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<mods:title id="75EBC3B64BB4C2CD35945B95752F68CE">Craniodental Morphology And Phylogeny Of Marsupials</mods:title>
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<taxonomicName id="4C4617C8F6FD68EFD915F9B31E2FF9F2" ID-CoL="34Q" authority="Gill, 1872" authorityName="Gill" authorityYear="1872" box="[798,1101,1565,1589]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Didelphimorphia" pageId="198" pageNumber="198" phylum="Chordata" rank="order">
Didelphimorphia
<bibRefCitation id="EFD711BAF6FD68EFD9E8F9B31E2FF9F2" author="Gill, T." box="[995,1101,1565,1589]" pageId="198" pageNumber="198" pagination="1 - 98" refId="ref205030" refString="Gill, T. 1872. Arrangement of the families of mammals with analytical tables. Smithsonian Miscellaneous Collections 11 (1): 1 - 98." type="journal article" year="1872">Gill, 1872</bibRefCitation>
</taxonomicName>
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</paragraph>
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<subSubSection id="C35C3FC0F6FD68EFD8C7F9E119D1F940" pageId="198" pageNumber="198" type="synonymic_list">
<paragraph id="8BF96C4BF6FD68EFD8C7F9E119D1F940" blockId="198.[684,1211,1614,1738]" pageId="198" pageNumber="198">
CONTENTS:
<taxonomicName id="4C4617C8F6FD68EFD956F9E0198AF9A1" baseAuthorityName="sensu Voss and Jansa" baseAuthorityYear="2009" box="[861,1000,1614,1638]" class="Mammalia" family="Didelphidae" kingdom="Animalia" order="Didelphimorphia" pageId="198" pageNumber="198" phylum="Chordata" rank="family">Didelphidae</taxonomicName>
(see
<tableCitation id="C6C459F0F6FD68EFDE3DF9E01EE8F9A0" box="[1078,1162,1614,1639]" captionStart="TABLE 2" captionStartId="11.[609,689,226,250]" captionTargetPageId="11" captionText="TABLE 2 Assumed Classification of Marsupial Ingroup Genera Scored for Phylogenetic Analysisa" httpUri="http://table.plazi.org/id/DF393CC3F6306822D86AFF4C1E2EFED9" pageId="198" pageNumber="198" tableUuid="DF393CC3F6306822D86AFF4C1E2EFED9">table 2</tableCitation>
for included terminal taxa).
</paragraph>
</subSubSection>
<footnote id="E85D7045F6FD68EFDA87F9331B68F90E" pageId="198" pageNumber="198">
<paragraph id="8BF96C4BF6FD68EFDA87F9331B68F90E" blockId="198.[108,636,1693,1737]" pageId="198" pageNumber="198">
<superScript id="7C33C103F6FD68EFDA87F9331AF8F96E" attach="right" box="[140,154,1693,1705]" fontSize="5" pageId="198" pageNumber="198">29</superScript>
The phylogenetic analysis of
<bibRefCitation id="EFD711BAF6FD68EFDBAEF9301834F976" author="Wilson" box="[421,598,1694,1713]" etAl="et al." firstAuthor="Wilson" pageId="198" pageNumber="198" 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>
is a notable exception.
</paragraph>
</footnote>
<subSubSection id="C35C3FC0F6FD68EED8C7F93F1B40FDE4" lastPageId="199" lastPageNumber="199" pageId="198" pageNumber="198" type="description">
<paragraph id="8BF96C4BF6FD68EFD8C7F93F188EF90D" blockId="198.[684,1211,1614,1738]" pageId="198" pageNumber="198">STEM AGE: 56.2 Mya (95% HPD: 54.758.6 Mya).</paragraph>
<paragraph id="8BF96C4BF6FC68EEDA87FF4C1ACFFEDC" blockId="199.[108,638,225,1738]" pageId="199" pageNumber="199">CROWN AGE: 27.1 Mya (95% HPD: 23.731.1 Mya).</paragraph>
<paragraph id="8BF96C4BF6FC68EEDA87FE8A1B40FDE4" blockId="199.[108,638,225,1738]" pageId="199" pageNumber="199">UNAMBIGUOUS CRANIODENTAL SYNAPOMORPHIES: Premolariform P2 and premolariform P3 subequal in height (char. 119: 21; ci = 0.118); premolariform P3 with well-developed anterior and posterior cutting edges (char. 124: 1→0; ci = 0.667); M4 erupts before P3 (char. 130: 2→0; ci = 0.089); and p2 distinctly taller than p3 (char. 156: 2→0; ci = 0.118).</paragraph>
</subSubSection>
<subSubSection id="C35C3FC0F6FC68E0DA86FD821BFCFC8A" lastPageId="201" lastPageNumber="201" pageId="199" pageNumber="199" type="discussion">
<paragraph id="8BF96C4BF6FC68EEDA86FD821B39FB1E" blockId="199.[108,638,225,1738]" pageId="199" pageNumber="199">
COMMENTS: As noted by numerous authors (e.g.,
<bibRefCitation id="EFD711BAF6FC68EEDABDFDE31BBEFDA2" author="Aplin, K. P. &amp; M. Archer" box="[182,476,589,613]" pageId="199" pageNumber="199" 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>
;
<bibRefCitation id="EFD711BAF6FC68EEDBFBFDE3181BFDA2" author="Goin, F. J." box="[496,633,589,613]" pageId="199" pageNumber="199" 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="EFD711BAF6FC68EEDA67FDC01B30FD40" author="Forasiepi, A. M. &amp; F. Goin &amp; A. G. Martinelli" box="[108,338,622,647]" pageId="199" pageNumber="199" pagination="1252 - 1263" refId="ref203314" refString="Forasiepi, A. M., F. Goin, and A. G. Martinelli. 2009. Contribution to the knowledge of the Sparassocynidae (Mammalia, Metatheria, Didelphoidea), with comments on the age of the Aisol Formation (Neogene), Mendoza Province, Argentina. Journal of Vertebrate Paleontology 29 (4): 1252 - 1263." type="journal article" year="2009">Forasiepi et al., 2009</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6FC68EEDB57FDC11824FD40" author="Voss and Jansa" box="[348,582,622,647]" firstAuthor="Voss" pageId="199" pageNumber="199" refId="ref232342" refString="Voss, R. S., and S. A. Jansa. 2021. Opossums: an adaptive radiation of New World marsupials, Baltimore: Johns Hopkins University Press." type="book" year="2021">Voss and Jansa, 2021</bibRefCitation>
), the name “
<taxonomicName id="4C4617C8F6FC68EEDAB4FD211BE7FD60" authorityName="Gill" authorityYear="1872" box="[191,389,655,679]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Didelphimorphia" pageId="199" pageNumber="199" phylum="Chordata" rank="order">Didelphimorphia</taxonomicName>
” has long been used for a nonmonophyletic assemblage of dentally plesiomorphic metatherians that included both marsupials and stem metatherians. By contrast, we follow the stem-based phylogenetic definition of
<taxonomicName id="4C4617C8F6FC68EEDAB6FC9B1B1DFC8A" authorityName="Gill" authorityYear="1872" box="[189,383,821,845]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Didelphimorphia" pageId="199" pageNumber="199" phylum="Chordata" rank="order">Didelphimorphia</taxonomicName>
proposed by
<bibRefCitation id="EFD711BAF6FC68EED81DFC9B1B5AFCA9" author="Beck, R. M. D. &amp; M. L. Taglioretti" pageId="199" pageNumber="199" pagination="385 - 417" refId="ref192802" refString="Beck, R. M. D., and M. L. Taglioretti. 2020. A nearly complete juvenile skull of the marsupial Sparassocynus derivatus from the Pliocene of Argentina, the affinities of &quot; sparassocynids, &quot; and the diversification of opossums (Marsupialia; Didelphimorphia; Didelphidae). Journal of Mammalian Evolution 27: 385 - 417." type="journal article" year="2020">Beck and Taglioretti (2020)</bibRefCitation>
, namely, the most inclusive clade containing
<taxonomicName id="4C4617C8F6FC68EEDB25FCD9187EFC48" authorityName="Linnaeus" authorityYear="1758" box="[302,540,887,911]" class="Mammalia" family="Didelphidae" genus="Didelphis" kingdom="Animalia" order="Didelphimorphia" pageId="199" pageNumber="199" phylum="Chordata" rank="species" species="marsupialis">
<emphasis id="B932B059F6FC68EEDB25FCD9187EFC48" box="[302,540,887,911]" italics="true" pageId="199" pageNumber="199">Didelphis marsupialis</emphasis>
</taxonomicName>
, but not
<taxonomicName id="4C4617C8F6FC68EEDA66FC361B09FC77" baseAuthorityName="Tomes" baseAuthorityYear="1863" box="[109,363,920,944]" class="Mammalia" family="Caenolestidae" genus="Caenolestes" kingdom="Animalia" order="Paucituberculata" pageId="199" pageNumber="199" phylum="Chordata" rank="species" species="fuliginosus">
<emphasis id="B932B059F6FC68EEDA66FC361B09FC77" box="[109,363,920,944]" italics="true" pageId="199" pageNumber="199">Caenolestes fuliginosus</emphasis>
</taxonomicName>
or
<taxonomicName id="4C4617C8F6FC68EEDB9DFC361814FC77" baseAuthorityName="Pallas" baseAuthorityYear="1766" box="[406,630,920,944]" class="Mammalia" family="Phalangeridae" genus="Phalanger" kingdom="Animalia" order="Diprotodontia" pageId="199" pageNumber="199" phylum="Chordata" rank="species" species="orientalis">
<emphasis id="B932B059F6FC68EEDB9DFC361814FC77" box="[406,630,920,944]" italics="true" pageId="199" pageNumber="199">Phalanger orientalis</emphasis>
</taxonomicName>
. The family
<taxonomicName id="4C4617C8F6FC68EEDAE7FC171ACFFC35" authority="(sensu Voss and Jansa, 2009)" baseAuthorityName="sensu Voss and Jansa" baseAuthorityYear="2009" class="Mammalia" family="Didelphidae" kingdom="Animalia" order="Didelphimorphia" pageId="199" pageNumber="199" phylum="Chordata" rank="family">
Didelphidae (sensu
<bibRefCitation id="EFD711BAF6FC68EEDBC5FC141AC4FC35" author="Voss and Jansa" firstAuthor="Voss" pageId="199" pageNumber="199" pagination="1 - 177" refId="ref232295" refString="Voss, R. S., and S. A. Jansa. 2009. Phylogenetic relationships and classification of didelphid marsupials, an extant radiation of New World metatherian mammals. Bulletin of the American Museum of Natural History 322: 1 - 177." type="journal article" year="2009">Voss and Jansa, 2009</bibRefCitation>
)
</taxonomicName>
comprises the entire didelphimorphian crown clade. All four of our fossil didelphimorphian terminals (†
<taxonomicName id="4C4617C8F6FC68EEDB4FFBB31BBEFBF3" box="[324,476,1053,1076]" class="Mammalia" family="Sparassocynidae" genus="Hesperocynus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Didelphimorphia" pageId="199" pageNumber="199" phylum="Chordata" rank="genus">
<emphasis id="B932B059F6FC68EEDB4FFBB31BBEFBF3" box="[324,476,1053,1076]" italics="true" pageId="199" pageNumber="199">Hesperocynus</emphasis>
</taxonomicName>
, †
<taxonomicName id="4C4617C8F6FC68EEDBF0FBB31AF1FB92" baseAuthorityName="Beck and Taglioretti" baseAuthorityYear="2020" class="Mammalia" family="Didelphidae" genus="Sparassocynus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Didelphimorphia" pageId="199" pageNumber="199" phylum="Chordata" rank="genus">
<emphasis id="B932B059F6FC68EEDBF0FBB31AF1FB92" italics="true" pageId="199" pageNumber="199">Sparassocynus</emphasis>
</taxonomicName>
, †
<taxonomicName id="4C4617C8F6FC68EEDAB9FB931B3CFB92" authorityName="Reig" authorityYear="1952" box="[178,350,1085,1109]" class="Mammalia" family="Didelphidae" genus="Thylatheridium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Didelphimorphia" pageId="199" pageNumber="199" phylum="Chordata" rank="genus">
<emphasis id="B932B059F6FC68EEDAB9FB931B3CFB92" box="[178,350,1085,1109]" italics="true" pageId="199" pageNumber="199">Thylatheridium</emphasis>
</taxonomicName>
, and †
<taxonomicName id="4C4617C8F6FC68EEDBBAFB931826FB92" box="[433,580,1085,1109]" class="Mammalia" family="Didelphidae" genus="Thylophorops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Didelphimorphia" pageId="199" pageNumber="199" phylum="Chordata" rank="genus">
<emphasis id="B932B059F6FC68EEDBBAFB931826FB92" box="[433,580,1085,1109]" italics="true" pageId="199" pageNumber="199">Thylophorops</emphasis>
</taxonomicName>
) fall within the crown clade, so craniodental synapomorphies for
<taxonomicName id="4C4617C8F6FC68EEDB4FFBD1186FFB50" authorityName="Gill" authorityYear="1872" box="[324,525,1151,1175]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Didelphimorphia" pageId="199" pageNumber="199" phylum="Chordata" rank="order">Didelphimorphia</taxonomicName>
are also synapomorphies of
<taxonomicName id="4C4617C8F6FC68EEDB43FB0E1BB3FB7F" baseAuthorityName="sensu Voss and Jansa" baseAuthorityYear="2009" box="[328,465,1184,1208]" class="Mammalia" family="Didelphidae" kingdom="Animalia" order="Didelphimorphia" pageId="199" pageNumber="199" phylum="Chordata" rank="family">Didelphidae</taxonomicName>
in our applications of these names.
</paragraph>
<paragraph id="8BF96C4BF6FC68EEDA87FB4D19ECFDA2" blockId="199.[108,638,225,1738]" lastBlockId="199.[683,1213,226,1605]" pageId="199" pageNumber="199">
All the craniodental features that optimize as unambiguous synapomorphies of
<taxonomicName id="4C4617C8F6FC68EEDBECFAAA1AF9FAFA" authorityName="Gill" authorityYear="1872" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Didelphimorphia" pageId="199" pageNumber="199" phylum="Chordata" rank="order">Didelphimorphia</taxonomicName>
/
<taxonomicName id="4C4617C8F6FC68EEDAA8FA8B1B4BFAFA" baseAuthorityName="sensu Voss and Jansa" baseAuthorityYear="2009" box="[163,297,1317,1341]" class="Mammalia" family="Didelphidae" kingdom="Animalia" order="Didelphimorphia" pageId="199" pageNumber="199" phylum="Chordata" rank="family">Didelphidae</taxonomicName>
show high levels of homoplasy, as indicated by their low consistency indices (see above). One potential craniodental synapomorphy of
<taxonomicName id="4C4617C8F6FC68EEDABCFA261B16FA67" authorityName="Gill" authorityYear="1872" box="[183,372,1416,1440]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Didelphimorphia" pageId="199" pageNumber="199" phylum="Chordata" rank="order">Didelphimorphia</taxonomicName>
/
<taxonomicName id="4C4617C8F6FC68EEDB77FA261860FA67" baseAuthorityName="sensu Voss and Jansa" baseAuthorityYear="2009" box="[380,514,1416,1440]" class="Mammalia" family="Didelphidae" kingdom="Animalia" order="Didelphimorphia" pageId="199" pageNumber="199" phylum="Chordata" rank="family">Didelphidae</taxonomicName>
that shows less homoplasy is precocious fusion of the suture between the interparietal and supraoccipital (char. 31, state 1). This feature was first noted in this context by
<bibRefCitation id="EFD711BAF6FC68EEDB1AF9A31B9DF9E3" author="Voss and Jansa" box="[273,511,1548,1572]" firstAuthor="Voss" pageId="199" pageNumber="199" pagination="1 - 177" refId="ref232295" refString="Voss, R. S., and S. A. Jansa. 2009. Phylogenetic relationships and classification of didelphid marsupials, an extant radiation of New World metatherian mammals. Bulletin of the American Museum of Natural History 322: 1 - 177." type="journal article" year="2009">Voss and Jansa (2009)</bibRefCitation>
, who stated that it is unique to didelphids within
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, but we also observed it in dactylopsiline petaurids (see char. 31). Nevertheless, interparietal-supraoccipital fusion optimizes as a synapomorphy of
<taxonomicName id="4C4617C8F6FC68EEDB3AF91C1B8CF90D" authorityName="Gill" authorityYear="1872" box="[305,494,1714,1738]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Didelphimorphia" pageId="199" pageNumber="199" phylum="Chordata" rank="order">Didelphimorphia</taxonomicName>
/
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only under Accelerated Transformation. The reason is that it could not be scored in key nondidelphimorphian taxa that would help unambiguously resolve the branch along which this feature arose, either because suitably preserved material of an appropriate ontogenetic age is unavailable (e.g., for paucituberculatans, †
<taxonomicName id="4C4617C8F6FC68EED8B1FE671929FE26" box="[698,843,457,481]" class="Mammalia" family="Yalkaparidontidae" genus="Yalkaparidon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="199" pageNumber="199" phylum="Chordata" rank="genus">
<emphasis id="B932B059F6FC68EED8B1FE671929FE26" box="[698,843,457,481]" italics="true" pageId="199" pageNumber="199">Yalkaparidon</emphasis>
</taxonomicName>
, †
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<emphasis id="B932B059F6FC68EED962FE671E7BFE26" box="[873,1049,457,481]" italics="true" pageId="199" pageNumber="199">Mimoperadectes</emphasis>
</taxonomicName>
, and our outgroup terminals), or because this character is inapplicable (as is the case for †
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<emphasis id="B932B059F6FC68EEDE00FDA51ED4FDE4" box="[1035,1206,523,547]" italics="true" pageId="199" pageNumber="199">Herpetotherium</emphasis>
</taxonomicName>
, in which a distinct interparietal appears to be absent; see char. 30).
</paragraph>
<paragraph id="8BF96C4BF6FC68EED8C6FDC119B6FAB2" blockId="199.[683,1213,226,1605]" pageId="199" pageNumber="199">
In agreement with
<bibRefCitation id="EFD711BAF6FC68EED9A4FDC01E57FD40" author="Goin, F. J." box="[943,1077,622,647]" pageId="199" pageNumber="199" refId="ref205607" refString="Goin, F. J. 1991. Los Didelphoidea (Mammalia, Marsupialia) del Cenozoico tardio de la region Pampeana. Ph. D. dissertation, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, La Plata." type="book" year="1991">Goin (1991</bibRefCitation>
,
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), our dated total-evidence analysis (
<figureCitation id="137D70CEF6FC68EED9F4FD211E2BFD6F" box="[1023,1097,655,680]" 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="199" pageNumber="199">fig. 33</figureCitation>
) indicates that the didelphimorphian crown clade is a relatively young radiation, with its earliest divergence estimated as late Oligocene or early Miocene. This estimate is also broadly congruent with the molecular node-dating analysis of
<bibRefCitation id="EFD711BAF6FC68EEDE88FC9B194EFCA9" author="Jansa, S. A. &amp; F. K. Barker &amp; R. S. Voss" pageId="199" pageNumber="199" pagination="684 - 695" refId="ref210091" refString="Jansa, S. A., F. K. Barker, and R. S. Voss. 2014. The early diversification history of didelphid marsupials: a window into South America's &quot; splendid isolation. &quot; Evolution 68 (3): 684 - 695." type="journal article" year="2014">Jansa et al. (2014)</bibRefCitation>
and with the total-evidence tip-andnode dating analysis of
<bibRefCitation id="EFD711BAF6FC68EED9C5FCD91897FC77" author="Beck, R. M. D. &amp; M. L. Taglioretti" pageId="199" pageNumber="199" pagination="385 - 417" refId="ref192802" refString="Beck, R. M. D., and M. L. Taglioretti. 2020. A nearly complete juvenile skull of the marsupial Sparassocynus derivatus from the Pliocene of Argentina, the affinities of &quot; sparassocynids, &quot; and the diversification of opossums (Marsupialia; Didelphimorphia; Didelphidae). Journal of Mammalian Evolution 27: 385 - 417." type="journal article" year="2020">Beck and Taglioretti (2020)</bibRefCitation>
; however, it is considerably younger than estimated divergence dates suggested by several other molecular studies, which have dated the last common ancestor of living didelphimorphians to the early Oligocene or Eocene (
<bibRefCitation id="EFD711BAF6FC68EEDE65FBB21943FB92" author="Steiner" etAl="et al." firstAuthor="Steiner" pageId="199" pageNumber="199" pagination="363 - 379" refId="ref228396" refString="Steiner, C., M. - k. Tilak, E. J. P. Douzery, and F. M. Catzeflis. 2005. New DNA data from a transthyretin nuclear intron suggest an Oligocene to Miocene diversification of living South America opossums (Marsupialia: Didelphidae). Molecular Phylogenetics and Evolution 35: 363 - 379." type="journal article" year="2005">Steiner et al., 2005</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6FC68EED927FB931E7AFB92" author="Meredith, R. W. &amp; M. Westerman &amp; J. A. Case &amp; M. S. Springer" box="[812,1048,1085,1109]" pageId="199" pageNumber="199" 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="EFD711BAF6FC68EEDE28FB931E3BFB92" author="Meredith, R. W." box="[1059,1113,1085,1109]" pageId="199" pageNumber="199" pagination="521 - 524" refId="ref216858" refString="Meredith, R. W., et al. 2011. Impacts of the Cretaceous Terrestrial Revolution and KPg extinction on mammal diversification. Science 334: 521 - 524." type="journal article" year="2011">2011</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6FC68EEDE68FB931946FBB0" author="Mitchell, K. J." pageId="199" pageNumber="199" 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="EFD711BAF6FC68EED924FBF01989FBB1" author="Vilela" box="[815,1003,1118,1142]" etAl="et al." firstAuthor="Vilela" pageId="199" pageNumber="199" pagination="414 - 427" refId="ref232150" refString="Vilela, J. F., J. A. De Oliveira, and C. A. D. Russo. 2015. The diversification of the genus Monodelphis and the chronology of Didelphidae (Didelphimorphia). Zoological Journal of the Linnean Society 174 (2): 414 - 427." type="journal article" year="2015">Vilela et al., 2015</bibRefCitation>
;
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; see table 6). Our estimated crown age is likewise broadly congruent with the relevant fossil record (reviewed by
<bibRefCitation id="EFD711BAF6FC68EED9DEFB6C1881FB3C" author="Beck, R. M. D. &amp; M. L. Taglioretti" pageId="199" pageNumber="199" pagination="385 - 417" refId="ref192802" refString="Beck, R. M. D., and M. L. Taglioretti. 2020. A nearly complete juvenile skull of the marsupial Sparassocynus derivatus from the Pliocene of Argentina, the affinities of &quot; sparassocynids, &quot; and the diversification of opossums (Marsupialia; Didelphimorphia; Didelphidae). Journal of Mammalian Evolution 27: 385 - 417." type="journal article" year="2020">Beck and Taglioretti, 2020</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6FC68EED8F9FB4D19AAFB3C" author="Castro, M. C. &amp; M. J. Dahur &amp; G. S. Ferreira" box="[754,968,1251,1275]" pageId="199" pageNumber="199" pagination="583 - 598" refId="ref196923" refString="Castro, M. C., M. J. Dahur, and G. S. Ferreira. 2021. Amazonia as the origin and diversification area of Didelphidae (Mammalia: Metatheria), and a review of the fossil record of the clade. Journal of Mammalian Evolution 28: 583 - 598." type="journal article" year="2021">Castro et al., 2021</bibRefCitation>
), in which the oldest known (probable crown-clade) didelphids are from the early Miocene (Colhuehuapian SALMA, ~21.020.1 Mya;
<bibRefCitation id="EFD711BAF6FC68EED962FAE81E4FFA99" author="Goin, F." box="[873,1069,1350,1374]" pageId="199" pageNumber="199" pagination="29 - 71" refId="ref206561" refString="Goin, F., et al. 2007 a. Los Metatheria sudamericanos de comienzos del Neogeno (Mioceno Temprano, Edad-mamifero Colhuehuapense). Parte I: Introduccion, Didelphimorphia y Sparassodonta. Ameghiniana 44 (1): 29 - 71." type="journal article" year="2007">Goin et al., 2007a</bibRefCitation>
;
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;
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).
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</paragraph>
<paragraph id="8BF96C4BF6FC68E0D8C7FA261BFCFC8A" blockId="199.[683,1213,226,1605]" lastBlockId="201.[108,637,226,845]" lastPageId="201" lastPageNumber="201" pageId="199" pageNumber="199">
The comparatively long branch between the divergence of
<taxonomicName id="4C4617C8F6FC68EED955FA071E4AFA06" authorityName="Gill" authorityYear="1872" box="[862,1064,1449,1473]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Didelphimorphia" pageId="199" pageNumber="199" phylum="Chordata" rank="order">Didelphimorphia</taxonomicName>
from other marsupials and the first diversification of lineages ancestral to living didelphids suggests a long history (presumably largely or entirely South American;
<bibRefCitation id="EFD711BAF6FC68EED970F9801E27F982" author="Jansa, S. A. &amp; F. K. Barker &amp; R. S. Voss" box="[891,1093,1581,1605]" pageId="199" pageNumber="199" pagination="684 - 695" refId="ref210091" refString="Jansa, S. A., F. K. Barker, and R. S. Voss. 2014. The early diversification history of didelphid marsupials: a window into South America's &quot; splendid isolation. &quot; Evolution 68 (3): 684 - 695." type="journal article" year="2014">Jansa et al., 2014</bibRefCitation>
; Voss and Jansa, 2021) of stem didelphimorphians; because none have been reported to date—in effect, a protracted ghost lineage. However, given the lack of dental synapomorphies characterizing
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Didelphimorphia (
<bibRefCitation id="EFD711BAF6F268E0DBC9FEC81AC5FE58" author="Voss and Jansa" firstAuthor="Voss" pageId="201" pageNumber="201" pagination="1 - 177" refId="ref232295" refString="Voss, R. S., and S. A. Jansa. 2009. Phylogenetic relationships and classification of didelphid marsupials, an extant radiation of New World metatherian mammals. Bulletin of the American Museum of Natural History 322: 1 - 177." type="journal article" year="2009">Voss and Jansa, 2009</bibRefCitation>
)
</taxonomicName>
, confidently distinguishing stem didelphimorphians from other dentally plesiomorphic marsupialiforms is likely to be difficult in the absence of well-preserved cranial specimens. Indeed, it seems likely that stem didelphimorphians are represented but remain unidentified among the rich dental record of Palaeogene marsupialiforms from South America (
<bibRefCitation id="EFD711BAF6F268E0DAD6FD211BC3FD60" author="Goin, F. J. &amp; M. O. Woodburne &amp; A. N. Zimicz &amp; G. M. Martin &amp; L. Chornogubsky" box="[221,417,655,679]" pageId="201" pageNumber="201" 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>
). Postcranial material may prove useful in future attempts to identify stem didelphimorphians, because putative synapomorphies have been identified in the tarsus (
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;
<bibRefCitation id="EFD711BAF6F268E0DBB6FCBA1B97FCEB" author="Szalay" box="[445,501,788,812]" firstAuthor="Szalay" pageId="201" pageNumber="201" 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">1994</bibRefCitation>
;
<bibRefCitation id="EFD711BAF6F268E0D80FFCBA1A94FC8A" author="Szalay and Sargis" firstAuthor="Szalay" pageId="201" pageNumber="201" pagination="139 - 302" refId="ref229157" refString="Szalay, F. S., and E. J. Sargis. 2001. Model-based analysis of postcranial osteology of marsupials from the Palaeocene of Itaborai (Brazil) and the phylogenetics and biogeography of Metatheria. Geodiversitas 23 (2): 139 - 302." type="journal article" year="2001">Szalay and Sargis, 2001</bibRefCitation>
;
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, 2009
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).
</paragraph>
</subSubSection>
</treatment>
</document>