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<taxonomicName id="4C4617C8F6F668E4DAD7FC64186FFC25" authority="Szalay, 1982" authorityName="Szalay" authorityYear="1982" box="[220,525,970,994]" genus="Australidelphia" kingdom="Animalia" pageId="205" pageNumber="205" rank="genus">
Australidelphia
<bibRefCitation id="EFD711BAF6F668E4DB80FC64186FFC25" author="Szalay, F. S." box="[395,525,970,994]" pageId="205" pageNumber="205" pagination="621 - 640" refId="ref228988" refString="Szalay, F. S. 1982 a. A new appraisal of marsupial phylogeny and classification. In M. Archer (editor), Carnivorous marsupials: 621 - 640. Mosman, New South Wales: Royal Zoological Society of New South Wales." type="book chapter" year="1982">Szalay, 1982</bibRefCitation>
</taxonomicName>
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<paragraph id="8BF96C4BF6F668E4DA87FC551B57FB92" blockId="205.[108,636,1019,1738]" pageId="205" pageNumber="205">
CONTENTS:
<taxonomicName id="4C4617C8F6F668E4DB1BFC551BAEFBD4" baseAuthorityName="Kealy and Beck" baseAuthorityYear="2017" box="[272,460,1019,1043]" class="Mammalia" kingdom="Animalia" order="Dasyuromorphia" pageId="205" pageNumber="205" phylum="Chordata" rank="order">Dasyuromorphia</taxonomicName>
,
<taxonomicName id="4C4617C8F6F668E4DBD2FC55181AFBD4" authorityName="Owen" authorityYear="1866" box="[473,632,1019,1043]" class="Mammalia" kingdom="Animalia" order="Diprotodontia" pageId="205" pageNumber="205" phylum="Chordata" rank="order">Diprotodontia</taxonomicName>
,
<taxonomicName id="4C4617C8F6F668E4DA67FBB21B44FBF3" authorityName="Ameghino" authorityYear="1889" box="[108,294,1052,1076]" class="Mammalia" kingdom="Animalia" order="Microbiotheria" pageId="205" pageNumber="205" phylum="Chordata" rank="order">Microbiotheria</taxonomicName>
,
<taxonomicName id="4C4617C8F6F668E4DB42FBB2184EFBF3" authorityName="Kirsch" authorityYear="1977" box="[329,556,1052,1076]" class="Mammalia" kingdom="Animalia" order="Notoryctemorphia" pageId="205" pageNumber="205" phylum="Chordata" rank="order">Notoryctemorphia</taxonomicName>
, and
<taxonomicName id="4C4617C8F6F668E4DA67FB931B53FB92" baseAuthorityName="Travouillon and Phillips" baseAuthorityYear="2018" box="[108,305,1085,1109]" class="Mammalia" kingdom="Animalia" order="Peramelemorphia" pageId="205" pageNumber="205" phylum="Chordata" rank="order">Peramelemorphia</taxonomicName>
.
</paragraph>
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<subSubSection id="C35C3FC0F6F668E4DA87FBF11A98FADC" pageId="205" pageNumber="205" type="description">
<paragraph id="8BF96C4BF6F668E4DA87FBF11ACEFB5F" blockId="205.[108,636,1019,1738]" pageId="205" pageNumber="205">STEM AGE: 55.1 Mya (95% HPD: 54.656.6 Mya).</paragraph>
<paragraph id="8BF96C4BF6F668E4DA87FB0F1ACFFB1D" blockId="205.[108,636,1019,1738]" pageId="205" pageNumber="205">CROWN AGE: 48.0 Mya (95% HPD: 44.350.9 Mya).</paragraph>
<paragraph id="8BF96C4BF6F668E4DA87FB4D1A98FADC" blockId="205.[108,636,1019,1738]" pageId="205" pageNumber="205">UNAMBIGUOUS CRANIODENTAL SYNAPOMORPHIES: None.</paragraph>
</subSubSection>
<subSubSection id="C35C3FC0F6F668E7DA87FA8B1E66FA67" lastPageId="206" lastPageNumber="206" pageId="205" pageNumber="205" type="discussion">
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COMMENTS: Recent phylogenetic analyses based on molecular data (e.g.,
<bibRefCitation id="EFD711BAF6F668E4DBD0FAE81AC1FAB8" author="Phillips" etAl="et al." firstAuthor="Phillips" pageId="205" pageNumber="205" pagination="1533 - 1538" refId="ref221252" refString="Phillips, M. J., Y. H. Lin, G. L. Harrison, and D. Penny. 2001. Mitochondrial genomes of a bandicoot and a brushtail possum confirm the monophyly of australidelphian marsupials. Proceedings of the Royal Society of London B, Biological Sciences 268 (1475): 1533 - 1538." type="journal article" year="2001">Phillips et al., 2001</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F668E4DABAFAC91B9CFAB8" author="Amrine-Madsen, H." box="[177,510,1383,1407]" pageId="205" pageNumber="205" pagination="186 - 196" refId="ref188949" refString="Amrine-Madsen, H., et al. 2003 b. Nuclear gene sequences provide evidence for the monophyly of australidelphian marsupials. Molecular Phylogenetics and Evolution 28 (2): 186 - 196." type="journal article" year="2003">Amrine-Madsen et al., 2003b</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F668E4D800FAC91AA8FA67" author="Nilsson" etAl="et al." firstAuthor="Nilsson" pageId="205" pageNumber="205" pagination="189 - 196" refId="ref219362" refString="Nilsson, M. A., U. Arnason, P. B. S. Spencer, and A. Janke. 2004. Marsupial relationships and a timeline for marsupial radiation in South Gondwana. Gene 340: 189 - 196." type="journal article" year="2004">Nilsson et al., 2004</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F668E4DADEFA261BCBFA67" author="Phillips" box="[213,425,1416,1440]" etAl="et al." firstAuthor="Phillips" pageId="205" pageNumber="205" pagination="122 - 137" refId="ref221314" refString="Phillips, M. J., P. A. McLenachan, C. Down, G. C. Gibb, and D. Penny. 2006. Combined mitochondrial and nuclear DNA sequences resolve the interrelations of the major Australasian marsupial radiations. Systematic Biology 55 (1): 122 - 137." type="journal article" year="2006">Phillips et al., 2006</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F668E4DBBFFA261858FA67" author="Beck, R. M. D." box="[436,570,1416,1440]" pageId="205" pageNumber="205" pagination="175 - 189" refId="ref192516" refString="Beck, R. M. D. 2008 a. A dated phylogeny of marsupials using a molecular supermatrix and multiple fossil constraints. Journal of Mammalogy 89 (1): 175 - 189." type="journal article" year="2008">Beck, 2008a</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F668E4D84DFA271B59FA06" author="Meredith, R. W. &amp; M. Westerman &amp; J. A. Case &amp; M. S. Springer" pageId="205" pageNumber="205" pagination="1 - 36" refId="ref216587" refString="Meredith, R. W., M. Westerman, J. A. Case, and M. S. Springer. 2008 b. A phylogeny and timescale for marsupial evolution based on sequences for five nuclear genes. Journal of Mammalian Evolution 15 (1): 1 - 36." type="journal article" year="2008">Meredith et al., 2008b</bibRefCitation>
,
<bibRefCitation id="EFD711BAF6F668E4DB41FA071BEEFA06" author="Meredith, R. W. &amp; C. Krajewski &amp; M. Westerman &amp; M. S. Springer" box="[330,396,1449,1473]" pageId="205" pageNumber="205" pagination="383 - 406" refId="ref216745" refString="Meredith, R. W., C. Krajewski, M. Westerman, and M. S. Springer. 2009 c. Relationships and divergence times among the orders and families of Marsupialia. Museum of Northern Arizona Bulletin 65: 383 - 406." type="journal article" year="2009">
2009
<bibRefCitation id="EFD711BAF6F668E4DB8AFA071BEEFA06" author="Rodgers, J. C." box="[385,396,1449,1473]" pageId="205" pageNumber="205" refId="ref224441" refString="Rodgers, J. C. 2011. Comparative morphology of the vestibular semicircular canals in therian mammals. Ph. D. dissertation, Faculty of the Graduate School, University of Texas at Austin, Austin." type="book" year="2011">c</bibRefCitation>
</bibRefCitation>
, 2011;
<bibRefCitation id="EFD711BAF6F668E4DBEBFA071AC1FA25" author="Nilsson" etAl="et al." firstAuthor="Nilsson" pageId="205" pageNumber="205" pagination="1000436" refId="ref219407" refString="Nilsson, M. A., et al. 2010. Tracking marsupial evolution using archaic genomic retroposon insertions. PLoS Biology 8 (7): e 1000436." type="journal article" year="2010">Nilsson et al., 2010</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F668E4DABBFA641BF0FA25" author="Mitchell, K. J." box="[176,402,1482,1506]" pageId="205" pageNumber="205" pagination="2322 - 2330" refId="ref217043" refString="Mitchell, K. J., et al. 2014. Molecular phylogeny, biogeography, and habitat preference evolution of marsupials. Molecular Biology and Evolution 31 (9): 2322 - 2330." type="journal article" year="2014">Mitchell et al., 2014</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F668E4DB94FA64181AFA25" author="Gallus, S. &amp; A. Janke &amp; V. Kumar &amp; M. A. Nilsson" box="[415,632,1482,1506]" pageId="205" pageNumber="205" pagination="985 - 992" refId="ref204175" refString="Gallus, S., A. Janke, V. Kumar, and M. A. Nilsson. 2015 a. Disentangling the relationship of the Australian marsupial orders using retrotransposon and evolutionary network analyses. Genome Biology and Evolution 7 (4): 985 - 992." type="journal article" year="2015">Gallus et al., 2015a</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F668E4DA67FA451B39F9C4" author="Duchene, D. A." box="[108,347,1515,1539]" pageId="205" pageNumber="205" pagination="400 - 412" refId="ref200659" refString="Duchene, D. A., et al. 2018. Analysis of phylogenomic tree space resolves relationships among marsupial families. Systematic Biology 67 (3): 400 - 412." type="journal article" year="2018">Duchêne et al., 2018</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F668E4DB61FA421AC0F9E3" author="Alvarez-Carretero, S." pageId="205" pageNumber="205" pagination="263 - 267" refId="ref188636" refString="Alvarez-Carretero, S., et al. 2021. A species-level timeline of mammal evolution integrating phylogenomic data. Nature 602: 263 - 267." type="journal article" year="2021">Álvarez-Carretero et al., 2021</bibRefCitation>
) and others based on total-evidence datasets (e.g.,
<bibRefCitation id="EFD711BAF6F668E4DADFF9831BFCF982" author="Asher, R. J. &amp; I. Horovitz &amp; M. R. Sanchez-Villagra" box="[212,414,1581,1605]" pageId="205" pageNumber="205" pagination="240 - 250" refId="ref191802" refString="Asher, R. J., I. Horovitz, and M. R. Sanchez-Villagra. 2004. First combined cladistic analysis of marsupial mammal interrelationships. Molecular Phylogenetics and Evolution 33: 240 - 250." type="journal article" year="2004">Asher et al., 2004</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F668E4DBA6F983181AF982" author="Beck, R. M. D. &amp; H. Godthelp &amp; V. Weisbecker &amp; M. Archer &amp; S. J. Hand" box="[429,632,1581,1605]" pageId="205" pageNumber="205" pagination="1858" refId="ref192866" refString="Beck, R. M. D., H. Godthelp, V. Weisbecker, M. Archer, and S. J. Hand. 2008 a. Australia's oldest marsupial fossils and their biogeographical implications. PLoS One 3 (3): e 1858." type="journal article" year="2008">
Beck et al., 2008
<bibRefCitation id="EFD711BAF6F668E4D867F983181AF982" author="Rambaut, A. &amp; M. A. Suchard &amp; D. Xie &amp; A. J. Drummond" box="[620,632,1581,1605]" pageId="205" pageNumber="205" refId="ref223023" refString="Rambaut, A., M. A. Suchard, D. Xie, and A. J. Drummond. 2014. Tracer v 1.6." type="book" year="2014">a</bibRefCitation>
</bibRefCitation>
, 2014,
<bibRefCitation id="EFD711BAF6F668E4DAB8F9E01A88F9A1" author="Beck, R. M. D. &amp; N. M. Warburton &amp; M. Archer &amp; S. J. Hand &amp; K. P. Aplin" box="[179,234,1614,1638]" pageId="205" pageNumber="205" pagination="151 - 171" refId="ref193033" refString="Beck, R. M. D., N. M. Warburton, M. Archer, S. J. Hand, and K. P. Aplin. 2016. Going underground: postcranial morphology of the early Miocene marsupial mole Naraboryctes philcreaseri and the evolution of fossoriality in notoryctemorphians. Memoirs of Museum Victoria 74: 151 - 171." type="journal article" year="2016">2016</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F668E4DAF1F9E01B8BF9A0" author="Horovitz, I." box="[250,489,1614,1639]" pageId="205" pageNumber="205" pagination="8278" refId="ref209439" refString="Horovitz, I., et al. 2009. Cranial anatomy of the earliest marsupials and the origin of opossums. PLoS One 4 (12): e 8278." type="journal article" year="2009">Horovitz et al., 2009</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F668E4DBF2F9E0181AF9A0" author="Beck, R. M. D." box="[505,632,1614,1639]" pageId="205" pageNumber="205" pagination="715 - 729" refId="ref192633" refString="Beck, R. M. D. 2012. An ' ameridelphian' marsupial from the early Eocene of Australia supports a complex model of Southern Hemisphere marsupial biogeography. Naturwissenschaften 99 (9): 715 - 729." type="journal article" year="2012">Beck, 2012</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F668E4DA67F9DE1B02F94F" author="Maga, A. M. &amp; R. M. D. Beck" box="[108,352,1647,1672]" pageId="205" pageNumber="205" pagination="0181712" refId="ref214955" refString="Maga, A. M., and R. M. D. Beck. 2017. Skeleton of an unusual, cat-sized marsupial relative (Metatheria: Marsupialiformes) from the middle Eocene (Lutetian: 44 - 43 million years ago) of Turkey. PLoS One 12 (8): e 0181712." type="journal article" year="2017">Maga and Beck, 2017</bibRefCitation>
) have consistently recovered monophyly of Australidelphia, and australidelphian monophyly has also been found in most (e.g.,
<bibRefCitation id="EFD711BAF6F668E4D92EFF4C1EDAFF3D" author="Horovitz, I. &amp; M. R. Sanchez-Villagra" box="[805,1208,226,250]" pageId="205" pageNumber="205" pagination="181 - 212" refId="ref209364" refString="Horovitz, I., and M. R. Sanchez-Villagra. 2003. A morphological analysis of marsupial mammal higherlevel phylogenetic relationships. Cladistics 19: 181 - 212." type="journal article" year="2003">Horovitz and Sánchez-Villagra, 2003</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F668E4D8A6FEAD199EFEDC" author="Sanchez-Villagra" box="[685,1020,259,283]" etAl="et al." firstAuthor="Sanchez-Villagra" pageId="205" pageNumber="205" pagination="83 - 97" refId="ref226509" refString="Schmelzle, T., M. R. Sanchez-Villagra, and W. Maier. 2007. Vestibular labyrinth diversity in diprotodontian marsupial mammals. Mammal Study 32: 83 - 97." type="journal article" year="2007">Sánchez-Villagra et al., 2007</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F668E4DE06FEAD1881FEFB" author="Horovitz, I." pageId="205" pageNumber="205" pagination="109 - 141" refId="ref209395" refString="Horovitz, I., et al. 2008. The anatomy of Herpetotherium cf. fugax Cope, 1873, a metatherian from the Oligocene of North America. Palaeontographica Abteilung A 284 (4 - 6): 109 - 141." type="journal article" year="2008">Horovitz et al., 2008</bibRefCitation>
,
<bibRefCitation id="EFD711BAF6F668E4D8F9FE8A194BFEFB" author="Horovitz, I." box="[754,809,292,316]" pageId="205" pageNumber="205" pagination="8278" refId="ref209439" refString="Horovitz, I., et al. 2009. Cranial anatomy of the earliest marsupials and the origin of opossums. PLoS One 4 (12): e 8278." type="journal article" year="2009">2009</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F668E4D93CFE8A1E75FEFB" author="Lorente, M. &amp; L. Chornogubsky &amp; F. J. Goin" box="[823,1047,292,316]" pageId="205" pageNumber="205" pagination="533 - 547" refId="ref213633" refString="Lorente, M., L. Chornogubsky, and F. J. Goin. 2016. On the existence of non-microbiotherian australidelphian marsupials (Diprotodontia) in the Eocene of Patagonia. Palaeontology 59 (4): 533 - 547." type="journal article" year="2016">Lorente et al., 2016</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F668E4DE2DFE8A1937FE9A" author="Carneiro, L. M. &amp; E. V. Oliveira" pageId="205" pageNumber="205" pagination="355 - 372" refId="ref196481" refString="Carneiro, L. M., and E. V. Oliveira. 2017 a. Systematic affinities of the extinct metatherian Eobrasilia coutoi Simpson, 1947, a South American early Eocene Stagodontidae: implications for &quot; Eobrasiliinae. &quot; Revista Brasileira de Paleontologia 20 (3): 355 - 372." type="journal article" year="2017">Carneiro and Oliveira, 2017a</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F668E4D969FEEB1E2AFE9A" author="Carneiro, L. M. &amp; E. V. Oliveira &amp; F. J. Goin" box="[866,1096,325,349]" pageId="205" pageNumber="205" pagination="120 - 131" refId="ref196583" refString="Carneiro, L. M., E. V. Oliveira, and F. J. Goin. 2018. Austropediomys marshalli gen. et sp. nov., a new Pediomyoidea (Mammalia, Metatheria) from the Paleogene of Brazil: paleobiogeographic implications. Revista Brasileira de Paleontologia 21 (2): 120 - 131." type="journal article" year="2018">Carneiro et al., 2018</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F668E4DE5EFEEB1881FEB9" author="Carneiro, L. M." pageId="205" pageNumber="205" pagination="20180440" refId="ref196433" refString="Carneiro, L. M. 2019. A new protodidelphid (Mammalia, Marsupialia, Didelphimorphia) from the Itaborai Basin and its implications for the evolution of the Protodidelphidae. Anais da Academia Brasileira de Ciencias 91 (Suppl. 2): e 20180440." type="journal article" year="2019">Carneiro, 2019</bibRefCitation>
) but not all (e.g.,
<bibRefCitation id="EFD711BAF6F668E4D9B0FEC81881FE58" author="Ladeveze, S. &amp; C. de Muizon" pageId="205" pageNumber="205" pagination="746 - 784" refId="ref212638" refString="Ladeveze, S., and C. de Muizon. 2010. Evidence of early evolution of Australidelphia (Metatheria, Mammalia) in South America: phylogenetic relationships of the metatherians from the late Palaeocene of Itaborai (Brazil) based on teeth and petrosal bones. Zoological Journal of the Linnean Society 159 (3): 746 - 784." type="journal article" year="2010">Ladevèze and Muizon, 2010</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F668E4D8E4FE2919A2FE58" author="Wilson" box="[751,960,391,415]" etAl="et al." firstAuthor="Wilson" pageId="205" pageNumber="205" pagination="13734" refId="ref234583" refString="Wilson, G. P., E. G. Ekdale, J. W. Hoganson, J. J. Calede, and A. Vander Linden. 2016. A large carnivorous mammal from the Late Cretaceous and the North American origin of marsupials. Nature Communications 7: 13734." type="journal article" year="2016">Wilson et al., 2016</bibRefCitation>
) recent morphological analyses. This overall pattern is confirmed here: our morphological analyses did not recover australidelphian monophyly (figs. 30, 31), whereas it was strongly supported in all our molecular (figs. 2729) and total evidence (figs. 32,
<figureCitation id="137D70CEF6F668E4DE39FD821E2CFD83" box="[1074,1102,556,580]" 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="205" pageNumber="205">33</figureCitation>
) analyses. Given that
<bibRefCitation id="EFD711BAF6F668E4D921FDE319ABFDA2" author="Szalay" box="[810,969,589,613]" firstAuthor="Szalay" pageId="205" pageNumber="205" pagination="621 - 640" refId="ref228988" refString="Szalay, F. S. 1982 a. A new appraisal of marsupial phylogeny and classification. In M. Archer (editor), Carnivorous marsupials: 621 - 640. Mosman, New South Wales: Royal Zoological Society of New South Wales." type="book chapter" year="1982" yearSuffix="a">Szalay (1982a)</bibRefCitation>
first proposed monophyly of Australidelphia based primarily on shared derived features of the tarsus (specifically the presence of a continuous lower ankle joint, and a tripartite calcaneocuboid facet; see also
<bibRefCitation id="EFD711BAF6F668E4D8A7FD5D1956FCCC" author="Szalay" box="[684,820,755,779]" firstAuthor="Szalay" pageId="205" pageNumber="205" refId="ref229131" refString="Szalay, F. S. 1994. Evolutionary history of the marsupials and an analysis of osteological characters, Cambridge: Cambridge University Press." type="book" year="1994">Szalay, 1994</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F668E4D94AFD5D19DDFCCC" author="Beck, R. M. D." box="[833,959,755,779]" pageId="205" pageNumber="205" pagination="715 - 729" refId="ref192633" refString="Beck, R. M. D. 2012. An ' ameridelphian' marsupial from the early Eocene of Australia supports a complex model of Southern Hemisphere marsupial biogeography. Naturwissenschaften 99 (9): 715 - 729." type="journal article" year="2012">Beck, 2012</bibRefCitation>
), the addition of postcranial characters to our craniodental matrix may ultimately result in morphological support for Australidelphia (as in Horovitz and SánchezVillagra, 2003).
</paragraph>
<paragraph id="8BF96C4BF6F668E7D8C6FC371BD9FEDC" blockId="205.[684,1213,226,1738]" lastBlockId="206.[108,637,226,1738]" lastPageId="206" lastPageNumber="206" pageId="205" pageNumber="205">
No craniodental feature optimizes as an unambiguous synapomorphy of Australidelphia in our dated total-evidence analysis, but three optimize as synapomorphies under Accelerated Transformation—extracranial course of mandibular nerve fully enclosed by medial outgrowths of the auditory bulla (char. 52: 0→1; ci = 0.231); posterior limb of ectotympanic in contact with, but suturally distinct from, pars canalicularis of the petrosal and/or posttympanic process of the squamosal (char. 60: 0→1; ci = 0.333); and prootic canal foramen on tympanic face of petrosal absent (char. 69: 0→1; ci = 0.083)—but all three traits show high levels of homoplasy and undergo subsequent reversals within Australidelphia (see file S
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the online supplement). Our failure to identify compelling craniodental synapomorphies for Australidelphia likely reflects the fact that the ancestral australidelphian probably had a relatively generalized cranium and dentition that was little different from the plesiomorphic marsupial condition (see
<bibRefCitation id="EFD711BAF6F668E4D944F9C11E6BF940" author="Szalay" box="[847,1033,1647,1672]" firstAuthor="Szalay" page="346" pageId="205" pageNumber="205" refId="ref229131" refString="Szalay, F. S. 1994. Evolutionary history of the marsupials and an analysis of osteological characters, Cambridge: Cambridge University Press." type="book" year="1994">Szalay, 1994: 346</bibRefCitation>
) and that different lineages within Australidelphia subsequently evolved very disparate apomorphies of the dentition, cranium, or both (see also comments by
<bibRefCitation id="EFD711BAF6F568E7DAD2FEAD1BCDFEDC" author="Archer, M." box="[217,431,259,283]" pageId="206" pageNumber="206" 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">Archer, 1984c: 782</bibRefCitation>
).
</paragraph>
<paragraph id="8BF96C4BF6F568E7DA86FE8A1BDCFC8A" blockId="206.[108,637,226,1738]" pageId="206" pageNumber="206">
Our estimate for the first split within Australidelphia is in the early to middle Eocene. This is younger than the early or middle Paleocene †
<taxonomicName id="4C4617C8F6F568E7DAB2FE291BC0FE58" authorityName="Marshall &amp; de Muizon" authorityYear="1988" box="[185,418,391,415]" class="Mammalia" family="Microbiotheriidae" genus="Khasia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Microbiotheria" pageId="206" pageNumber="206" phylum="Chordata" rank="species" species="cordillerensis">
<emphasis id="B932B059F6F568E7DAB2FE291BC0FE58" box="[185,418,391,415]" italics="true" pageId="206" pageNumber="206">Khasia cordillerensis</emphasis>
</taxonomicName>
, a taxon that was originally described as a microbiotherian (
<bibRefCitation id="EFD711BAF6F568E7D848FE061B11FE26" author="Marshall, L. G. &amp; C. de Muizon" pageId="206" pageNumber="206" pagination="23 - 55" refId="ref215460" refString="Marshall, L. G., and C. de Muizon. 1988. The dawn of the age of mammals in South America. National Geographic Research 4 (1): 23 - 55." type="journal article" year="1988">Marshall and Muizon, 1988</bibRefCitation>
; see also
<bibRefCitation id="EFD711BAF6F568E7DBD7FE671815FE26" author="Muizon, C. de" box="[476,631,457,481]" pageId="206" pageNumber="206" pagination="575 - 624" refId="ref217663" refString="Muizon, C. de. 1991. La fauna de mamiferos de Tiupampa (Paleoceno Inferior, Formacion Santa Lucia), Bolivia. In R. Suarez-Soruco (editor), Fosiles y facies de Bolivia, vol. 1. Vertebrados: 575 - 624. Santa Cruz, Bolivia: Revista Technica de Yacimientos Petroliferos Fiscales Bolivianos." type="book chapter" year="1991">Muizon, 1991</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F568E7DA66FE441B4CFDC5" author="Cozzuol, M. A. &amp; F. Goin &amp; M. De Los Reyes &amp; A. Ranzi" box="[109,302,490,514]" pageId="206" pageNumber="206" pagination="663 - 667" refId="ref198725" refString="Cozzuol, M. A., F. Goin, M. De Los Reyes, and A. Ranzi. 2006. The oldest species of Didelphis (Mammalia, Marsupialia, Didelphidae), from the late Miocene of Amazonia. Journal of Mammalogy 87 (4): 663 - 667." type="journal article" year="2006">Goin et al., 2006</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F568E7DB30FE44187BFDC5" author="Muizon, C. de &amp; S. Ladeveze &amp; C. Selva &amp; R. Vignaud &amp; F. Goussard" box="[315,537,490,514]" pageId="206" pageNumber="206" pagination="363 - 459" refId="ref218213" refString="Muizon, C. de, S. Ladeveze, C. Selva, R. Vignaud, and F. Goussard. 2018. Allqokirus australis (Sparassodonta, Metatheria) from the early Palaeocene of Tiupampa (Bolivia) and the rise of the metatherian carnivorous radiation in South America. Geodiversitas 40 (16): 363 - 459." type="journal article" year="2018">Muizon et al., 2018</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F568E7D82CFE441B2CFDE4" author="Ladeveze, S. &amp; C. Selva &amp; C. de Muizon" pageId="206" pageNumber="206" pagination="1463 - 1479" refId="ref212907" refString="Ladeveze, S., C. Selva, and C. de Muizon. 2020. What are &quot; opossum-like &quot; fossils? The phylogeny of herpetotheriid and peradectid metatherians, based on new features from the petrosal anatomy. Journal of Systematic Palaeontology 18 (17): 1463 - 1479." type="journal article" year="2020">Muizon and Ladevèze, 2020</bibRefCitation>
) and hence a crown-clade australidelphian. However, as noted above, several subsequent authors have argued that †
<taxonomicName id="4C4617C8F6F568E7D84EFDE31AE8FD41" class="Mammalia" family="Microbiotheriidae" genus="Khasia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Microbiotheria" pageId="206" pageNumber="206" phylum="Chordata" rank="genus">
<emphasis id="B932B059F6F568E7D84EFDE31AE8FD41" italics="true" pageId="206" pageNumber="206">Khasia</emphasis>
</taxonomicName>
is more likely a “pediomyoid” (
<bibRefCitation id="EFD711BAF6F568E7DBE4FDC01A88FD60" author="Oliveira and Goin" firstAuthor="Oliveira" pageId="206" pageNumber="206" pagination="299 - 320" refId="ref220101" refString="Oliveira, E. V., and F. J. Goin. 2006. Marsupiais do inicio do Terciario do Brasil: origem, irradiacao e historia biogeografica. In N. C. Caceres and E. L. A. Monteiro Filho (editors), Os Marsupiais do Brasil: biologia, ecologia e evolucao: 299 - 320. Campo Grande: UFMS." type="book chapter" year="2006">Oliveira and Goin, 2006</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F568E7DAFFFD211BCEFD60" author="Goin, F. J. &amp; M. O. Woodburne &amp; A. N. Zimicz &amp; G. M. Martin &amp; L. Chornogubsky" box="[244,428,655,679]" pageId="206" pageNumber="206" refId="ref207099" refString="Goin, F. J., M. O. Woodburne, A. N. Zimicz, G. M. Martin, and L. Chornogubsky. 2016. A brief history of South American metatherians: Evolutionary contexts and intercontinental dispersals. Dordrecht: Springer." type="book" year="2016">Goin et al., 2016</bibRefCitation>
), a hypothesis supported by the morphological phylogenetic analysis of
<bibRefCitation id="EFD711BAF6F568E7DAB5FD7C1BDAFD2D" author="Carneiro, L. M. &amp; E. V. Oliveira &amp; F. J. Goin" box="[190,440,722,746]" pageId="206" pageNumber="206" pagination="120 - 131" refId="ref196583" refString="Carneiro, L. M., E. V. Oliveira, and F. J. Goin. 2018. Austropediomys marshalli gen. et sp. nov., a new Pediomyoidea (Mammalia, Metatheria) from the Paleogene of Brazil: paleobiogeographic implications. Revista Brasileira de Paleontologia 21 (2): 120 - 131." type="journal article" year="2018">Carneiro et al. (2018)</bibRefCitation>
. †
<taxonomicName id="4C4617C8F6F568E7DBD8FD7C187DFD2D" box="[467,543,722,746]" class="Mammalia" family="Microbiotheriidae" genus="Khasia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Microbiotheria" pageId="206" pageNumber="206" phylum="Chordata" rank="genus">
<emphasis id="B932B059F6F568E7DBD8FD7C187DFD2D" box="[467,543,722,746]" italics="true" pageId="206" pageNumber="206">Khasia</emphasis>
</taxonomicName>
has not been included here because it is represented only by dental specimens (
<bibRefCitation id="EFD711BAF6F568E7DBDFFCBA1B6AFC8A" author="Marshall, L. G. &amp; C. de Muizon" pageId="206" pageNumber="206" pagination="23 - 55" refId="ref215460" refString="Marshall, L. G., and C. de Muizon. 1988. The dawn of the age of mammals in South America. National Geographic Research 4 (1): 23 - 55." type="journal article" year="1988">Marshall and Muizon, 1988</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F568E7DB1FFC9B1BD0FC8A" author="Muizon, C. de" box="[276,434,821,845]" pageId="206" pageNumber="206" pagination="575 - 624" refId="ref217663" refString="Muizon, C. de. 1991. La fauna de mamiferos de Tiupampa (Paleoceno Inferior, Formacion Santa Lucia), Bolivia. In R. Suarez-Soruco (editor), Fosiles y facies de Bolivia, vol. 1. Vertebrados: 575 - 624. Santa Cruz, Bolivia: Revista Technica de Yacimientos Petroliferos Fiscales Bolivianos." type="book chapter" year="1991">Muizon, 1991</bibRefCitation>
).
</paragraph>
<paragraph id="8BF96C4BF6F568E7DA86FCF8183CF9C4" blockId="206.[108,637,226,1738]" pageId="206" pageNumber="206">
If †
<taxonomicName id="4C4617C8F6F568E7DABEFCF81B60FCA9" box="[181,258,854,878]" class="Mammalia" family="Microbiotheriidae" genus="Khasia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Microbiotheria" pageId="206" pageNumber="206" phylum="Chordata" rank="genus">
<emphasis id="B932B059F6F568E7DABEFCF81B60FCA9" box="[181,258,854,878]" italics="true" pageId="206" pageNumber="206">Khasia</emphasis>
</taxonomicName>
is discounted, the oldest known australidelphian is probably †
<taxonomicName id="4C4617C8F6F568E7DBC9FCD91AD2FC77" class="Mammalia" family="Marsupialia" genus="Djarthia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="206" pageNumber="206" phylum="Chordata" rank="species" species="murgonensis">
<emphasis id="B932B059F6F568E7DBC9FCD91AD2FC77" italics="true" pageId="206" pageNumber="206">Djarthia murgonensis</emphasis>
</taxonomicName>
from the Tingamarra fossil site in eastern
<collectingCountry id="F3512CDBF6F568E7DA90FC171B67FC16" box="[155,261,953,977]" name="Australia" pageId="206" pageNumber="206">Australia</collectingCountry>
, which has been radiometrically dated as earliest Eocene (~54.6 Mya; (
<bibRefCitation id="EFD711BAF6F568E7D835FC741B53FBD4" author="Godthelp, H. &amp; M. Archer &amp; R. L. Cifelli &amp; S. J. Hand &amp; C. F. Gilkeson" pageId="206" pageNumber="206" pagination="514 - 516" refId="ref205513" refString="Godthelp, H., M. Archer, R. L. Cifelli, S. J. Hand, and C. F. Gilkeson. 1992. Earliest known Australian Tertiary mammal fauna. Nature 356: 514 - 516." type="journal article" year="1992">Godthelp et al., 1992</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F568E7DB4BFC551855FBD4" author="Godthelp, H. &amp; S. Wroe &amp; M. Archer" box="[320,567,1019,1043]" pageId="206" pageNumber="206" pagination="289 - 313" refId="ref205557" refString="Godthelp, H., S. Wroe, and M. Archer. 1999. A new marsupial from the early Eocene Tingamarra Local Fauna of Murgon, southeastern Queensland: a prototypical Australian marsupial? Journal of Mammalian Evolution 6 (3): 289 - 313." type="journal article" year="1999">Godthelp et al., 1999</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F568E7D84EFC551A98FBF3" author="Beck, R. M. D. &amp; H. Godthelp &amp; V. Weisbecker &amp; M. Archer &amp; S. J. Hand" pageId="206" pageNumber="206" pagination="1858" refId="ref192866" refString="Beck, R. M. D., H. Godthelp, V. Weisbecker, M. Archer, and S. J. Hand. 2008 a. Australia's oldest marsupial fossils and their biogeographical implications. PLoS One 3 (3): e 1858." type="journal article" year="2008">Beck et al., 2008a</bibRefCitation>
). We have not included †
<taxonomicName id="4C4617C8F6F568E7D817FBB2181EFBF3" authorityName="Godthelp, Wroe &amp; Archer" authorityYear="1999" box="[540,636,1052,1076]" class="Mammalia" family="Marsupialia" genus="Djarthia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="206" pageNumber="206" phylum="Chordata" rank="genus">
<emphasis id="B932B059F6F568E7D817FBB2181EFBF3" box="[540,636,1052,1076]" italics="true" pageId="206" pageNumber="206">Djarthia</emphasis>
</taxonomicName>
here due to its incompleteness (the only craniodental remains are incomplete dental specimens and isolated petrosals;
<bibRefCitation id="EFD711BAF6F568E7DBCFFBD11AC7FB7E" author="Godthelp, H. &amp; S. Wroe &amp; M. Archer" pageId="206" pageNumber="206" pagination="289 - 313" refId="ref205557" refString="Godthelp, H., S. Wroe, and M. Archer. 1999. A new marsupial from the early Eocene Tingamarra Local Fauna of Murgon, southeastern Queensland: a prototypical Australian marsupial? Journal of Mammalian Evolution 6 (3): 289 - 313." type="journal article" year="1999">Godthelp et al., 1999</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F568E7DABFFB0E1BEAFB7E" author="Beck, R. M. D. &amp; H. Godthelp &amp; V. Weisbecker &amp; M. Archer &amp; S. J. Hand" box="[180,392,1184,1209]" pageId="206" pageNumber="206" pagination="1858" refId="ref192866" refString="Beck, R. M. D., H. Godthelp, V. Weisbecker, M. Archer, and S. J. Hand. 2008 a. Australia's oldest marsupial fossils and their biogeographical implications. PLoS One 3 (3): e 1858." type="journal article" year="2008">Beck et al., 2008a</bibRefCitation>
), but isolated tarsals referred to this taxon exhibit characteristic australidelphian synapomorphies (
<bibRefCitation id="EFD711BAF6F568E7DBF1FB4D1AD5FADB" author="Beck, R. M. D. &amp; H. Godthelp &amp; V. Weisbecker &amp; M. Archer &amp; S. J. Hand" pageId="206" pageNumber="206" pagination="1858" refId="ref192866" refString="Beck, R. M. D., H. Godthelp, V. Weisbecker, M. Archer, and S. J. Hand. 2008 a. Australia's oldest marsupial fossils and their biogeographical implications. PLoS One 3 (3): e 1858." type="journal article" year="2008">Beck et al., 2008a</bibRefCitation>
). However, †
<taxonomicName id="4C4617C8F6F568E7DB63FAAA1BAFFADB" authorityName="Godthelp, Wroe &amp; Archer" authorityYear="1999" box="[360,461,1284,1308]" class="Mammalia" family="Marsupialia" genus="Djarthia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="206" pageNumber="206" phylum="Chordata" rank="genus">
<emphasis id="B932B059F6F568E7DB63FAAA1BAFFADB" box="[360,461,1284,1308]" italics="true" pageId="206" pageNumber="206">Djarthia</emphasis>
</taxonomicName>
falls outside crown-clade Australidelphia in most published analyses (with the notable exception of
<bibRefCitation id="EFD711BAF6F568E7D834FAE81B7CFAB8" author="Maga, A. M. &amp; R. M. D. Beck" pageId="206" pageNumber="206" pagination="0181712" refId="ref214955" refString="Maga, A. M., and R. M. D. Beck. 2017. Skeleton of an unusual, cat-sized marsupial relative (Metatheria: Marsupialiformes) from the middle Eocene (Lutetian: 44 - 43 million years ago) of Turkey. PLoS One 12 (8): e 0181712." type="journal article" year="2017">Maga and Beck, 2017</bibRefCitation>
: fig. 38, in which it is sister to
<taxonomicName id="4C4617C8F6F568E7DA65FA261B5AFA67" baseAuthorityName="Kealy and Beck" baseAuthorityYear="2017" box="[110,312,1416,1440]" class="Mammalia" kingdom="Animalia" order="Dasyuromorphia" pageId="206" pageNumber="206" phylum="Chordata" rank="order">Dasyuromorphia</taxonomicName>
), and its position is unresolved with respect to the crown clade in others (
<bibRefCitation id="EFD711BAF6F568E7DA97FA641B04FA25" author="Beck, R. M. D. &amp; H. Godthelp &amp; V. Weisbecker &amp; M. Archer &amp; S. J. Hand" box="[156,358,1482,1506]" pageId="206" pageNumber="206" pagination="1858" refId="ref192866" refString="Beck, R. M. D., H. Godthelp, V. Weisbecker, M. Archer, and S. J. Hand. 2008 a. Australia's oldest marsupial fossils and their biogeographical implications. PLoS One 3 (3): e 1858." type="journal article" year="2008">
Beck et al., 2008
<bibRefCitation id="EFD711BAF6F568E7DB51FA641B04FA25" author="Rambaut, A. &amp; M. A. Suchard &amp; D. Xie &amp; A. J. Drummond" box="[346,358,1482,1506]" pageId="206" pageNumber="206" refId="ref223023" refString="Rambaut, A., M. A. Suchard, D. Xie, and A. J. Drummond. 2014. Tracer v 1.6." type="book" year="2014">a</bibRefCitation>
</bibRefCitation>
, 2014,
<bibRefCitation id="EFD711BAF6F568E7DBBDFA641B8DFA25" author="Beck, R. M. D. &amp; N. M. Warburton &amp; M. Archer &amp; S. J. Hand &amp; K. P. Aplin" box="[438,495,1482,1506]" pageId="206" pageNumber="206" pagination="151 - 171" refId="ref193033" refString="Beck, R. M. D., N. M. Warburton, M. Archer, S. J. Hand, and K. P. Aplin. 2016. Going underground: postcranial morphology of the early Miocene marsupial mole Naraboryctes philcreaseri and the evolution of fossoriality in notoryctemorphians. Memoirs of Museum Victoria 74: 151 - 171." type="journal article" year="2016">2016</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F568E7DBF1FA641815FA25" author="Beck, R. M. D." box="[506,631,1482,1506]" pageId="206" pageNumber="206" pagination="715 - 729" refId="ref192633" refString="Beck, R. M. D. 2012. An ' ameridelphian' marsupial from the early Eocene of Australia supports a complex model of Southern Hemisphere marsupial biogeography. Naturwissenschaften 99 (9): 715 - 729." type="journal article" year="2012">Beck, 2012</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F568E7DA66FA421B2EF9C4" author="Lorente, M. &amp; L. Chornogubsky &amp; F. J. Goin" box="[109,332,1515,1539]" pageId="206" pageNumber="206" pagination="533 - 547" refId="ref213633" refString="Lorente, M., L. Chornogubsky, and F. J. Goin. 2016. On the existence of non-microbiotherian australidelphian marsupials (Diprotodontia) in the Eocene of Patagonia. Palaeontology 59 (4): 533 - 547." type="journal article" year="2016">Lorente et al., 2016</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F568E7DB53FA421833F9C4" author="Maga, A. M. &amp; R. M. D. Beck" box="[344,593,1515,1540]" pageId="206" pageNumber="206" pagination="0181712" refId="ref214955" refString="Maga, A. M., and R. M. D. Beck. 2017. Skeleton of an unusual, cat-sized marsupial relative (Metatheria: Marsupialiformes) from the middle Eocene (Lutetian: 44 - 43 million years ago) of Turkey. PLoS One 12 (8): e 0181712." type="journal article" year="2017">Maga and Beck, 2017</bibRefCitation>
).
</paragraph>
<paragraph id="8BF96C4BF6F568E7DA87F9A219ECFE58" blockId="206.[108,637,226,1738]" lastBlockId="206.[684,1214,226,1441]" pageId="206" pageNumber="206">
The next-oldest definitive australidelphian remains are isolated tarsals from the La Barda locality in
<collectingCountry id="F3512CDBF6F568E7DAEAF9E01B33F9A1" box="[225,337,1614,1638]" name="Argentina" pageId="206" pageNumber="206">Argentina</collectingCountry>
, which has been radiometrically dated as middle Eocene (between ~48 and 43 Mya;
<bibRefCitation id="EFD711BAF6F568E7DAC3F93E1BF7F96E" author="Tejedor" box="[200,405,1680,1705]" etAl="et al." firstAuthor="Tejedor" pageId="206" pageNumber="206" pagination="1 - 43" refId="ref229849" refString="Tejedor, M. F., et al. 2009. New early Eocene mammalian fauna from western Patagonia, Argentina. American Museum Novitates 3638: 1 - 43." type="journal article" year="2009">Tejedor et al., 2009</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F568E7DB94F93F1813F96F" author="Lorente, M. &amp; L. Chornogubsky &amp; F. J. Goin" box="[415,625,1680,1704]" pageId="206" pageNumber="206" pagination="533 - 547" refId="ref213633" refString="Lorente, M., L. Chornogubsky, and F. J. Goin. 2016. On the existence of non-microbiotherian australidelphian marsupials (Diprotodontia) in the Eocene of Patagonia. Palaeontology 59 (4): 533 - 547." type="journal article" year="2016">Lorente et al., 2016</bibRefCitation>
). The La Barda australidelphian tarsals fell within
<taxonomicName id="4C4617C8F6F568E7D8A5FF4C1928FF3D" authorityName="Owen" authorityYear="1866" box="[686,842,226,250]" class="Mammalia" kingdom="Animalia" order="Diprotodontia" pageId="206" pageNumber="206" phylum="Chordata" rank="order">Diprotodontia</taxonomicName>
(members of which are otherwise known only from
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, New
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, and adjacent islands) in the phylogenetic analysis of
<bibRefCitation id="EFD711BAF6F568E7D8A7FEEB19F7FE9A" author="Lorente, M. &amp; L. Chornogubsky &amp; F. J. Goin" box="[684,917,325,349]" pageId="206" pageNumber="206" pagination="533 - 547" refId="ref213633" refString="Lorente, M., L. Chornogubsky, and F. J. Goin. 2016. On the existence of non-microbiotherian australidelphian marsupials (Diprotodontia) in the Eocene of Patagonia. Palaeontology 59 (4): 533 - 547." type="journal article" year="2016">Lorente et al. (2016)</bibRefCitation>
, but we consider that this biogeographically anomalous relationship warrants further testing.
</paragraph>
<paragraph id="8BF96C4BF6F568E7D8C7FE061E28FB91" blockId="206.[684,1214,226,1441]" pageId="206" pageNumber="206">
The woodburnodontid microbiotherian †
<taxonomicName id="4C4617C8F6F568E7D8B1FE6719C9FE26" box="[698,939,457,481]" class="Mammalia" family="Woodburnodontidae" genus="Woodburnodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Microbiotheria" pageId="206" pageNumber="206" phylum="Chordata" rank="species" species="casei">
<emphasis id="B932B059F6F568E7D8B1FE6719C9FE26" box="[698,939,457,481]" italics="true" pageId="206" pageNumber="206">Woodburnodon casei</emphasis>
</taxonomicName>
from the Cucullaea I Allomember of the La Meseta Formation on Seymour Island, off the Antarctic Penninsula, is a more compelling candidate for the oldest definitive crown-clade australidelphian because it preserves distinctive dental features that are characteristic of microbiotherians (
<bibRefCitation id="EFD711BAF6F568E7DE31FD21188DFD0E" author="Goin, F. J." pageId="206" pageNumber="206" pagination="597 - 603" refId="ref206658" refString="Goin, F. J., et al. 2007 c. New marsupial (Mammalia) from the Eocene of Antarctica, and the origins and affinities of the Microbiotheria. Revista de la Asociacion Geologica Argentina 62 (4): 597 - 603." type="journal article" year="2007">Goin et al., 2007c</bibRefCitation>
), and it has been recovered within
<taxonomicName id="4C4617C8F6F568E7DE7BFD1E196CFD2D" authorityName="Ameghino" authorityYear="1889" class="Mammalia" kingdom="Animalia" order="Microbiotheria" pageId="206" pageNumber="206" phylum="Chordata" rank="order">Microbiotheria</taxonomicName>
in several phylogenetic analyses (
<bibRefCitation id="EFD711BAF6F568E7DE82FD7C19AAFCCC" author="Carneiro, L. M. &amp; E. V. Oliveira" pageId="206" pageNumber="206" pagination="497 - 507" refId="ref196536" refString="Carneiro, L. M., and E. V. Oliveira. 2017 b. The Eocene South American metatherian Zeusdelphys complicatus is not a protodidelphid but a hatcheriform: Paleobiogeographic implications. Acta Palaeontologica Polonica 62 (3): 497 - 507." type="journal article" year="2017">Carneiro and Oliveira, 2017b</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F568E7D9DEFD5D1EDAFCCC" author="Carneiro, L. M. &amp; E. V. Oliveira &amp; F. J. Goin" box="[981,1208,755,779]" pageId="206" pageNumber="206" pagination="120 - 131" refId="ref196583" refString="Carneiro, L. M., E. V. Oliveira, and F. J. Goin. 2018. Austropediomys marshalli gen. et sp. nov., a new Pediomyoidea (Mammalia, Metatheria) from the Paleogene of Brazil: paleobiogeographic implications. Revista Brasileira de Paleontologia 21 (2): 120 - 131." type="journal article" year="2018">Carneiro et al., 2018</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F568E7D8A5FCBA1903FCEB" author="Carneiro, L. M." box="[686,865,788,812]" pageId="206" pageNumber="206" pagination="20180440" refId="ref196433" refString="Carneiro, L. M. 2019. A new protodidelphid (Mammalia, Marsupialia, Didelphimorphia) from the Itaborai Basin and its implications for the evolution of the Protodidelphidae. Anais da Academia Brasileira de Ciencias 91 (Suppl. 2): e 20180440." type="journal article" year="2019">Carneiro, 2019</bibRefCitation>
). The age of the Cucullaea I Allomember has proved controversial (
<bibRefCitation id="EFD711BAF6F568E7DE5DFC9B196FFCA9" author="Crame, J. A." pageId="206" pageNumber="206" pagination="114743" refId="ref198824" refString="Crame, J. A., et al. 2014. The early origin of the Antarctic marine fauna and its evolutionary implications. PLoS One 9 (12): e 114743." type="journal article" year="2014">Crame et al., 2014</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F568E7D910FCF81984FCA9" author="Kemp, D. B." box="[795,998,854,878]" pageId="206" pageNumber="206" pagination="583 - 586" refId="ref211003" refString="Kemp, D. B., et al. 2014. A cool temperate climate on the Antarctic Peninsula through the latest Cretaceous to early Paleogene. Geology 42 (7): 583 - 586." type="journal article" year="2014">Kemp et al., 2014</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F568E7D9FFFCF81EDBFCA9" author="Gelfo, J. N. &amp; G. M. Lopez &amp; S. N. Santillana" box="[1012,1209,854,878]" pageId="206" pageNumber="206" pagination="445 - 455" refId="ref204712" refString="Gelfo, J. N., G. M. Lopez, and S. N. Santillana. 2017. Eocene ungulate mammals from west Antarctica: implications from their fossil record and a new species. Antarctic Science 29 (05): 445 - 455." type="journal article" year="2017">Gelfo et al., 2017</bibRefCitation>
,
<bibRefCitation id="EFD711BAF6F568E7D8A7FCD91880FC48" author="Gelfo, J. N. &amp; F. J. Goin &amp; N. Bauza &amp; M. Reguero" box="[684,738,887,911]" pageId="206" pageNumber="206" pagination="274 - 292" refId="ref204762" refString="Gelfo, J. N., F. J. Goin, N. Bauza, and M. Reguero. 2019. The fossil record of Antarctic land mammals: commented review and hypotheses for future research. Advances in Polar Science 30 (3): 274 - 292." type="journal article" year="2019">2019</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F568E7D8E6FCD919C1FC48" author="Zimicz, A. N. &amp; F. J. Goin" box="[749,931,887,911]" pageId="206" pageNumber="206" pagination="975 - 992" refId="ref237724" refString="Zimicz, A. N., and F. J. Goin. 2020. A reassessment of the genus Groeberia Patterson, 1952 (Mammalia, Metatheria): functional and phylogenetic implications. Journal of Systematic Palaeontology 18 (12): 975 - 992." type="journal article" year="2020">Goin et al., 2020</bibRefCitation>
), but it now appears to be about 40 Mya (
<bibRefCitation id="EFD711BAF6F568E7D945FC361E44FC77" author="Douglas, P. M." box="[846,1062,920,944]" pageId="206" pageNumber="206" pagination="6582 - 6587" refId="ref200562" refString="Douglas, P. M., et al. 2014. Pronounced zonal heterogeneity in Eocene southern high-latitude sea surface temperatures. Proceedings of the National Academy of Sciences of the United States of America 111 (18): 6582 - 6587." type="journal article" year="2014">Douglas et al., 2014</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F568E7DE3BFC36196EFC16" author="Amenabar, C. R. &amp; M. Montes &amp; F. Nozal &amp; S. Santillana" pageId="206" pageNumber="206" pagination="351 - 366" refId="ref188835" refString="Amenabar, C. R., M. Montes, F. Nozal, and S. Santillana. 2019. Dinoflagellate cysts of the La Meseta Formation (middle to late Eocene), Antarctic Peninsula: implications for biostratigraphy, palaeoceanography and palaeoenvironment. Geological Magazine 157 (3): 351 - 366." type="journal article" year="2019">Amenábar et al., 2019</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F568E7D912FC1419B8FC16" author="Mors, T. &amp; M. Reguero &amp; D. Vasilyan" box="[793,986,953,977]" pageId="206" pageNumber="206" pagination="5051" refId="ref217290" refString="Mors, T., M. Reguero, and D. Vasilyan. 2020. First fossil frog from Antarctica: implications for Eocene high latitude climate conditions and Gondwanan cosmopolitanism of Australobatrachia. Scientific Reports 10: 5051." type="journal article" year="2020">Mörs et al., 2020</bibRefCitation>
). This postdates our estimate for the first divergence within Australidelphia (see above) and also for the split between
<taxonomicName id="4C4617C8F6F568E7D8A7FBB21936FBF3" authorityName="Ameghino" authorityYear="1889" box="[684,852,1052,1076]" class="Mammalia" kingdom="Animalia" order="Microbiotheria" pageId="206" pageNumber="206" phylum="Chordata" rank="order">Microbiotheria</taxonomicName>
and
<taxonomicName id="4C4617C8F6F568E7D99DFBB21E54FBF3" authorityName="Owen" authorityYear="1866" box="[918,1078,1052,1076]" class="Mammalia" kingdom="Animalia" order="Diprotodontia" pageId="206" pageNumber="206" phylum="Chordata" rank="order">Diprotodontia</taxonomicName>
(median = 45.6 Mya; 95% HPD: 41.448.8 Mya).
</paragraph>
<paragraph id="8BF96C4BF6F568E7D8C7FBF01E66FA67" blockId="206.[684,1214,226,1441]" pageId="206" pageNumber="206">
Older putative microbiotherians have been reported from the early Eocene (probably 51.4 56.0 Mya;
<bibRefCitation id="EFD711BAF6F568E7D929FB0E1985FB7E" author="Clyde, W. C." box="[802,999,1184,1209]" pageId="206" pageNumber="206" pagination="289 - 306" refId="ref197816" refString="Clyde, W. C., et al. 2014. New age constraints for the Salamanca Formation and lower Rio Chico Group in the western San Jorge Basin, Patagonia, Argentina: implications for Cretaceous-Paleogene extinction recovery and land mammal age correlations. Geological Society of America Bulletin 126 (3 - 4): 289 - 306." type="journal article" year="2014">Clyde et al., 2014</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F568E7D9FFFB0E1892FB1D" author="Woodburne" etAl="et al." firstAuthor="Woodburne" pageId="206" pageNumber="206" pagination="109 - 119" refId="ref235572" refString="Woodburne, M. O., et al. 2014 a. Revised timing of the South American early Paleogene land mammal ages. Journal of South American Earth Sciences 54: 109 - 119." type="journal article" year="2014" yearSuffix="a">
Woodburne et al., 2014
<bibRefCitation id="EFD711BAF6F568E7D8EFFB6C1892FB1D" author="a" box="[740,752,1218,1242]" etAl="et al." firstAuthor="a" pageId="206" pageNumber="206" pagination="109 - 119" refId="ref235572" refString="Woodburne, M. O., et al. 2014 a. Revised timing of the South American early Paleogene land mammal ages. Journal of South American Earth Sciences 54: 109 - 119." type="journal article" year="2014" yearSuffix="b">a</bibRefCitation>
</bibRefCitation>
, 2014b;
<bibRefCitation id="EFD711BAF6F568E7D95DFB6C1E51FB1D" author="Krause, J. M." box="[854,1075,1218,1242]" pageId="206" pageNumber="206" pagination="886 - 903" refId="ref212274" refString="Krause, J. M., et al. 2017. New age constraints for early Paleogene strata of central Patagonia, Argentina: Implications for the timing of South American Land Mammal Ages. Geological Society of America Bulletin 129 (7 - 8): 886 - 903." type="journal article" year="2017">Krause et al., 2017</bibRefCitation>
) Las
<collectingCountry id="F3512CDBF6F568E7DE73FB6C1ED9FB1D" box="[1144,1211,1218,1242]" name="Indonesia" pageId="206" pageNumber="206">Flores</collectingCountry>
Local Fauna of southern
<collectingCountry id="F3512CDBF6F568E7D9B5FB4D1E49FB3C" box="[958,1067,1251,1275]" name="Argentina" pageId="206" pageNumber="206">Argentina</collectingCountry>
(
<bibRefCitation id="EFD711BAF6F568E7DE30FB4D1EDAFB3C" author="Goin, F. J." box="[1083,1208,1251,1275]" pageId="206" pageNumber="206" pagination="30 - 42" refId="ref205827" refString="Goin, F. J. 2003. Early marsupial radiations in South America. In M. Jones, C. Dickman, and M. Archer (editors), Predators with pouches: the biology of carnivorous marsupials: 30 - 42. Collingwood, Australia: CSIRO (Commonwealth Scientific and Industrial Research Organization)." type="book chapter" year="2003">Goin, 2003</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F568E7D8A6FAAA195DFADB" author="Zimicz" box="[685,831,1284,1308]" firstAuthor="Zimicz" pageId="206" pageNumber="206" refId="ref237684" refString="Zimicz, A. N. 2012. Ecomorfologia de los marsupiales paleogenos de America del Sur. Ph. D. dissertation, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, La Plata." type="book" year="2012">Zimicz, 2012</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F568E7D940FAAA1E3FFADB" author="Woodburne" box="[843,1117,1284,1308]" etAl="et al." firstAuthor="Woodburne" pageId="206" pageNumber="206" pagination="109 - 119" refId="ref235572" refString="Woodburne, M. O., et al. 2014 a. Revised timing of the South American early Paleogene land mammal ages. Journal of South American Earth Sciences 54: 109 - 119." type="journal article" year="2014" yearSuffix="a">Woodburne et al., 2014a</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F568E7DE61FAAA196DFAFA" author="Goin, F. J. &amp; M. O. Woodburne &amp; A. N. Zimicz &amp; G. M. Martin &amp; L. Chornogubsky" pageId="206" pageNumber="206" refId="ref207099" refString="Goin, F. J., M. O. Woodburne, A. N. Zimicz, G. M. Martin, and L. Chornogubsky. 2016. A brief history of South American metatherians: Evolutionary contexts and intercontinental dispersals. Dordrecht: Springer." type="book" year="2016">Goin et al., 2016</bibRefCitation>
), which is incongruent with our estimate for the time of the Microbiotheria-Diprotodontia split, but these potentially important fossils have yet to be described.
</paragraph>
</subSubSection>
</treatment>
</document>