemorphia) occupying a dasyurid-like ecological niche across Australia. Journal of Systematic Palaeontology 12 (3): 265 - 290." type="journal article" year="2014">Gurovich et al., 2014</bibRefCitation>
;
<bibRefCitation author="Chamberlain, P. M. &amp; K. J. Travouillon &amp; M. Archer &amp; S. J. Hand" box="[108,387,1350,1374]" pageId="211" pageNumber="211" pagination="197 - 206" refId="ref197024" refString="Chamberlain, P. M., K. J. Travouillon, M. Archer, and S. J. Hand. 2015. Kutjamarcoot brevirostrum gen. et sp. nov., a new short-snouted, early Miocene bandicoot (Marsupialia: Peramelemorphia) from the Kutjamarpu Local Fauna (Wipajiri Formation) in South Australia. Alcheringa: an Australasian Journal of Palaeontology 40 (2): 197 - 206." type="journal article" year="2015">Chamberlain et al., 2015</bibRefCitation>
).
</paragraph>
<paragraph blockId="211.[108,638,226,1638]" lastBlockId="211.[684,1212,226,977]" pageId="211" pageNumber="211">
The oldest putative perameloid is †
<emphasis italics="true" pageId="211" pageNumber="211">Bulbadon warburtonae</emphasis>
from the late Oligocene Ditjimanka Local Fauna (Faunal Zone B) of the Etadunna Formation, which is known from a single partial mandible, and has been tentatively identified as a thylacomyid (
<bibRefCitation author="Travouillon" box="[367,625,1548,1572]" etAl="et al." firstAuthor="Travouillon" pageId="211" pageNumber="211" pagination="109 - 125" refId="ref231075" refString="Travouillon, K. J., R. M. D. Beck, and J. A. Case. 2021. Upper Oligocene - lower-Middle Miocene peramelemorphians from the Etadunna, Namba and Wipajiri formations of South Australia. Alcheringa: an Australasian Journal of Palaeontology 45 (1): 109 - 125." type="journal article" year="2021">Travouillon et al., 2021</bibRefCitation>
). A best-fit age-model of paleomagnetic data indicates that the Etadunna Formation spans 23.626.1 Mya (
<bibRefCitation author="Metzger, C. A. &amp; G. J. Retallack" box="[868,1201,226,250]" pageId="211" pageNumber="211" pagination="871 - 885" refId="ref216925" refString="Metzger, C. A., and G. J. Retallack. 2010. Paleosol record of Neogene climate change in the Australian outback. Australian Journal of Earth Sciences 57: 871 - 885." type="journal article" year="2010">Metzger and Retallack, 2010</bibRefCitation>
), so, if †
<emphasis box="[763,868,259,283]" italics="true" pageId="211" pageNumber="211">Bulbadon</emphasis>
is a thylacomyid, then it predates our estimate for the crown age of Perameloidea. However, †
<emphasis box="[929,1034,325,349]" italics="true" pageId="211" pageNumber="211">Bulbadon</emphasis>
did not form a clade with other thylacomyids in the dated total-evidence analyses of
<bibRefCitation author="Travouillon" etAl="et al." firstAuthor="Travouillon" pageId="211" pageNumber="211" pagination="109 - 125" refId="ref231075" refString="Travouillon, K. J., R. M. D. Beck, and J. A. Case. 2021. Upper Oligocene - lower-Middle Miocene peramelemorphians from the Etadunna, Namba and Wipajiri formations of South Australia. Alcheringa: an Australasian Journal of Palaeontology 45 (1): 109 - 125." type="journal article" year="2021">Travouillon et al. (2021)</bibRefCitation>
and a lack of resolution meant that a position for †
<emphasis box="[847,954,457,481]" italics="true" pageId="211" pageNumber="211">Bulbadon</emphasis>
outside Perameloidea could not be ruled out in their analyses. Two taxa from the middle Miocene of Riversleigh World Heritage Area, both known only from isolated teeth, may also be early perameloids (
<bibRefCitation author="Travouillon" box="[691,988,622,647]" etAl="et al." firstAuthor="Travouillon" pageId="211" pageNumber="211" pagination="375 - 382" refId="ref230665" refString="Travouillon, K. J., S. J. Hand, M. Archer, and K. H. Black. 2014 a. Earliest modern bandicoot and bilby (Marsupialia, Peramelidae and Thylacomyidae) from the Miocene of the Riversleigh World Heritage Area, northwestern Queensland, Australia. Journal of Vertebrate Paleontology 34 (2): 375 - 382." type="journal article" year="2014" yearSuffix="a">Travouillon et al., 2014a</bibRefCitation>
): †
<taxonomicName authorityName="Travouillon, Hand, Archer &amp; Black" authorityYear="2014" box="[1026,1212,622,646]" class="Mammalia" family="Peramelidae" genus="Crash" higherTaxonomySource="GBIF" kingdom="Animalia" order="Peramelemorphia" pageId="211" pageNumber="211" phylum="Chordata" rank="species">
<emphasis box="[1026,1212,622,646]" italics="true" pageId="211" pageNumber="211">Crash bandicoot</emphasis>
</taxonomicName>
(which appears to be a peramelid), and †
<taxonomicName authorityName="Travouillon, Hand, Archer &amp; Black" authorityYear="2014" class="Mammalia" family="Thylacomyidae" genus="Liyamayi" higherTaxonomySource="GBIF" kingdom="Animalia" order="Peramelemorphia" pageId="211" pageNumber="211" phylum="Chordata" rank="species">
<emphasis italics="true" pageId="211" pageNumber="211">Liyamayi dayi</emphasis>
</taxonomicName>
(which was described as a thylacomyid, although its position in published phylogenies is highly labile;
<bibRefCitation author="Travouillon" etAl="et al." firstAuthor="Travouillon" pageId="211" pageNumber="211" pagination="375 - 382" refId="ref230665" refString="Travouillon, K. J., S. J. Hand, M. Archer, and K. H. Black. 2014 a. Earliest modern bandicoot and bilby (Marsupialia, Peramelidae and Thylacomyidae) from the Miocene of the Riversleigh World Heritage Area, northwestern Queensland, Australia. Journal of Vertebrate Paleontology 34 (2): 375 - 382." type="journal article" year="2014" yearSuffix="a">
Travouillon et al., 2014
<bibRefCitation author="a" box="[741,837,788,812]" firstAuthor="a" pageId="211" pageNumber="211" pagination="1408" refId="ref237143" refString="Yates, A. M. 2015 b. New craniodental remains of Wakaleo alcootaensis (Diprotodontia: Thylacoleonidae) a carnivorous marsupial from the late Miocene Alcoota Local Fauna of the Northern Territory, Australia. PeerJ 3: e 1408." type="journal article" year="2015" yearSuffix="b">
a, 2015
<bibRefCitation author="b" box="[823,907,788,812]" firstAuthor="b" pageId="211" pageNumber="211" pagination="1408" refId="ref237143" refString="Yates, A. M. 2015 b. New craniodental remains of Wakaleo alcootaensis (Diprotodontia: Thylacoleonidae) a carnivorous marsupial from the late Miocene Alcoota Local Fauna of the Northern Territory, Australia. PeerJ 3: e 1408." type="journal article" year="2021" yearSuffix="b">b, 2021</bibRefCitation>
</bibRefCitation>
</bibRefCitation>
;
<bibRefCitation author="Chamberlain, P. M. &amp; K. J. Travouillon &amp; M. Archer &amp; S. J. Hand" box="[921,1208,788,812]" pageId="211" pageNumber="211" pagination="197 - 206" refId="ref197024" refString="Chamberlain, P. M., K. J. Travouillon, M. Archer, and S. J. Hand. 2015. Kutjamarcoot brevirostrum gen. et sp. nov., a new short-snouted, early Miocene bandicoot (Marsupialia: Peramelemorphia) from the Kutjamarpu Local Fauna (Wipajiri Formation) in South Australia. Alcheringa: an Australasian Journal of Palaeontology 40 (2): 197 - 206." type="journal article" year="2015">Chamberlain et al., 2015</bibRefCitation>
;
<bibRefCitation author="Kear, B. P. &amp; K. P. Aplin &amp; M. Westerman" box="[684,873,821,845]" pageId="211" pageNumber="211" pagination="37537" refId="ref210965" refString="Kear, B. P., K. P. Aplin, and M. Westerman. 2016. Bandicoot fossils and DNA elucidate lineage antiquity amongst xeric-adapted Australasian marsupials. Scientific Reports 6: 37537." type="journal article" year="2016">Kear et al., 2016</bibRefCitation>
). If †
<taxonomicName authorityName="Travouillon, Hand, Archer &amp; Black" authorityYear="2014" box="[932,995,821,845]" class="Mammalia" family="Peramelidae" genus="Crash" higherTaxonomySource="GBIF" kingdom="Animalia" order="Peramelemorphia" pageId="211" pageNumber="211" phylum="Chordata" rank="genus">
<emphasis box="[932,995,821,845]" italics="true" pageId="211" pageNumber="211">Crash</emphasis>
</taxonomicName>
and †
<taxonomicName authorityName="Travouillon, Hand, Archer &amp; Black" authorityYear="2014" box="[1068,1170,822,845]" class="Mammalia" family="Thylacomyidae" genus="Liyamayi" higherTaxonomySource="GBIF" kingdom="Animalia" order="Peramelemorphia" pageId="211" pageNumber="211" phylum="Chordata" rank="genus">
<emphasis box="[1068,1170,822,845]" italics="true" pageId="211" pageNumber="211">Liyamayi</emphasis>
</taxonomicName>
are indeed early perameloids, then their middle Miocene age is broadly congruent with our middle to early Miocene age estimate for the onset of diversfication of Perameloidea.
</paragraph>
</subSubSection>
</treatment>
</document>