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<document id="4ADBF1E2E216764F314A9AD3793F9FD6" ID-CLB-Dataset="3273" ID-DOI="10.1093/zoolinnean/zly040" ID-GBIF-Dataset="685b49cb-9812-445d-842d-c5566f2de2cb" ID-ISSN="0024-4082" ID-Zenodo-Dep="5715169" ID-ZooBank="D50F5678-824D-477B-9DC9-F39DD40002B9" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1619628097858" checkinUser="carolina" docAuthor="Scott, Craig S." docDate="2019" docId="03B6879EFFBFD91CFF40AFA9DDF3FDA5" docLanguage="en" docName="ZoolJLinnSoc.185.431-458.pdf.imf" docOrigin="Zoological Journal of the Linnean Society 185" docStyle="DocumentStyle:36B3BD6A90C22AB4F7F465C853188CC8.7:ZoolJLinnSoc.2017-2023.journal_article" docStyleId="36B3BD6A90C22AB4F7F465C853188CC8" docStyleName="ZoolJLinnSoc.2017-2023.journal_article" docStyleVersion="7" docTitle="Ferrequitherium Scott 2019" docType="treatment" docVersion="8" lastPageNumber="446" masterDocId="FF8FFFE6FFB2D90CFFB8A93DDF4AFFEE" masterDocTitle="Horolodectidae: a new family of unusual eutherians (Mammalia: Theria) from the Palaeocene of Alberta, Canada" masterLastPageNumber="458" masterPageNumber="431" pageNumber="444" updateTime="1698937877048" updateUser="ExternalLinkService">
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<mods:title id="C7C2704CE0EF526EF46229AAF34A787D">Horolodectidae: a new family of unusual eutherians (Mammalia: Theria) from the Palaeocene of Alberta, Canada</mods:title>
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<mods:namePart id="F8EE19FF0B46038CE28517AC63862273">Scott, Craig S.</mods:namePart>
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<mods:date id="334550174B3B34763F32AB41DB6498B5">2019</mods:date>
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<mods:number id="33F5632232EBF5B807624BD7CAD7E761">2018-08-23</mods:number>
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<mods:identifier id="40AFA37B0D69ECAB5C453EAF68A17FD6" type="DOI">10.1093/zoolinnean/zly040</mods:identifier>
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<treatment id="03B6879EFFBFD91CFF40AFA9DDF3FDA5" ID-DOI="http://doi.org/10.5281/zenodo.5730331" ID-GBIF-Taxon="190872618" ID-Zenodo-Dep="5730331" LSID="urn:lsid:plazi:treatment:03B6879EFFBFD91CFF40AFA9DDF3FDA5" httpUri="http://treatment.plazi.org/id/03B6879EFFBFD91CFF40AFA9DDF3FDA5" lastPageId="16" lastPageNumber="446" pageId="13" pageNumber="444">
<subSubSection id="C3056503FFBFD901FF40AFA9DD79F924" pageId="13" pageNumber="444" type="nomenclature">
<paragraph id="8BA03688FFBFD901FF40AFA9DD79F924" blockId="13.[248,694,1684,1738]" pageId="13" pageNumber="444">
<heading id="D0E881E4FFBFD901FF40AFA9DD79F924" centered="true" fontSize="9" level="2" pageId="13" pageNumber="444" reason="2">
AFFINITIES OF
<taxonomicName id="4C1F4D0BFFBFD901FE1BAFA9DDCAF945" authorityName="Scott" authorityYear="2019" box="[419,640,1684,1707]" class="Mammalia" family="Zhelestidae" genus="Ferrequitherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Proteutheria" pageId="13" pageNumber="451" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBFD901FE1BAFA9DDCAF945" box="[419,640,1684,1707]" italics="true" pageId="13" pageNumber="444">FERREQUITHERIUM</emphasis>
</taxonomicName>
AND
<emphasis id="B96BEA9AFFBFD901FEC3AF8EDD79F924" box="[379,563,1715,1738]" italics="true" pageId="13" pageNumber="444">HOROLODECTES</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C3056503FFBFD91CFF1BAFE6DDF3FDA5" lastPageId="16" lastPageNumber="447" pageId="13" pageNumber="444" type="description">
<paragraph id="8BA03688FFBFD901FF1BAFE6DBF6FC25" blockId="13.[163,778,1755,1899]" lastBlockId="13.[827,1443,704,971]" pageId="13" pageNumber="444">
To examine the evolutionary relationships of
<taxonomicName id="4C1F4D0BFFBFD901FF1BAFC7DE21F8E1" authorityName="Scott" authorityYear="2019" box="[163,363,1786,1807]" class="Mammalia" family="Zhelestidae" genus="Ferrequitherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Proteutheria" pageId="13" pageNumber="444" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBFD901FF1BAFC7DE21F8E1" box="[163,363,1786,1807]" italics="true" pageId="13" pageNumber="444">Ferrequitherium</emphasis>
</taxonomicName>
and
<taxonomicName id="4C1F4D0BFFBFD901FE14AFC7DD0DF8E1" authorityName="Scott, Fox &amp; Webb" authorityYear="2006" box="[428,583,1786,1807]" class="Mammalia" family="Horolodectidae" genus="Horolodectes" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="13" pageNumber="444" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBFD901FE14AFC7DD0DF8E1" box="[428,583,1786,1807]" italics="true" pageId="13" pageNumber="444">Horolodectes</emphasis>
</taxonomicName>
, a phylogenetic analysis was performed using a newly constructed character–taxon matrix. The matrix consists of 81 characters, compiled through a combination of direct observation of specimens and descriptions from the literature. The specimens and literature used in the analyses, character list and data matrix are presented in Supporting Information, Appendices S2–S4. In addition to
<taxonomicName id="4C1F4D0BFFBFD901FC32AA06DB0FFCBE" authorityName="Scott" authorityYear="2019" box="[906,1093,827,848]" class="Mammalia" family="Zhelestidae" genus="Ferrequitherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Proteutheria" pageId="13" pageNumber="444" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBFD901FC32AA06DB0FFCBE" box="[906,1093,827,848]" italics="true" pageId="13" pageNumber="444">Ferrequitherium</emphasis>
</taxonomicName>
and
<taxonomicName id="4C1F4D0BFFBFD901FBC5AA06DA47FCBE" authorityName="Scott, Fox &amp; Webb" authorityYear="2006" box="[1149,1293,827,848]" class="Mammalia" family="Horolodectidae" genus="Horolodectes" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="13" pageNumber="444" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBFD901FBC5AA06DA47FCBE" box="[1149,1293,827,848]" italics="true" pageId="13" pageNumber="444">Horolodectes</emphasis>
</taxonomicName>
, the analysis includes several of the taxa discussed in Scott
<emphasis id="B96BEA9AFFBFD901FAD0AA67DAE9FC80" box="[1384,1443,857,879]" italics="true" pageId="13" pageNumber="444">et al.</emphasis>
(2006) as potential near-relatives of
<taxonomicName id="4C1F4D0BFFBFD901FB5FAA45DA31FC63" authorityName="Scott, Fox &amp; Webb" authorityYear="2006" box="[1255,1403,888,909]" class="Mammalia" family="Horolodectidae" genus="Horolodectes" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="13" pageNumber="444" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBFD901FB5FAA45DA31FC63" box="[1255,1403,888,909]" italics="true" pageId="13" pageNumber="444">Horolodectes</emphasis>
</taxonomicName>
, as well as additional taxa with dental morphologies that resemble that of
<taxonomicName id="4C1F4D0BFFBFD901FC43AA88DBFCFC24" authorityName="Scott" authorityYear="2019" box="[1019,1206,949,970]" class="Mammalia" family="Zhelestidae" genus="Ferrequitherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Proteutheria" pageId="13" pageNumber="444" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBFD901FC43AA88DBFCFC24" box="[1019,1206,949,970]" italics="true" pageId="13" pageNumber="444">Ferrequitherium</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BA03688FFBFD901FC83AADEDB4AFC14" blockId="13.[827,1443,994,1171]" box="[827,1024,994,1018]" pageId="13" pageNumber="444">
<emphasis id="B96BEA9AFFBFD901FC83AADEDB4AFC14" box="[827,1024,994,1018]" italics="true" pageId="13" pageNumber="444">Taxon sampling</emphasis>
</paragraph>
<paragraph id="8BA03688FFBFD901FC83AD3EDA01FB7D" blockId="13.[827,1443,994,1171]" pageId="13" pageNumber="444">
The ingroup includes the following taxa:
<taxonomicName id="4C1F4D0BFFBFD901FC83AD1FDC98FBD6" authorityName="Scott, Fox &amp; Webb" authorityYear="2006" box="[827,978,1058,1080]" class="Mammalia" family="Horolodectidae" genus="Horolodectes" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="13" pageNumber="456" phylum="Chordata" rank="genus">Horolodectes</taxonomicName>
<emphasis id="B96BEA9AFFBFD901FC62AD1FDB4DFBD9" box="[986,1031,1058,1079]" italics="true" pageId="13" pageNumber="444">and</emphasis>
Ferrequitherium:
<taxonomicName id="4C1F4D0BFFBFD901FB5BAD1FDAE9FBD9" authorityName="Scott" authorityYear="2019" box="[1251,1443,1058,1079]" class="Mammalia" family="Zhelestidae" genus="Ferrequitherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Proteutheria" pageId="13" pageNumber="444" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBFD901FB5BAD1FDAE9FBD9" box="[1251,1443,1058,1079]" italics="true" pageId="13" pageNumber="444">Ferrequitherium</emphasis>
</taxonomicName>
and its hypothesized close relative
<taxonomicName id="4C1F4D0BFFBFD901FB62AD7DDA25FBBB" authorityName="Scott, Fox &amp; Webb" authorityYear="2006" box="[1242,1391,1088,1109]" class="Mammalia" family="Horolodectidae" genus="Horolodectes" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="13" pageNumber="444" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBFD901FB62AD7DDA25FBBB" box="[1242,1391,1088,1109]" italics="true" pageId="13" pageNumber="444">Horolodectes</emphasis>
</taxonomicName>
, are each monotypic and are represented by
<taxonomicName id="4C1F4D0BFFBFD901FAB3AD5DDA25FB9B" authorityName="Scott" authorityYear="2019" box="[1291,1391,1119,1141]" class="Mammalia" family="Zhelestidae" genus="Ferrequitherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Proteutheria" pageId="13" pageNumber="444" phylum="Chordata" rank="species" species="sweeti">
<emphasis id="B96BEA9AFFBFD901FAB3AD5DDA51FB9B" box="[1291,1307,1120,1141]" italics="true" pageId="13" pageNumber="444">F</emphasis>
.
<emphasis id="B96BEA9AFFBFD901FA91AD5DDA25FB9B" box="[1321,1391,1120,1141]" italics="true" pageId="13" pageNumber="444">sweeti</emphasis>
</taxonomicName>
and
<taxonomicName id="4C1F4D0BFFBFD901FC83AD43DBFEFB7D" authority="Scott, Fox &amp; Webb 2006" authorityName="Scott, Fox &amp; Webb" authorityYear="2006" box="[827,1204,1149,1171]" class="Mammalia" family="Horolodectidae" genus="Horolodectes" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="13" pageNumber="444" phylum="Chordata" rank="species" species="sunae">
<emphasis id="B96BEA9AFFBFD901FC83AD43DC1AFB7D" box="[827,848,1150,1171]" italics="true" pageId="13" pageNumber="444">H</emphasis>
.
<emphasis id="B96BEA9AFFBFD901FCE4AD43DCD4FB7D" box="[860,926,1150,1171]" italics="true" pageId="13" pageNumber="444">sunae</emphasis>
<bibRefCitation id="EF8E4B79FFBFD901FC1DAD43DBFEFB7D" author="Scott CS &amp; Webb MW &amp; Fox RC" box="[933,1204,1149,1171]" pageId="13" pageNumber="444" pagination="1009 - 1025" refId="ref20508" refString="Scott CS, Webb MW, Fox RC. 2006. Horolodectes sunae, an enigmatic mammal from the Late Paleocene of Alberta, Canada. Journal of Paleontology 80: 1009 - 1025." type="journal article" year="2006">Scott, Fox &amp; Webb 2006</bibRefCitation>
</taxonomicName>
, respectively.
</paragraph>
<paragraph id="8BA03688FFBFD902FC83AD8CDD53FD49" blockId="13.[827,1444,1201,1898]" lastBlockId="14.[145,761,197,679]" lastPageId="14" lastPageNumber="445" pageId="13" pageNumber="444">
<emphasis id="B96BEA9AFFBFD901FC83AD8CDCB7FB28" box="[827,1021,1201,1222]" italics="true" pageId="13" pageNumber="444">
<taxonomicName id="4C1F4D0BFFBFD901FC83AD8CDCBDFB28" authorityName="Simpson" authorityYear="1937" box="[827,1015,1201,1222]" class="Mammalia" family="Pentacodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pantolesta" pageId="13" pageNumber="444" phylum="Chordata" rank="family">Pentacodontidae</taxonomicName>
:
</emphasis>
The
<taxonomicName id="4C1F4D0BFFBFD901FB8FAD8CDAE9FB29" authority="Simpson, 1937" authorityName="Simpson" authorityYear="1937" box="[1079,1443,1201,1223]" class="Mammalia" family="Pentacodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pantolesta" pageId="13" pageNumber="444" phylum="Chordata" rank="family">
Pentacodontidae
<bibRefCitation id="EF8E4B79FFBFD901FB44AD8CDAE9FB29" author="Simpson GG" box="[1276,1443,1201,1223]" pageId="13" pageNumber="444" pagination="1 - 287" refId="ref20798" refString="Simpson GG. 1937. The Fort Union of the Crazy Mountain Field, Montana and its mammalian faunas. Bulletin of the United States National Museum 169: 1 - 287." type="journal article" year="1937">Simpson, 1937</bibRefCitation>
</taxonomicName>
are a family of unusual, Palaeocene–Eocene-age eutherians characterized by moderately to greatly enlarged upper and lower fourth premolars and relatively unspecialized molars (
<bibRefCitation id="EF8E4B79FFBFD901FB82AC11DBAAFAAF" author="Simpson GG" box="[1082,1248,1324,1346]" pageId="13" pageNumber="444" pagination="1 - 287" refId="ref20798" refString="Simpson GG. 1937. The Fort Union of the Crazy Mountain Field, Montana and its mammalian faunas. Bulletin of the United States National Museum 169: 1 - 287." type="journal article" year="1937">Simpson, 1937</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBFD901FB55AC11DC38FA8E" author="Gingerich PD &amp; Houde P &amp; Krause DW" pageId="13" pageNumber="444" pagination="957 - 970" refId="ref17776" refString="Gingerich PD, Houde P, Krause DW. 1983. A new earliest Tiffanian (Late Paleocene) mammalian fauna from Bangtail Plateau, western Crazy Mountain Basin, Montana. Journal of Paleontology 57: 957 - 970." type="journal article" year="1983">
Gingerich
<emphasis id="B96BEA9AFFBFD901FADDAC10DAD7FAAF" box="[1381,1437,1324,1346]" italics="true" pageId="13" pageNumber="444">et al.</emphasis>
, 1983
</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBFD901FC38AC76DB29FA8E" author="Gunnell GF &amp; Bown TM &amp; Bloch JI &amp; Boyer DM" box="[896,1123,1355,1377]" pageId="13" pageNumber="444" pagination="63 - 81" refId="ref17916" refString="Gunnell GF, Bown TM, Bloch JI, Boyer DM. 2008. Proteutheria. In: Janis CM, Gunnell GF, Uhen MD, eds. Evolution of tertiary mammals of North America, Vol. 2: small mammals, Xenarthrans, and marine mammals. Cambridge: Cambridge University Press, 63 - 81." type="book chapter" year="2008">
Gunnell
<emphasis id="B96BEA9AFFBFD901FC5DAC71DB54FA8E" box="[997,1054,1355,1377]" italics="true" pageId="13" pageNumber="444">et al.</emphasis>
, 2008
</bibRefCitation>
).
<bibRefCitation id="EF8E4B79FFBFD901FBC1AC76DA66FA8F" author="Simpson GG" box="[1145,1324,1355,1377]" pageId="13" pageNumber="444" pagination="1 - 287" refId="ref20798" refString="Simpson GG. 1937. The Fort Union of the Crazy Mountain Field, Montana and its mammalian faunas. Bulletin of the United States National Museum 169: 1 - 287." type="journal article" year="1937">Simpson (1937)</bibRefCitation>
originally restricted Pentacodontinae (then considered a subfamily) to
<taxonomicName id="4C1F4D0BFFBFD901FC09ACB5DBACFA70" authority="Simpson, 1935" authorityName="Simpson" authorityYear="1935" box="[945,1254,1416,1438]" class="Mammalia" family="Pantolestidae" genus="Aphronorus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="13" pageNumber="444" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBFD901FC09ACB5DB72FA73" box="[945,1080,1416,1437]" italics="true" pageId="13" pageNumber="444">Aphronorus</emphasis>
<bibRefCitation id="EF8E4B79FFBFD901FB87ACB5DBACFA70" author="Simpson GG" box="[1087,1254,1416,1438]" pageId="13" pageNumber="444" pagination="221 - 244" refId="ref20746" refString="Simpson GG. 1935. New Paleocene mammals from the Fort Union of Montana. Proceedings of the United States National Museum 83: 221 - 244." type="journal article" year="1935">Simpson, 1935</bibRefCitation>
</taxonomicName>
and
<taxonomicName id="4C1F4D0BFFBFD901FA98ACB5DCF5FA52" authority="Scott, 1892" authorityName="Scott" authorityYear="1892" class="Mammalia" family="Pantolestidae" genus="Pentacodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="13" pageNumber="444" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBFD901FA98ACB5DAE8FA73" box="[1312,1442,1416,1437]" italics="true" pageId="13" pageNumber="444">Pentacodon</emphasis>
<bibRefCitation id="EF8E4B79FFBFD901FC83AC9ADCF5FA52" author="Scott WB" box="[827,959,1447,1468]" pageId="13" pageNumber="444" pagination="291 - 323" refId="ref20200" refString="Scott WB. 1892. A revision of the North American Creodonta with notes on some genera which have been referred to that group. Proceedings of the Academy of Natural Sciences of Philadelphia 44: 291 - 323." type="journal article" year="1892">Scott, 1892</bibRefCitation>
</taxonomicName>
, but the family has since been expanded to include several other North American and, potentially, European taxa (
<bibRefCitation id="EF8E4B79FFBFD901FBEDACD8DA3AFA17" author="McKenna MC &amp; Bell SK" box="[1109,1392,1508,1530]" pageId="13" pageNumber="444" refId="ref19265" refString="McKenna MC, Bell SK. 1997. Classification of mammals above the species level. New York: Columbia University Press." type="book" year="1997">McKenna &amp; Bell, 1997</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBFD901FA3AACD8DB67F9F6" author="De Bast E &amp; Smith T" pageId="13" pageNumber="444" pagination="741 - 785" refId="ref16952" refString="De Bast E, Smith T. 2017. The oldest Cenozoic mammal fauna of Europe: implication of the Hainin reference fauna for mammalian evolution and dispersals during the Paleocene. Journal of Systematic Palaeontology 15: 741 - 785." type="journal article" year="2017">De Bast &amp; Smith, 2017</bibRefCitation>
). The broader relationships of
<taxonomicName id="4C1F4D0BFFBFD901FC83AF1CDCBEF9D9" authorityName="Simpson" authorityYear="1937" box="[827,1012,1569,1591]" class="Mammalia" family="Pentacodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pantolesta" pageId="13" pageNumber="444" phylum="Chordata" rank="family">Pentacodontidae</taxonomicName>
remain unclear, although the opinions of
<bibRefCitation id="EF8E4B79FFBFD901FCEDAF7DDCB4F9BB" author="Matthew WD" box="[853,1022,1600,1622]" pageId="13" pageNumber="444" pagination="291 - 567" refId="ref19031" refString="Matthew WD. 1909. The Carnivora and Insectivora of the Bridger basin, Middle Eocene. Memoir of the American Museum of Natural History 9: 291 - 567." type="journal article" year="1909">Matthew (1909</bibRefCitation>
,
<bibRefCitation id="EF8E4B79FFBFD901FBB2AF7DDB0AF9BB" author="Matthew WD" box="[1034,1088,1600,1621]" pageId="13" pageNumber="444" pagination="565 - 657" refId="ref19105" refString="Matthew WD. 1918. A revision of the lower Eocene Wasatch and Wind River faunas. Bulletin of the American Museum of Natural History 38: 565 - 657." type="journal article" year="1918">1918</bibRefCitation>
),
<bibRefCitation id="EF8E4B79FFBFD901FBEBAF7DDA4AF9B8" author="Simpson GG" box="[1107,1280,1600,1622]" pageId="13" pageNumber="444" pagination="1 - 287" refId="ref20798" refString="Simpson GG. 1937. The Fort Union of the Crazy Mountain Field, Montana and its mammalian faunas. Bulletin of the United States National Museum 169: 1 - 287." type="journal article" year="1937">Simpson (1937)</bibRefCitation>
and
<bibRefCitation id="EF8E4B79FFBFD901FA8EAF7CDCC8F99B" author="McKenna MC" pageId="13" pageNumber="444" pagination="21 - 46" refId="ref19226" refString="McKenna MC. 1975. Toward a phylogenetic classification of the Mammalia. In: Luckett WP, Szalay FS, eds. Phylogeny of the Primates. New York: Plenum Publishing Company, 21 - 46." type="book chapter" year="1975">McKenna (1975)</bibRefCitation>
that link the group with pantolestids, as either a subfamily of
<taxonomicName id="4C1F4D0BFFBFD901FC5AAF40DB36F97D" authorityName="Cope" authorityYear="1884" box="[994,1148,1661,1683]" class="Mammalia" family="Pantolestidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="13" pageNumber="444" phylum="Chordata" rank="family">Pantolestidae</taxonomicName>
or a family within a more inclusive
<taxonomicName id="4C1F4D0BFFBFD901FC1CAFA1DB56F95C" authorityName="McKenna" authorityYear="1975" box="[932,1052,1692,1714]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pantolesta" pageId="13" pageNumber="444" phylum="Chordata" rank="order">Pantolesta</taxonomicName>
, have been accepted in recent studies (e.g.
<bibRefCitation id="EF8E4B79FFBFD901FC2FAF86DBD0F93E" author="McKenna MC &amp; Bell SK" box="[919,1178,1723,1745]" pageId="13" pageNumber="444" refId="ref19265" refString="McKenna MC, Bell SK. 1997. Classification of mammals above the species level. New York: Columbia University Press." type="book" year="1997">McKenna &amp; Bell, 1997</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBFD901FB1FAF86DAD7F93E" author="Boyer DM &amp; Georgi JA" box="[1191,1437,1723,1745]" pageId="13" pageNumber="444" pagination="239 - 280" refId="ref16226" refString="Boyer DM, Georgi JA. 2007. Cranial morphology of a pantolestid eutherian mammal from the Eocene Bridger Formation, Wyoming, USA: implications for relationships and habitat. Journal of Mammalian Evolution 14: 239 - 280." type="journal article" year="2007">Boyer &amp; Georgi, 2007</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBFD901FC83AFE4DC85F901" author="Rankin BD" box="[827,975,1753,1775]" pageId="13" pageNumber="444" pagination="1199 - 1211" refId="ref19728" refString="Rankin BD. 2014. New pantolestids (Mammalia, Eutheria) from the Late Paleocene (Late Middle Tiffanian) Roche Percee local fauna, southeastern Saskatchewan, Canada. Journal of Paleontology 88: 1199 - 1211." type="journal article" year="2014">Rankin, 2014</bibRefCitation>
). Scott
<emphasis id="B96BEA9AFFBFD901FB99AFE7DB12F900" box="[1057,1112,1753,1775]" italics="true" pageId="13" pageNumber="444">et al.</emphasis>
(2006) noted several similarities in the dentitions of
<taxonomicName id="4C1F4D0BFFBFD901FBF4AFC5DB97F8E3" authorityName="Scott, Fox &amp; Webb" authorityYear="2006" box="[1100,1245,1784,1805]" class="Mammalia" family="Horolodectidae" genus="Horolodectes" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="13" pageNumber="444" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBFD901FBF4AFC5DB97F8E3" box="[1100,1245,1784,1805]" italics="true" pageId="13" pageNumber="444">Horolodectes</emphasis>
</taxonomicName>
and
<taxonomicName id="4C1F4D0BFFBFD901FAAEAFC5DAD6F8E3" authorityName="Simpson" authorityYear="1935" box="[1302,1436,1784,1805]" class="Mammalia" family="Pantolestidae" genus="Aphronorus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="13" pageNumber="444" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBFD901FAAEAFC5DAD6F8E3" box="[1302,1436,1784,1805]" italics="true" pageId="13" pageNumber="444">Aphronorus</emphasis>
</taxonomicName>
, and the genus is therefore included in this study.
<taxonomicName id="4C1F4D0BFFBFD901FC83AE08DC8FF8A4" authorityName="Simpson" authorityYear="1935" box="[827,965,1845,1866]" class="Mammalia" family="Pantolestidae" genus="Aphronorus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="13" pageNumber="444" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBFD901FC83AE08DC8FF8A4" box="[827,965,1845,1866]" italics="true" pageId="13" pageNumber="444">Aphronorus</emphasis>
</taxonomicName>
was scored using
<taxonomicName id="4C1F4D0BFFBFD901FB26AE0BDC39F884" authority="Simpson, 1935" authorityName="Simpson" authorityYear="1935" class="Mammalia" family="Pantolestidae" genus="Aphronorus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="13" pageNumber="444" phylum="Chordata" rank="species" species="fraudator">
<emphasis id="B96BEA9AFFBFD901FB26AE0BDA65F8A4" box="[1182,1327,1845,1867]" italics="true" pageId="13" pageNumber="444">A. fraudator</emphasis>
<bibRefCitation id="EF8E4B79FFBFD901FA80AE08DC39F884" author="Simpson GG" pageId="13" pageNumber="444" pagination="221 - 244" refId="ref20746" refString="Simpson GG. 1935. New Paleocene mammals from the Fort Union of Montana. Proceedings of the United States National Museum 83: 221 - 244." type="journal article" year="1935">Simpson, 1935</bibRefCitation>
</taxonomicName>
, the
<typeStatus id="54A4882AFFBFD901FC1BAE68DC9EF884" box="[931,980,1877,1898]" pageId="13" pageNumber="444">type</typeStatus>
species, and
<taxonomicName id="4C1F4D0BFFBFD901FBE6AE69DA71F884" authority="Gazin, 1969" authorityName="Gazin" authorityYear="1969" box="[1118,1339,1876,1898]" class="Mammalia" family="Pantolestidae" genus="Aphronorus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="13" pageNumber="444" phylum="Chordata" rank="species" species="orieli">
<emphasis id="B96BEA9AFFBFD901FBE6AE69DB25F887" box="[1118,1135,1876,1897]" italics="true" pageId="13" pageNumber="444">A</emphasis>
.
<emphasis id="B96BEA9AFFBFD901FBC1AE69DBF9F887" box="[1145,1203,1876,1897]" italics="true" pageId="13" pageNumber="444">orieli</emphasis>
<bibRefCitation id="EF8E4B79FFBFD901FB0FAE69DA71F884" author="Gazin CL" box="[1207,1339,1876,1898]" pageId="13" pageNumber="444" pagination="1 - 16" refId="ref17510" refString="Gazin CL. 1969. A new occurrence of Paleocene mammals in the Evanston Formation, southwestern Wyoming. Smithsonian Contributions to Paleobiology 2: 1 - 16." type="journal article" year="1969">Gazin, 1969</bibRefCitation>
</taxonomicName>
. A second putative pentacodontid,
<taxonomicName id="4C1F4D0BFFBCD902FE1FA9F8DD8BFF35" authority="Gazin, 1956" authorityName="Gazin" authorityYear="1956" box="[423,705,197,219]" class="Mammalia" family="Pantolestidae" genus="Bisonalveus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="14" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBCD902FE1FA9F8DD7AFF34" box="[423,560,197,218]" italics="true" pageId="14" pageNumber="445">Bisonalveus</emphasis>
<bibRefCitation id="EF8E4B79FFBCD902FD8FA9F8DD8BFF35" author="Gazin CL" box="[567,705,197,219]" pageId="14" pageNumber="445" pagination="1 - 57" refId="ref17485" refString="Gazin CL. 1956. Paleocene mammalian faunas of the Bison Basin in south-central Wyoming. Smithsonian Miscellaneous Collections 131: 1 - 57." type="journal article" year="1956">Gazin, 1956</bibRefCitation>
</taxonomicName>
, was also included in this analysis. In the parts that can be compared, the dentition of this genus is similar to that of
<taxonomicName id="4C1F4D0BFFBCD902FF14A81CDE2CFED8" authorityName="Scott" authorityYear="2019" box="[172,358,289,310]" class="Mammalia" family="Zhelestidae" genus="Ferrequitherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Proteutheria" pageId="14" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBCD902FF14A81CDE2CFED8" box="[172,358,289,310]" italics="true" pageId="14" pageNumber="445">Ferrequitherium</emphasis>
</taxonomicName>
, particularly the upper molars and lower fourth premolar.
<taxonomicName id="4C1F4D0BFFBCD902FE2FA87DDD55FEBB" authorityName="Gazin" authorityYear="1956" box="[407,543,320,341]" class="Mammalia" family="Pantolestidae" genus="Bisonalveus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="14" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBCD902FE2FA87DDD55FEBB" box="[407,543,320,341]" italics="true" pageId="14" pageNumber="445">Bisonalveus</emphasis>
</taxonomicName>
has long been considered a pentacodontid (e.g.
<bibRefCitation id="EF8E4B79FFBCD902FE67A862DDD2FE9A" author="Van Valen L" box="[479,664,350,372]" pageId="14" pageNumber="445" pagination="217 - 284" refId="ref21261" refString="Van Valen L. 1967. New Paleocene insectivores and insectivore classification. Bulletin of the American Museum of Natural History 135: 217 - 284." type="journal article" year="1967">Van Valen, 1967</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FD1EA862DE26FE7C" author="Gingerich PD &amp; Houde P &amp; Krause DW" pageId="14" pageNumber="445" pagination="957 - 970" refId="ref17776" refString="Gingerich PD, Houde P, Krause DW. 1983. A new earliest Tiffanian (Late Paleocene) mammalian fauna from Bangtail Plateau, western Crazy Mountain Basin, Montana. Journal of Paleontology 57: 957 - 970." type="journal article" year="1983">Krause &amp; Gingerich, 1983</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FEC2A840DD1CFE7C" author="Youzwyshyn GP" box="[378,598,381,403]" pageId="14" pageNumber="445" refId="ref21847" refString="Youzwyshyn GP. 1988. Late Paleocene mammals from near Cochrane, southwestern Alberta. Unpublished M. S. Thesis, University of Alberta." type="book" year="1988">Youzwyshyn, 1988</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FDDDA843DF8DFE5C" author="Fox RC &amp; Scott CS" pageId="14" pageNumber="445" refId="ref17342" refString="Fox RC, Scott CS. 2005. First evidence of a venom delivery" type="book" year="2005">Fox &amp; Scott, 2005</bibRefCitation>
), although its evolutionary relationships have not been examined in detail, and at least one recent, largescale analysis has hypothesized a bizarre sister-group relationship between the genus and a clade, including pantodonts and tillodonts (
<bibRefCitation id="EF8E4B79FFBCD902FE68AB2BDDF4FDC2" author="Halliday TJ &amp; Upchurch P &amp; Goswami A" box="[464,702,534,556]" pageId="14" pageNumber="445" pagination="521 - 550" refId="ref18053" refString="Halliday TJ, Upchurch P, Goswami A. 2017. Resolving the relationships of Paleocene placental mammals. Biological Reviews of the Cambridge Philosophical Society 92: 521 - 550." type="journal article" year="2017">
Halliday
<emphasis id="B96BEA9AFFBCD902FD85AB2ADD32FDC5" box="[573,632,534,556]" italics="true" pageId="14" pageNumber="445">et al.</emphasis>
, 2017
</bibRefCitation>
; and see
<bibRefCitation id="EF8E4B79FFBCD902FF04AB08DE7CFDA4" author="Zack SP" box="[188,310,565,587]" pageId="14" pageNumber="445" refId="ref21873" refString="Zack SP. 2009. The phylogeny of eutherian mammals: a new analysis emphasizing dental and postcranial morphology of Paleogene taxa. Unpublished Ph. D. Dissertation, The Johns Hopkins University School of Medicine." type="book" year="2009">Zack, 2009</bibRefCitation>
for discussion).
<taxonomicName id="4C1F4D0BFFBCD902FE56AB08DD3CFDA4" authorityName="Gazin" authorityYear="1956" box="[494,630,565,586]" class="Mammalia" family="Pantolestidae" genus="Bisonalveus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="14" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBCD902FE56AB08DD3CFDA4" box="[494,630,565,586]" italics="true" pageId="14" pageNumber="445">Bisonalveus</emphasis>
</taxonomicName>
was scored using
<taxonomicName id="4C1F4D0BFFBCD902FF61AB69DD1EFD84" authority="Gazin, 1956" authorityName="Gazin" authorityYear="1956" box="[217,596,596,618]" class="Mammalia" family="Pantolestidae" genus="Bisonalveus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="14" pageNumber="445" phylum="Chordata" rank="species" species="browni">
<emphasis id="B96BEA9AFFBCD902FF61AB69DEF4FD87" box="[217,446,596,617]" italics="true" pageId="14" pageNumber="445">Bisonalveus browni</emphasis>
<bibRefCitation id="EF8E4B79FFBCD902FE7FAB69DD1EFD84" author="Gazin CL" box="[455,596,596,618]" pageId="14" pageNumber="445" pagination="1 - 57" refId="ref17485" refString="Gazin CL. 1956. Paleocene mammalian faunas of the Bison Basin in south-central Wyoming. Smithsonian Miscellaneous Collections 131: 1 - 57." type="journal article" year="1956">Gazin, 1956</bibRefCitation>
</taxonomicName>
, the
<typeStatus id="54A4882AFFBCD902FD37AB69DD8BFD87" box="[655,705,596,617]" pageId="14" pageNumber="445">type</typeStatus>
species, and an as-yet unnamed new species from the late Palaeocene of
<collectingRegion id="49DBF86AFFBCD902FE95ABACDECBFD49" box="[301,385,657,679]" country="Canada" name="Alberta" pageId="14" pageNumber="445">Alberta</collectingRegion>
(
<bibRefCitation id="EF8E4B79FFBCD902FE37ABACDD41FD48" author="Scott CS" box="[399,523,657,678]" pageId="14" pageNumber="445" refId="ref20268" refString="Scott CS. 2008. Late Paleocene mammals from near Red Deer, Alberta, and a phylogenetic analysis of the earliest Lipotyphla (Mammalia, Insectivora). Unpublished Ph. D. Dissertation, University of Alberta." type="book" year="2008">Scott, 2008</bibRefCitation>
).
</paragraph>
<paragraph id="8BA03688FFBCD902FF29ABF5DE28FB47" blockId="14.[145,762,712,1193]" pageId="14" pageNumber="445">
<taxonomicName id="4C1F4D0BFFBCD902FF29ABF5DE90FD33" authority="Simpson, 1936" authorityName="Simpson" authorityYear="1936" box="[145,474,712,733]" class="Mammalia" family="Pantolestidae" genus="Bessoecetor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="14" pageNumber="445" phylum="Chordata" rank="genus">
Bessoecetor
<bibRefCitation id="EF8E4B79FFBCD902FE91ABF5DE90FD33" author="Simpson GG" box="[297,474,712,733]" pageId="14" pageNumber="445" pagination="1 - 27" refId="ref20774" refString="Simpson GG. 1936. A new fauna from the Fort Union of Montana. American Museum Novitates 873: 1 - 27." type="journal article" year="1936">
<emphasis id="B96BEA9AFFBCD902FE91ABF5DE90FD33" box="[297,474,712,733]" italics="true" pageId="14" pageNumber="445">Simpson, 1936</emphasis>
</bibRefCitation>
</taxonomicName>
<emphasis id="B96BEA9AFFBCD902FE64ABF5DD39FD33" box="[476,627,712,733]" italics="true" pageId="14" pageNumber="445">
:
<taxonomicName id="4C1F4D0BFFBCD902FE53ABF5DD39FD33" authorityName="Simpson" authorityYear="1936" box="[491,627,712,733]" class="Mammalia" family="Pantolestidae" genus="Bessoecetor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="14" pageNumber="445" phylum="Chordata" rank="genus">Bessoecetor</taxonomicName>
</emphasis>
is the oldest discovered pantolestid, known principally from Palaeocene deposits in the northern part of the Western Interior of North America (
<bibRefCitation id="EF8E4B79FFBCD902FDF7AA19DDB8FCD7" author="Russell LS" box="[591,754,804,826]" pageId="14" pageNumber="445" pagination="162 - 178" refId="ref20073" refString="Russell LS. 1929. Paleocene vertebrates from Alberta. American Journal of Science 17: 162 - 178." type="journal article" year="1929">Russell, 1929</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FF29AA7FDE0FFCB6" author="Simpson GG" box="[145,325,834,856]" pageId="14" pageNumber="445" pagination="1 - 27" refId="ref20774" refString="Simpson GG. 1936. A new fauna from the Fort Union of Montana. American Museum Novitates 873: 1 - 27." type="journal article" year="1936">Simpson, 1936</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FEEEAA7EDDD6FCB6" author="Gingerich PD &amp; Houde P &amp; Krause DW" box="[342,668,834,856]" pageId="14" pageNumber="445" pagination="957 - 970" refId="ref17776" refString="Gingerich PD, Houde P, Krause DW. 1983. A new earliest Tiffanian (Late Paleocene) mammalian fauna from Bangtail Plateau, western Crazy Mountain Basin, Montana. Journal of Paleontology 57: 957 - 970." type="journal article" year="1983">Krause &amp; Gingerich, 1983</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FD15AA7FDE5DFC99" author="Cifelli RL &amp; Czaplewski NJ &amp; Rose KD" pageId="14" pageNumber="445" pagination="304 - 315" refId="ref16403" refString="Cifelli RL, Czaplewski NJ, Rose KD. 1995. Additions to knowledge of Paleocene mammals from the North Horn Formation, central Utah. Great Basin Naturalist 55: 304 - 315." type="journal article" year="1995">
Cifelli
<emphasis id="B96BEA9AFFBCD902FF29AA5FDF84FC98" box="[145,206,865,887]" italics="true" pageId="14" pageNumber="445">et al.</emphasis>
, 1995
</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FE9FAA5CDEB9FC98" author="Scott CS &amp; Fox RC &amp; Youzwyshyn GP" box="[295,499,865,887]" pageId="14" pageNumber="445" pagination="691 - 704" refId="ref20471" refString="Scott CS, Fox RC, Youzwyshyn GP. 2002. New earliest Tiffanian (Late Paleocene) mammals from Cochrane 2, southwestern Alberta, Canada. Acta Palaeontologica Polonica 47: 691 - 704." type="journal article" year="2002">
Scott
<emphasis id="B96BEA9AFFBCD902FED6AA5FDEE0FC98" box="[366,426,865,887]" italics="true" pageId="14" pageNumber="445">et al.</emphasis>
, 2002
</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FDBCAA5CDDC0FC98" author="Scott CS" box="[516,650,865,886]" pageId="14" pageNumber="445" refId="ref20268" refString="Scott CS. 2008. Late Paleocene mammals from near Red Deer, Alberta, and a phylogenetic analysis of the earliest Lipotyphla (Mammalia, Insectivora). Unpublished Ph. D. Dissertation, University of Alberta." type="book" year="2008">Scott, 2008</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FD22AA5CDF81FC7B" author="Rankin BD" pageId="14" pageNumber="445" pagination="1199 - 1211" refId="ref19728" refString="Rankin BD. 2014. New pantolestids (Mammalia, Eutheria) from the Late Paleocene (Late Middle Tiffanian) Roche Percee local fauna, southeastern Saskatchewan, Canada. Journal of Paleontology 88: 1199 - 1211." type="journal article" year="2014">Rankin, 2014</bibRefCitation>
). The genus was included in this analysis to test for potential broader pantolestan (i.e. exclusive of
<taxonomicName id="4C1F4D0BFFBCD902FF15AA80DE27FC3D" authorityName="Simpson" authorityYear="1937" box="[173,365,957,979]" class="Mammalia" family="Pentacodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pantolesta" pageId="14" pageNumber="445" phylum="Chordata" rank="family">Pentacodontidae</taxonomicName>
) affinities of
<taxonomicName id="4C1F4D0BFFBCD902FDB0AA80DD8CFC3C" authorityName="Scott" authorityYear="2019" box="[520,710,957,978]" class="Mammalia" family="Zhelestidae" genus="Ferrequitherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Proteutheria" pageId="14" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBCD902FDB0AA80DD8CFC3C" box="[520,710,957,978]" italics="true" pageId="14" pageNumber="445">Ferrequitherium</emphasis>
</taxonomicName>
and
<taxonomicName id="4C1F4D0BFFBCD902FF29AAE1DE6BFC1F" authorityName="Scott, Fox &amp; Webb" authorityYear="2006" box="[145,289,988,1009]" class="Mammalia" family="Horolodectidae" genus="Horolodectes" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="14" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBCD902FF29AAE1DE6BFC1F" box="[145,289,988,1009]" italics="true" pageId="14" pageNumber="445">Horolodectes</emphasis>
</taxonomicName>
.
<taxonomicName id="4C1F4D0BFFBCD902FE95AAE1DEE7FC1F" authorityName="Simpson" authorityYear="1936" box="[301,429,988,1009]" class="Mammalia" family="Pantolestidae" genus="Bessoecetor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="14" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBCD902FE95AAE1DEE7FC1F" box="[301,429,988,1009]" italics="true" pageId="14" pageNumber="445">Bessoecetor</emphasis>
</taxonomicName>
was scored using
<taxonomicName id="4C1F4D0BFFBCD902FDC1AAE1DEAFFBFE" authority="(Russell, 1929)" baseAuthorityName="Russell" baseAuthorityYear="1929" class="Mammalia" family="Pantolestidae" genus="Bessoecetor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="14" pageNumber="445" phylum="Chordata" rank="species" species="septentrionalis">
<emphasis id="B96BEA9AFFBCD902FDC1AAE1DE73FBE1" italics="true" pageId="14" pageNumber="445">Bessoecetor septentrionalis</emphasis>
(
<bibRefCitation id="EF8E4B79FFBCD902FEFEAAC7DE97FBFE" author="Russell LS" box="[326,477,1018,1040]" pageId="14" pageNumber="445" pagination="162 - 178" refId="ref20073" refString="Russell LS. 1929. Paleocene vertebrates from Alberta. American Journal of Science 17: 162 - 178." type="journal article" year="1929">Russell, 1929</bibRefCitation>
)
</taxonomicName>
: the species is one of the oldest and arguably most basal of the genus, and is particularly well documented, being known from dozens of specimens from
<collectingRegion id="49DBF86AFFBCD902FE2FAD6BDEA4FB82" box="[407,494,1110,1132]" country="Canada" name="Alberta" pageId="14" pageNumber="445">Alberta</collectingRegion>
and
<collectingRegion id="49DBF86AFFBCD902FD91AD6ADDDBFB82" box="[553,657,1111,1132]" country="United States of America" name="Montana" pageId="14" pageNumber="445">Montana</collectingRegion>
(see, e.g.
<bibRefCitation id="EF8E4B79FFBCD902FF29AD48DE7DFB65" author="Simpson GG" box="[145,311,1141,1163]" pageId="14" pageNumber="445" pagination="1 - 27" refId="ref20774" refString="Simpson GG. 1936. A new fauna from the Fort Union of Montana. American Museum Novitates 873: 1 - 27." type="journal article" year="1936">Simpson, 1936</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FEFCAD48DD27FB64" author="Gingerich PD &amp; Houde P &amp; Krause DW" box="[324,621,1141,1163]" pageId="14" pageNumber="445" pagination="957 - 970" refId="ref17776" refString="Gingerich PD, Houde P, Krause DW. 1983. A new earliest Tiffanian (Late Paleocene) mammalian fauna from Bangtail Plateau, western Crazy Mountain Basin, Montana. Journal of Paleontology 57: 957 - 970." type="journal article" year="1983">Krause &amp; Gingerich, 1983</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FDC2AD48DF82FB47" author="Scott CS &amp; Fox RC &amp; Youzwyshyn GP" pageId="14" pageNumber="445" pagination="691 - 704" refId="ref20471" refString="Scott CS, Fox RC, Youzwyshyn GP. 2002. New earliest Tiffanian (Late Paleocene) mammals from Cochrane 2, southwestern Alberta, Canada. Acta Palaeontologica Polonica 47: 691 - 704." type="journal article" year="2002">
Scott
<emphasis id="B96BEA9AFFBCD902FD03AD4BDDB9FB64" box="[699,755,1141,1163]" italics="true" pageId="14" pageNumber="445">et al.</emphasis>
, 2002
</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FF6CADA9DE18FB47" author="Scott CS" box="[212,338,1172,1193]" pageId="14" pageNumber="445" refId="ref20268" refString="Scott CS. 2008. Late Paleocene mammals from near Red Deer, Alberta, and a phylogenetic analysis of the earliest Lipotyphla (Mammalia, Insectivora). Unpublished Ph. D. Dissertation, University of Alberta." type="book" year="2008">Scott, 2008</bibRefCitation>
).
</paragraph>
<paragraph id="8BA03688FFBCD902FF29ADEDDD8BF9F6" blockId="14.[145,761,1232,1560]" pageId="14" pageNumber="445">
<taxonomicName id="4C1F4D0BFFBCD902FF29ADEDDEFEFB0B" authority="Marsh, 1889" authorityName="Marsh" authorityYear="1889" box="[145,436,1232,1254]" class="Mammalia" family="Cimolestidae" genus="Cimolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="14" pageNumber="445" phylum="Chordata" rank="genus">
Cimolestes
<bibRefCitation id="EF8E4B79FFBCD902FEA4ADEDDEFEFB0B" author="Marsh OC" box="[284,436,1232,1254]" pageId="14" pageNumber="445" pagination="81 - 92" refId="ref19011" refString="Marsh OC. 1889. Discovery of Cretaceous Mammalia. American Journal of Science 38: 81 - 92." type="journal article" year="1889">Marsh, 1889</bibRefCitation>
</taxonomicName>
: The genus
<taxonomicName id="4C1F4D0BFFBCD902FDFCADEDDD88FB0B" authorityName="Marsh" authorityYear="1889" box="[580,706,1232,1253]" class="Mammalia" family="Cimolestidae" genus="Cimolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="14" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBCD902FDFCADEDDD88FB0B" box="[580,706,1232,1253]" italics="true" pageId="14" pageNumber="445">Cimolestes</emphasis>
</taxonomicName>
was recently split into several genera, in recognition of the morphological disparity among its included species (
<bibRefCitation id="EF8E4B79FFBCD902FF75AC11DE76FAAF" author="Fox RC" box="[205,316,1323,1345]" pageId="14" pageNumber="445" pagination="1137 - 1149" refId="ref17311" refString="Fox RC. 2015. A revision of the Late Cretaceous - Paleocene eutherian mammal Cimolestes Marsh, 1889. Canadian Journal of Earth Sciences 52: 1137 - 1149." type="journal article" year="2015">Fox, 2015</bibRefCitation>
).
<bibRefCitation id="EF8E4B79FFBCD902FEEBAC11DE84FAAF" author="Fox RC" box="[339,462,1323,1345]" pageId="14" pageNumber="445" pagination="1137 - 1149" refId="ref17311" refString="Fox RC. 2015. A revision of the Late Cretaceous - Paleocene eutherian mammal Cimolestes Marsh, 1889. Canadian Journal of Earth Sciences 52: 1137 - 1149." type="journal article" year="2015">Fox (2015)</bibRefCitation>
limited
<taxonomicName id="4C1F4D0BFFBCD902FD89AC11DDE7FAAF" authorityName="Marsh" authorityYear="1889" box="[561,685,1324,1345]" class="Mammalia" family="Cimolestidae" genus="Cimolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="14" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBCD902FD89AC11DDE7FAAF" box="[561,685,1324,1345]" italics="true" pageId="14" pageNumber="445">Cimolestes</emphasis>
</taxonomicName>
to the
<typeStatus id="54A4882AFFBCD902FF29AC76DF8EFA8E" box="[145,196,1355,1376]" pageId="14" pageNumber="445">type</typeStatus>
species,
<taxonomicName id="4C1F4D0BFFBCD902FE96AC76DD75FAB1" authority="Marsh, 1889" authorityName="Marsh" authorityYear="1889" box="[302,575,1354,1376]" class="Mammalia" family="Cimolestidae" genus="Cimolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="14" pageNumber="445" phylum="Chordata" rank="species" species="incisus">
<emphasis id="B96BEA9AFFBCD902FE96AC76DE0AFA8E" box="[302,320,1355,1376]" italics="true" pageId="14" pageNumber="445">C</emphasis>
.
<emphasis id="B96BEA9AFFBCD902FEF7AC76DEEBFA8E" box="[335,417,1355,1376]" italics="true" pageId="14" pageNumber="445">incisus</emphasis>
<bibRefCitation id="EF8E4B79FFBCD902FE12AC77DD75FAB1" author="Marsh OC" box="[426,575,1354,1376]" pageId="14" pageNumber="445" pagination="81 - 92" refId="ref19011" refString="Marsh OC. 1889. Discovery of Cretaceous Mammalia. American Journal of Science 38: 81 - 92." type="journal article" year="1889">Marsh, 1889</bibRefCitation>
</taxonomicName>
, and
<taxonomicName id="4C1F4D0BFFBCD902FD39AC76DE0BFA90" authority="Clemens, 1973" authorityName="Clemens" authorityYear="1973" class="Mammalia" family="Cimolestidae" genus="Cimolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="14" pageNumber="445" phylum="Chordata" rank="species" species="stirtoni">
<emphasis id="B96BEA9AFFBCD902FD39AC76DDD9FA8E" box="[641,659,1355,1376]" italics="true" pageId="14" pageNumber="445">C</emphasis>
.
<emphasis id="B96BEA9AFFBCD902FD1AAC76DDB3FA8E" box="[674,761,1355,1376]" italics="true" pageId="14" pageNumber="445">stirtoni</emphasis>
<bibRefCitation id="EF8E4B79FFBCD902FF29AC54DE0BFA90" author="Clemens WA" box="[145,321,1385,1407]" pageId="14" pageNumber="445" pagination="1 - 102" refId="ref16470" refString="Clemens WA. 1973. Fossil mammals of the type Lance Formation Wyoming: part III. Eutheria and summary. University of California Publications in Geological Sciences 94: 1 - 102." type="journal article" year="1973">Clemens, 1973</bibRefCitation>
</taxonomicName>
, both from Late Cretaceous deposits of the Western Interior of North America, with the former being included in this analysis.
<taxonomicName id="4C1F4D0BFFBCD902FDD8AC9BDD97FA55" authorityName="Marsh" authorityYear="1889" box="[608,733,1446,1467]" class="Mammalia" family="Cimolestidae" genus="Cimolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="14" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBCD902FDD8AC9BDD97FA55" box="[608,733,1446,1467]" italics="true" pageId="14" pageNumber="445">Cimolestes</emphasis>
</taxonomicName>
is considered a basal cimolestan (
<bibRefCitation id="EF8E4B79FFBCD902FE49ACF8DDBBFA34" author="McKenna MC &amp; Bell SK" box="[497,753,1477,1499]" pageId="14" pageNumber="445" refId="ref19265" refString="McKenna MC, Bell SK. 1997. Classification of mammals above the species level. New York: Columbia University Press." type="book" year="1997">McKenna &amp; Bell, 1997</bibRefCitation>
) and was included in the analysis to test for broader cimolestan (i.e. exclusive of
<taxonomicName id="4C1F4D0BFFBCD902FE74AF3FDD0CF9F6" authorityName="McKenna" authorityYear="1975" box="[460,582,1538,1560]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pantolesta" pageId="14" pageNumber="445" phylum="Chordata" rank="order">Pantolesta</taxonomicName>
) affinities.
</paragraph>
<paragraph id="8BA03688FFBCD902FF29AF03DB86FC16" blockId="14.[145,762,1598,1896]" lastBlockId="14.[809,1426,197,1016]" pageId="14" pageNumber="445">
<taxonomicName id="4C1F4D0BFFBCD902FF29AF03DE95F9BD" authority="Matthew, 1918" authorityName="Matthew" authorityYear="1918" box="[145,479,1598,1620]" class="Mammalia" family="Apheliscidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Macroscelidea" pageId="14" pageNumber="445" phylum="Chordata" rank="family">
<emphasis id="B96BEA9AFFBCD902FF29AF03DE6CF9BD" box="[145,294,1598,1619]" italics="true" pageId="14" pageNumber="445">Apheliscidae</emphasis>
<bibRefCitation id="EF8E4B79FFBCD902FE96AF03DE95F9BD" author="Matthew WD" box="[302,479,1598,1620]" pageId="14" pageNumber="445" pagination="565 - 657" refId="ref19105" refString="Matthew WD. 1918. A revision of the lower Eocene Wasatch and Wind River faunas. Bulletin of the American Museum of Natural History 38: 565 - 657." type="journal article" year="1918">Matthew, 1918</bibRefCitation>
</taxonomicName>
: The
<taxonomicName id="4C1F4D0BFFBCD902FDA6AF03DDFEF9BA" authorityName="Matthew" authorityYear="1918" box="[542,692,1598,1620]" class="Mammalia" family="Apheliscidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Macroscelidea" pageId="14" pageNumber="445" phylum="Chordata" rank="family">Apheliscidae</taxonomicName>
are a family of small-bodied eutherians that have been traditionally allied with ‘Condylarthra’, a widely recognized wastebasket taxon consisting of primarily Palaeocene to Eocene age, bunodont mammals (
<bibRefCitation id="EF8E4B79FFBCD902FD83AF84DDB8F921" author="McKenna MC" box="[571,754,1721,1743]" pageId="14" pageNumber="445" pagination="1 - 130" refId="ref19193" refString="McKenna MC. 1960. Fossil Mammalia from the Early Wasatchian Four Mile fauna, Eocene of northwest Colorado. University of California Publications in Geological Sciences 37: 1 - 130." type="journal article" year="1960">McKenna, 1960</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FF29AFE5DE06F903" author="Van Valen L" box="[145,332,1751,1774]" pageId="14" pageNumber="445" pagination="217 - 284" refId="ref21261" refString="Van Valen L. 1967. New Paleocene insectivores and insectivore classification. Bulletin of the American Museum of Natural History 135: 217 - 284." type="journal article" year="1967">Van Valen, 1967</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FEE1AFE5DD5BF903" author="Archibald JD" box="[345,529,1752,1774]" pageId="14" pageNumber="445" pagination="292 - 331" refId="ref15846" refString="Archibald JD. 1998. Archaic ungulates (' Condylarthra'). In: Janis CM, Scott KM, Jacobs LL, eds. Evolution of tertiary mammals of North America, Vol. 1: terrestrial carnivores, ungulates, and ungulatelike mammals. Cambridge: Cambridge University Press, 292 - 331." type="book chapter" year="1998">Archibald, 1998</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FDA6AFE5DDA0F903" author="Zack SP &amp; Penkrot TA &amp; Krause DW &amp; Maas MC" box="[542,746,1751,1774]" pageId="14" pageNumber="445" pagination="809 - 830" refId="ref21947" refString="Zack SP, Penkrot TA, Krause DW, Maas MC. 2005 b. A new apheliscine ' condylarth' from the late Paleocene of Montana and Alberta and the phylogeny of ' hyopsodontids'. Acta Palaeontologica Polonica 50: 809 - 830." type="journal article" year="2005">
Zack
<emphasis id="B96BEA9AFFBCD902FDE5AFE5DDDDF903" box="[605,663,1752,1773]" italics="true" pageId="14" pageNumber="445">et al.</emphasis>
, 2005b
</bibRefCitation>
). ‘Condylarthra’ are thought to represent the initial radiation of ungulates, although compelling evidence linking any of the included lower-level ‘condylarth’ taxa with crown group ungulates remains elusive. A subset of
<taxonomicName id="4C1F4D0BFFBCD902FBA5A9F8DBF3FF35" authorityName="Matthew" authorityYear="1918" box="[1053,1209,197,219]" class="Mammalia" family="Apheliscidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Macroscelidea" pageId="14" pageNumber="445" phylum="Chordata" rank="family">Apheliscidae</taxonomicName>
, the Apheliscinae
<bibRefCitation id="EF8E4B79FFBCD902FC91A9DEDBA7FF17" author="Zack SP &amp; Penkrot TA &amp; Krause DW &amp; Maas MC" box="[809,1261,227,249]" pageId="14" pageNumber="445" pagination="809 - 830" refId="ref21947" refString="Zack SP, Penkrot TA, Krause DW, Maas MC. 2005 b. A new apheliscine ' condylarth' from the late Paleocene of Montana and Alberta and the phylogeny of ' hyopsodontids'. Acta Palaeontologica Polonica 50: 809 - 830." type="journal article" year="2005">Zack, Penkrot, Krause &amp; Maas, 2005b</bibRefCitation>
, is characterized by enlarged upper and lower fourth premolars, a weak p4 paraconid and metaconid, poorly developed molar cingulids, and inflated cusps (
<bibRefCitation id="EF8E4B79FFBCD902FB7DA802DAC0FEBB" author="Zack SP &amp; Penkrot TA &amp; Krause DW &amp; Maas MC" box="[1221,1418,319,341]" pageId="14" pageNumber="445" pagination="809 - 830" refId="ref21947" refString="Zack SP, Penkrot TA, Krause DW, Maas MC. 2005 b. A new apheliscine ' condylarth' from the late Paleocene of Montana and Alberta and the phylogeny of ' hyopsodontids'. Acta Palaeontologica Polonica 50: 809 - 830." type="journal article" year="2005">
Zack
<emphasis id="B96BEA9AFFBCD902FABAA87DDA70FEBA" box="[1282,1338,319,341]" italics="true" pageId="14" pageNumber="445">et al.</emphasis>
, 2005b
</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FC91A863DB5BFE9D" author="Penkrot TA &amp; Zack SP &amp; Rose KD &amp; Bloch JI" box="[809,1041,350,372]" pageId="14" pageNumber="445" pagination="73 - 106" refId="ref19662" refString="Penkrot TA, Zack SP, Rose KD, Bloch JI. 2008. Postcranial morphology of Apheliscus and Haplomylus (Condylarthra, Apheliscidae): evidence for a Paleocene Holarctic origin of Macroscelidea. In: Sargis EJ, Dagosto M, eds. Mammalian evolutionary morphology: a tribute to Frederick S. Szalay. The Netherlands: Springer, 73 - 106." type="book chapter" year="2008">
Penkrot
<emphasis id="B96BEA9AFFBCD902FC37A862DC80FE9D" box="[911,970,350,372]" italics="true" pageId="14" pageNumber="445">et al.</emphasis>
, 2008
</bibRefCitation>
). Apheliscines broadly resemble
<taxonomicName id="4C1F4D0BFFBCD902FC91A840DCA2FE7C" authorityName="Scott" authorityYear="2019" box="[809,1000,381,402]" class="Mammalia" family="Zhelestidae" genus="Ferrequitherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Proteutheria" pageId="14" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBCD902FC91A840DCA2FE7C" box="[809,1000,381,402]" italics="true" pageId="14" pageNumber="445">Ferrequitherium</emphasis>
</taxonomicName>
in these regards, and these are best seen in the dentition of
<taxonomicName id="4C1F4D0BFFBCD902FBF8A8A6DAC6FE5F" authority="Jepsen, 1930" authorityName="Jepsen" authorityYear="1930" box="[1088,1420,411,433]" class="Mammalia" family="Mioclaenidae" genus="Phenacodaptes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="14" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBCD902FBF8A8A6DBA7FE5E" box="[1088,1261,411,432]" italics="true" pageId="14" pageNumber="445">Phenacodaptes</emphasis>
<bibRefCitation id="EF8E4B79FFBCD902FB4DA8A1DAC6FE5F" author="Jepsen GL" box="[1269,1420,412,433]" pageId="14" pageNumber="445" pagination="117 - 131" refId="ref18298" refString="Jepsen GL. 1930. New vertebrate fossils from the Lower Eocene of the Bighorn Basin, Wyoming. Proceedings of the American Philosophical Society 69: 117 - 131." type="journal article" year="1930">Jepsen, 1930</bibRefCitation>
</taxonomicName>
, from the Late Palaeocene of
<collectingRegion id="49DBF86AFFBCD902FB2DA887DA43FE3E" box="[1173,1289,442,464]" country="United States of America" name="Wyoming" pageId="14" pageNumber="445">Wyoming</collectingRegion>
, the upper molars and lower fourth premolar of which resemble those of
<taxonomicName id="4C1F4D0BFFBCD902FC37A8CADB18FDE2" authorityName="Scott" authorityYear="2019" box="[911,1106,503,524]" class="Mammalia" family="Zhelestidae" genus="Ferrequitherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Proteutheria" pageId="14" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBCD902FC37A8CADB18FDE2" box="[911,1106,503,524]" italics="true" pageId="14" pageNumber="445">Ferrequitherium</emphasis>
</taxonomicName>
.
<taxonomicName id="4C1F4D0BFFBCD902FBDBA8CADCF6FDC5" authority="Jepsen, 1930" authorityName="Jepsen" authorityYear="1930" class="Mammalia" family="Mioclaenidae" genus="Phenacodaptes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="14" pageNumber="445" phylum="Chordata" rank="species" species="sabulosus">
<emphasis id="B96BEA9AFFBCD902FBDBA8CADADAFDE2" box="[1123,1424,503,524]" italics="true" pageId="14" pageNumber="445">Phenacodaptes sabulosus</emphasis>
<bibRefCitation id="EF8E4B79FFBCD902FC91AB2ADCF6FDC5" author="Jepsen GL" box="[809,956,534,556]" pageId="14" pageNumber="445" pagination="117 - 131" refId="ref18298" refString="Jepsen GL. 1930. New vertebrate fossils from the Lower Eocene of the Bighorn Basin, Wyoming. Proceedings of the American Philosophical Society 69: 117 - 131." type="journal article" year="1930">Jepsen, 1930</bibRefCitation>
</taxonomicName>
, the only known species of the genus, was scored in this analysis. In addition to
<taxonomicName id="4C1F4D0BFFBCD902FB67AB08DAC0FDA4" authorityName="Jepsen" authorityYear="1930" box="[1247,1418,565,586]" class="Mammalia" family="Mioclaenidae" genus="Phenacodaptes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="14" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBCD902FB67AB08DAC0FDA4" box="[1247,1418,565,586]" italics="true" pageId="14" pageNumber="445">Phenacodaptes</emphasis>
</taxonomicName>
, the Late Palaeocene
<taxonomicName id="4C1F4D0BFFBCD902FBB6AB6EDC9AFD66" authority="Zack, Penkrot, Krause &amp; Maas, 2005 b" authorityName="Zack, Penkrot, Krause &amp; Maas" authorityYear="2005" class="Mammalia" family="Apheliscidae" genus="Gingerichia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Macroscelidea" pageId="14" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBCD902FBB6AB6EDBDEFD86" box="[1038,1172,595,616]" italics="true" pageId="14" pageNumber="445">Gingerichia</emphasis>
<bibRefCitation id="EF8E4B79FFBCD902FB20AB6EDC9AFD66" author="Zack SP &amp; Penkrot TA &amp; Krause DW &amp; Maas MC" pageId="14" pageNumber="445" pagination="809 - 830" refId="ref21947" refString="Zack SP, Penkrot TA, Krause DW, Maas MC. 2005 b. A new apheliscine ' condylarth' from the late Paleocene of Montana and Alberta and the phylogeny of ' hyopsodontids'. Acta Palaeontologica Polonica 50: 809 - 830." type="journal article" year="2005">Zack, Penkrot, Krause &amp; Maas, 2005b</bibRefCitation>
</taxonomicName>
was also included;
<taxonomicName id="4C1F4D0BFFBCD902FB10AB4FDA64FD69" authorityName="Zack, Penkrot, Krause &amp; Maas" authorityYear="2005" box="[1192,1326,626,647]" class="Mammalia" family="Apheliscidae" genus="Gingerichia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Macroscelidea" pageId="14" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBCD902FB10AB4FDA64FD69" box="[1192,1326,626,647]" italics="true" pageId="14" pageNumber="445">Gingerichia</emphasis>
</taxonomicName>
is recognized as the basalmost apheliscine (
<bibRefCitation id="EF8E4B79FFBCD902FB07ABACDAC8FD49" author="Zack SP &amp; Penkrot TA &amp; Krause DW &amp; Maas MC" box="[1215,1410,657,679]" pageId="14" pageNumber="445" pagination="809 - 830" refId="ref21947" refString="Zack SP, Penkrot TA, Krause DW, Maas MC. 2005 b. A new apheliscine ' condylarth' from the late Paleocene of Montana and Alberta and the phylogeny of ' hyopsodontids'. Acta Palaeontologica Polonica 50: 809 - 830." type="journal article" year="2005">
Zack
<emphasis id="B96BEA9AFFBCD902FB43ABACDA78FD48" box="[1275,1330,657,678]" italics="true" pageId="14" pageNumber="445">et al.</emphasis>
, 2005b
</bibRefCitation>
). The genus is known from two species (
<taxonomicName id="4C1F4D0BFFBCD902FB44AB8DDBAEFD0A" authority="Zack, Penkrot, Krause &amp; Maas, 2005 b" authorityName="Zack, Penkrot, Krause &amp; Maas" authorityYear="2005" class="Mammalia" family="Apheliscidae" genus="Gingerichia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Macroscelidea" pageId="14" pageNumber="445" phylum="Chordata" rank="species" species="geoteretes">
<emphasis id="B96BEA9AFFBCD902FB44AB8DDA45FD2B" box="[1276,1295,688,709]" italics="true" pageId="14" pageNumber="445">G</emphasis>
.
<emphasis id="B96BEA9AFFBCD902FAA7AB8DDADBFD2B" box="[1311,1425,688,709]" italics="true" pageId="14" pageNumber="445">geoteretes</emphasis>
<bibRefCitation id="EF8E4B79FFBCD902FC91ABF3DBAEFD0A" author="Zack SP &amp; Penkrot TA &amp; Krause DW &amp; Maas MC" box="[809,1252,718,740]" pageId="14" pageNumber="445" pagination="809 - 830" refId="ref21947" refString="Zack SP, Penkrot TA, Krause DW, Maas MC. 2005 b. A new apheliscine ' condylarth' from the late Paleocene of Montana and Alberta and the phylogeny of ' hyopsodontids'. Acta Palaeontologica Polonica 50: 809 - 830." type="journal article" year="2005">Zack, Penkrot, Krause &amp; Maas, 2005b</bibRefCitation>
</taxonomicName>
and
<taxonomicName id="4C1F4D0BFFBCD902FAA7ABF3DBBBFCED" authority="Zack, Penkrot, Krause &amp; Maas, 2005 b" authorityName="Zack, Penkrot, Krause &amp; Maas" authorityYear="2005" class="Mammalia" family="Apheliscidae" genus="Gingerichia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Macroscelidea" pageId="14" pageNumber="445" phylum="Chordata" rank="species" species="hystrix">
<emphasis id="B96BEA9AFFBCD902FAA7ABF3DA78FD0D" box="[1311,1330,718,739]" italics="true" pageId="14" pageNumber="445">G</emphasis>
.
<emphasis id="B96BEA9AFFBCD902FAF8ABF3DADAFD0D" box="[1344,1424,718,739]" italics="true" pageId="14" pageNumber="445">hystrix</emphasis>
<bibRefCitation id="EF8E4B79FFBCD902FC91ABD0DBBBFCED" author="Zack SP &amp; Penkrot TA &amp; Krause DW &amp; Maas MC" box="[809,1265,749,771]" pageId="14" pageNumber="445" pagination="809 - 830" refId="ref21947" refString="Zack SP, Penkrot TA, Krause DW, Maas MC. 2005 b. A new apheliscine ' condylarth' from the late Paleocene of Montana and Alberta and the phylogeny of ' hyopsodontids'. Acta Palaeontologica Polonica 50: 809 - 830." type="journal article" year="2005">Zack, Penkrot, Krause &amp; Maas, 2005b</bibRefCitation>
</taxonomicName>
), and scoring included both of these species. To test for broader apheliscid (i.e. exclusive of Apheliscinae) affinities, the North American Palaeocene genus
<taxonomicName id="4C1F4D0BFFBCD902FB09AA74DC2AFC93" authority="Simpson, 1935" authorityName="Simpson" authorityYear="1935" class="Mammalia" family="Hyopsodontidae" genus="Litomylus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="14" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBCD902FB09AA74DA69FCB0" box="[1201,1315,841,862]" italics="true" pageId="14" pageNumber="445">Litomylus</emphasis>
<bibRefCitation id="EF8E4B79FFBCD902FA91AA74DC2AFC93" author="Simpson GG" pageId="14" pageNumber="445" pagination="221 - 244" refId="ref20746" refString="Simpson GG. 1935. New Paleocene mammals from the Fort Union of Montana. Proceedings of the United States National Museum 83: 221 - 244." type="journal article" year="1935">Simpson, 1935</bibRefCitation>
</taxonomicName>
was also included. Scoring decisions were based on
<taxonomicName id="4C1F4D0BFFBCD902FCF4AABBDA5BFC72" authority="Simpson, 1935" authorityName="Simpson" authorityYear="1935" box="[844,1297,902,924]" class="Mammalia" family="Hyopsodontidae" genus="Litomylus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="14" pageNumber="445" phylum="Chordata" rank="species" species="dissentaneus">
<emphasis id="B96BEA9AFFBCD902FCF4AABBDB17FC75" box="[844,1117,902,923]" italics="true" pageId="14" pageNumber="445">Litomylus dissentaneus</emphasis>
<bibRefCitation id="EF8E4B79FFBCD902FBDDAABBDA5BFC72" author="Simpson GG" box="[1125,1297,902,924]" pageId="14" pageNumber="445" pagination="221 - 244" refId="ref20746" refString="Simpson GG. 1935. New Paleocene mammals from the Fort Union of Montana. Proceedings of the United States National Museum 83: 221 - 244." type="journal article" year="1935">Simpson, 1935</bibRefCitation>
</taxonomicName>
:
<taxonomicName id="4C1F4D0BFFBCD902FAA5AABBDCF5FC54" authorityName="Simpson" authorityYear="1935" class="Mammalia" family="Hyopsodontidae" genus="Litomylus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Condylarthra" pageId="14" pageNumber="445" phylum="Chordata" rank="species" species="dissentaneus">
<emphasis id="B96BEA9AFFBCD902FAA5AABBDCF5FC54" italics="true" pageId="14" pageNumber="445">Litomylus dissentaneus</emphasis>
</taxonomicName>
is reasonably well known and is represented by several well-preserved specimens documenting significant parts of the dentition.
</paragraph>
<paragraph id="8BA03688FFBCD903FC91AD2BDD1CFDC2" blockId="14.[809,1426,1046,1896]" lastBlockId="15.[163,779,197,556]" lastPageId="15" lastPageNumber="446" pageId="14" pageNumber="445">
<emphasis id="B96BEA9AFFBCD902FC91AD2BDC93FBC5" box="[809,985,1046,1067]" italics="true" pageId="14" pageNumber="445">
<taxonomicName id="4C1F4D0BFFBCD902FC91AD2BDC99FBC5" baseAuthorityName="Murray" baseAuthorityYear="1866" box="[809,979,1046,1067]" class="Mammalia" family="Arctocyonidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Insectivora" pageId="14" pageNumber="445" phylum="Chordata" rank="family">Arctocyonidae</taxonomicName>
:
</emphasis>
<bibRefCitation id="EF8E4B79FFBCD902FC5AAD2BDB32FBC2" author="Webb MW" box="[994,1144,1046,1068]" pageId="14" pageNumber="445" refId="ref21331" refString="Webb MW. 1996. Late Paleocene mammals from near Drayton Valley, Alberta. Unpublished M. Sc. Thesis, University of Alberta." type="book" year="1996">Webb (1996)</bibRefCitation>
and Scott
<emphasis id="B96BEA9AFFBCD902FB47AD2ADA76FBC5" box="[1279,1340,1046,1068]" italics="true" pageId="14" pageNumber="445">et al.</emphasis>
(2006) noted similarities in the dentition of
<taxonomicName id="4C1F4D0BFFBCD902FB4FAD08DADBFBA4" authorityName="Scott, Fox &amp; Webb" authorityYear="2006" box="[1271,1425,1077,1098]" class="Mammalia" family="Horolodectidae" genus="Horolodectes" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="14" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBCD902FB4FAD08DADBFBA4" box="[1271,1425,1077,1098]" italics="true" pageId="14" pageNumber="445">Horolodectes</emphasis>
</taxonomicName>
and those of ‘Oxyclaenidae’ (
<emphasis id="B96BEA9AFFBCD902FB36AD69DB9BFB87" box="[1166,1233,1108,1129]" italics="true" pageId="14" pageNumber="445">sensu</emphasis>
<bibRefCitation id="EF8E4B79FFBCD902FB64AD6EDADBFB87" author="Matthew WD" box="[1244,1425,1107,1129]" pageId="14" pageNumber="445" pagination="4 - 103" refId="ref19062" refString="Matthew WD. 1915. A revision of the lower Eocene Wasatch and Wind River faunas, Part I: order Ferae (Carnivora). Suborder Creodonta. Bulletin of the American Museum of Natural History 34: 4 - 103." type="journal article" year="1915">Matthew, 1915</bibRefCitation>
and
<bibRefCitation id="EF8E4B79FFBCD902FCE6AD4FDB3AFB69" author="McKenna MC &amp; Bell SK" box="[862,1136,1138,1160]" pageId="14" pageNumber="445" refId="ref19265" refString="McKenna MC, Bell SK. 1997. Classification of mammals above the species level. New York: Columbia University Press." type="book" year="1997">McKenna &amp; Bell, 1997</bibRefCitation>
), a family of basal ‘condylarths’ with generally plesiomorphic dentitions (
<bibRefCitation id="EF8E4B79FFBCD902FC89AD92DCAFFB2A" author="Archibald JD" box="[817,997,1199,1221]" pageId="14" pageNumber="445" pagination="292 - 331" refId="ref15846" refString="Archibald JD. 1998. Archaic ungulates (' Condylarthra'). In: Janis CM, Scott KM, Jacobs LL, eds. Evolution of tertiary mammals of North America, Vol. 1: terrestrial carnivores, ungulates, and ungulatelike mammals. Cambridge: Cambridge University Press, 292 - 331." type="book chapter" year="1998">Archibald 1998</bibRefCitation>
). There is little consensus on either the monophyly of the various ‘oxyclaenids’ or how best to classify them (e.g.
<bibRefCitation id="EF8E4B79FFBCD902FBDAADD0DBB9FAEC" author="Cifelli RL" box="[1122,1267,1261,1283]" pageId="14" pageNumber="445" pagination="1 - 49" refId="ref16372" refString="Cifelli RL. 1983. The origin and affinities of the South American Condylarthra and early Tertiary Litopterna (Mammalia). American Museum Novitates 2772: 1 - 49." type="journal article" year="1983">Cifelli, 1983</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FABAADD0DCEBFACE" author="McKenna MC &amp; Bell SK" pageId="14" pageNumber="445" refId="ref19265" refString="McKenna MC, Bell SK. 1997. Classification of mammals above the species level. New York: Columbia University Press." type="book" year="1997">McKenna &amp; Bell, 1997</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FC17AC36DB26FACE" author="Archibald JD" box="[943,1132,1291,1313]" pageId="14" pageNumber="445" pagination="292 - 331" refId="ref15846" refString="Archibald JD. 1998. Archaic ungulates (' Condylarthra'). In: Janis CM, Scott KM, Jacobs LL, eds. Evolution of tertiary mammals of North America, Vol. 1: terrestrial carnivores, ungulates, and ungulatelike mammals. Cambridge: Cambridge University Press, 292 - 331." type="book chapter" year="1998">Archibald, 1998</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FBC3AC36DAC1FACE" author="Muizon C de &amp; Cifelli RL" box="[1147,1419,1291,1313]" pageId="14" pageNumber="445" pagination="47 - 150" refId="ref19288" refString="Muizon C de, Cifelli RL. 2000. The ' condylarths' (archaic Ungulata, Mammalia) from the Early Paleocene of Tiupampa (Bolivia): implications on the origin of the South American ungulates. Geodiversitas 22: 47 - 150." type="journal article" year="2000">Muizon &amp; Cifelli, 2000</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FC91AC17DB1DFAD1" author="Williamson TE &amp; Carr TD" box="[809,1111,1322,1344]" pageId="14" pageNumber="445" pagination="973 - 986" refId="ref21556" refString="Williamson TE, Carr TD. 2007. Revision of the problematic Early Paleocene genus Oxyclaenus (Mammalia: Oxyclaenidae) and a new species of Carcinodon. Journal of Vertebrate Paleontology 27: 973 - 986." type="journal article" year="2007">Williamson &amp; Carr, 2007</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FBDFAC17DACBFAD1" author="De Bast E &amp; Smith T" box="[1127,1409,1322,1344]" pageId="14" pageNumber="445" pagination="964 - 976" refId="ref16910" refString="De Bast E, Smith T. 2013. Reassessment of the small ' arctocyonid' Prolatidens waudruae from the Early Paleocene of Belgium, and its phylogenetic relationships with ungulate-like mammals. Journal of Vertebrate Paleontology 33: 964 - 976." type="journal article" year="2013">De Bast &amp; Smith, 2013</bibRefCitation>
). Given these continuing uncertainties, the nomen
<taxonomicName id="4C1F4D0BFFBCD902FC91AC5ADC9EFA93" baseAuthorityName="Murray" baseAuthorityYear="1866" box="[809,980,1383,1405]" class="Mammalia" family="Arctocyonidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Insectivora" pageId="14" pageNumber="445" phylum="Chordata" rank="family">Arctocyonidae</taxonomicName>
is used here to refer to the possibly paraphyletic or even polyphyletic assemblage of basal ‘condylarths’ exclusive of
<taxonomicName id="4C1F4D0BFFBCD902FBE0AC98DB97FA54" authorityName="Cope" authorityYear="1884" box="[1112,1245,1445,1466]" class="Mammalia" family="Oxyclaenidae" genus="Oxyclaenus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Procreodi" pageId="14" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBCD902FBE0AC98DB97FA54" box="[1112,1245,1445,1466]" italics="true" pageId="14" pageNumber="445">Oxyclaenus</emphasis>
</taxonomicName>
(which may be more closely related to mesonychids;
<bibRefCitation id="EF8E4B79FFBCD902FB54ACFEDCEEFA19" author="Williamson TE &amp; Carr TD" pageId="14" pageNumber="445" pagination="973 - 986" refId="ref21556" refString="Williamson TE, Carr TD. 2007. Revision of the problematic Early Paleocene genus Oxyclaenus (Mammalia: Oxyclaenidae) and a new species of Carcinodon. Journal of Vertebrate Paleontology 27: 973 - 986." type="journal article" year="2007">Williamson &amp; Carr, 2007</bibRefCitation>
), while recognizing that some of the constituent arctocyonid taxa may be monophyletic (e.g. Arctocyoninae
<emphasis id="B96BEA9AFFBCD902FC60AF1DDB52F9DB" box="[984,1048,1568,1589]" italics="true" pageId="14" pageNumber="445">sensu</emphasis>
<bibRefCitation id="EF8E4B79FFBCD902FBA7AF22DB93F9DA" author="Archibald JD" box="[1055,1241,1567,1589]" pageId="14" pageNumber="445" pagination="292 - 331" refId="ref15846" refString="Archibald JD. 1998. Archaic ungulates (' Condylarthra'). In: Janis CM, Scott KM, Jacobs LL, eds. Evolution of tertiary mammals of North America, Vol. 1: terrestrial carnivores, ungulates, and ungulatelike mammals. Cambridge: Cambridge University Press, 292 - 331." type="book chapter" year="1998">Archibald, 1998</bibRefCitation>
). Two arctocyonids were included in this analysis:
<taxonomicName id="4C1F4D0BFFBCD902FB63AF03DB0CF99C" authority="Sloan &amp; Van Valen, 1965" authorityName="Sloan &amp; Van Valen" authorityYear="1965" class="Mammalia" family="Tupaiidae" genus="Protungulatum" higherTaxonomySource="GBIF" kingdom="Animalia" order="Insectivora" pageId="14" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBCD902FB63AF03DADBF9BD" box="[1243,1425,1598,1619]" italics="true" pageId="14" pageNumber="445">Protungulatum</emphasis>
<bibRefCitation id="EF8E4B79FFBCD902FC91AF60DB0CF99C" author="Sloan RE &amp; Van Valen L" box="[809,1094,1628,1651]" pageId="14" pageNumber="445" pagination="220 - 227" refId="ref20919" refString="Sloan RE, Van Valen L. 1965. Cretaceous Mammals from Montana. Science (New York, N. Y.) 148: 220 - 227." type="journal article" year="1965">Sloan &amp; Van Valen, 1965</bibRefCitation>
</taxonomicName>
, and
<taxonomicName id="4C1F4D0BFFBCD902FB3CAF60DAC4F99C" authority="Cope, 1885" authorityName="Cope" authorityYear="1885" box="[1156,1422,1628,1650]" class="Mammalia" family="Arctocyonidae" genus="Loxolophus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Procreodi" pageId="14" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBCD902FB3CAF60DA4CF99C" box="[1156,1286,1629,1650]" italics="true" pageId="14" pageNumber="445">Loxolophus</emphasis>
<bibRefCitation id="EF8E4B79FFBCD902FAB6AF60DAC4F99C" author="Cope ED" box="[1294,1422,1628,1650]" pageId="14" pageNumber="445" pagination="385 - 386" refId="ref16852" refString="Cope ED. 1885. The oldest Tertiary Mammalia. American Naturalist 19: 385 - 386." type="journal article" year="1885">Cope, 1885</bibRefCitation>
</taxonomicName>
.
<taxonomicName id="4C1F4D0BFFBCD902FC91AF46DC96F97E" authorityName="Sloan &amp; Van Valen" authorityYear="1965" box="[809,988,1659,1680]" class="Mammalia" family="Tupaiidae" genus="Protungulatum" higherTaxonomySource="GBIF" kingdom="Animalia" order="Insectivora" pageId="14" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBCD902FC91AF46DC96F97E" box="[809,988,1659,1680]" italics="true" pageId="14" pageNumber="445">Protungulatum</emphasis>
</taxonomicName>
is widely regarded as the most basal ‘condylarth’, although its systematic position as a placental (i.e. within the eutherian crown group) or as a stem eutherian continues to be debated (e.g.
<bibRefCitation id="EF8E4B79FFBCD902FAF6AFEADCE4F8E5" author="Wible JR &amp; Rougier GW &amp; Novacek MJ &amp; Asher RJ" pageId="14" pageNumber="445" pagination="1003 - 1006" refId="ref21474" refString="Wible JR, Rougier GW, Novacek MJ, Asher RJ. 2007. Cretaceous eutherians and Laurasian origin for placental mammals near the K / T boundary. Nature 447: 1003 - 1006." type="journal article" year="2007">
Wible
<emphasis id="B96BEA9AFFBCD902FC91AFCADC2FF8E5" box="[809,869,1782,1804]" italics="true" pageId="14" pageNumber="445">et al.</emphasis>
, 2007
</bibRefCitation>
,
<bibRefCitation id="EF8E4B79FFBCD902FC05AFCBDCBCF8E5" author="Wible JR &amp; Rougier GW &amp; Novacek MJ &amp; Asher RJ" box="[957,1014,1782,1803]" pageId="14" pageNumber="445" pagination="1 - 123" refId="ref21510" refString="Wible JR, Rougier GW, Novacek MJ, Asher RJ. 2009. The eutherian mammal Maelestes gobiensis from the Late Cretaceous of Mongolia and the phylogeny of Cretaceous Eutheria. Bulletin of the American Museum of Natural History 327: 1 - 123." type="journal article" year="2009">2009</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FBBEAFCBDA47F8E5" author="Spaulding M &amp; O'Leary MA &amp; Gatesy J" box="[1030,1293,1782,1804]" pageId="14" pageNumber="445" pagination="7062" refId="ref20949" refString="Spaulding M, O'Leary MA, Gatesy J. 2009. Relationships of Cetacea (Artiodactyla) among mammals: increased taxon sampling alters interpretations of key fossils and character evolution. PLoS ONE 4: e 7062." type="journal article" year="2009">
Spaulding
<emphasis id="B96BEA9AFFBCD902FB31AFCADB8FF8E5" box="[1161,1221,1782,1804]" italics="true" pageId="14" pageNumber="445">et al.</emphasis>
, 2009
</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FAA4AFCBDCECF8C4" author="Archibald JD &amp; Zhang Y &amp; Harper T &amp; Cifelli RL" pageId="14" pageNumber="445" pagination="153 - 161" refId="ref16085" refString="Archibald JD, Zhang Y, Harper T, Cifelli RL. 2011. Protungulatum, confirmed Cretaceous occurrence of an otherwise Paleocene eutherian (placental?) mammal. Journal of Mammalian Evolution 18: 153 - 161." type="journal article" year="2011">
Archibald
<emphasis id="B96BEA9AFFBCD902FC91AE28DC28F8C4" box="[809,866,1813,1834]" italics="true" pageId="14" pageNumber="445">et al.</emphasis>
, 2011
</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FC0CAE28DBDEF8C4" author="O'Leary MA &amp; Bloch JI &amp; Flynn JJ &amp; Gaudin TJ &amp; Giallombardo A &amp; Giannini NP &amp; Goldberg SL &amp; Kraatz BP &amp; Luo ZX &amp; Meng J &amp; Ni X &amp; Novacek MJ &amp; Perini FA &amp; Randall ZS &amp; Rougier GW &amp; Sargis EJ &amp; Silcox MT &amp; Simmons NB &amp; Spaulding M &amp; Velazco PM &amp; Weksler M &amp; Wible JR &amp; Cirranello AL" box="[948,1172,1813,1834]" pageId="14" pageNumber="445" pagination="662 - 667" refId="ref19502" refString="O'Leary MA, Bloch JI, Flynn JJ, Gaudin TJ, Giallombardo A, Giannini NP, Goldberg SL, Kraatz BP, Luo ZX, Meng J, Ni X, Novacek MJ, Perini FA, Randall ZS, Rougier GW, Sargis EJ, Silcox MT, Simmons NB, Spaulding M, Velazco PM, Weksler M, Wible JR, Cirranello AL. 2013. The placental mammal ancestor and the post-K-Pg radiation of placentals. Science (New York, N. Y.) 339: 662 - 667." type="journal article" year="2013">
O’Leary
<emphasis id="B96BEA9AFFBCD902FBAFAE28DB1AF8C4" box="[1047,1104,1813,1834]" italics="true" pageId="14" pageNumber="445">et al.</emphasis>
, 2013
</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FB1AAE28DAC0F8C4" author="Halliday TJD &amp; Upchurch P &amp; Goswami A" box="[1186,1418,1812,1835]" pageId="14" pageNumber="445" pagination="20153026" refId="ref18015" refString="Halliday TJD, Upchurch P, Goswami A. 2016. Eutherians experienced elevated evolutionary rates in the immediate aftermath of the Cretaceous - Paleogene mass extinction. Proceedings of the Royal Society B 283: 20153026." type="journal article" year="2016">
Halliday
<emphasis id="B96BEA9AFFBCD902FAB5AE28DA75F8C4" box="[1293,1343,1813,1834]" italics="true" pageId="14" pageNumber="445">et al</emphasis>
., 2016
</bibRefCitation>
,
<bibRefCitation id="EF8E4B79FFBCD902FC91AE0EDC2AF8A6" author="Halliday TJ &amp; Upchurch P &amp; Goswami A" box="[809,864,1843,1864]" pageId="14" pageNumber="445" pagination="521 - 550" refId="ref18053" refString="Halliday TJ, Upchurch P, Goswami A. 2017. Resolving the relationships of Paleocene placental mammals. Biological Reviews of the Cambridge Philosophical Society 92: 521 - 550." type="journal article" year="2017">2017</bibRefCitation>
). The genus is first known from Late Cretaceous deposits of North America (
<bibRefCitation id="EF8E4B79FFBCD902FBDBAE6FDA5AF889" author="Johnston PA" box="[1123,1296,1874,1896]" pageId="14" pageNumber="445" pagination="512 - 519" refId="ref18361" refString="Johnston PA. 1980. First record of Mesozoic mammals from Saskatchewan. Canadian Journal of Earth Sciences 17: 512 - 519." type="journal article" year="1980">Johnston, 1980</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBCD902FAA6AE6FDAC1F889" author="Fox RC" box="[1310,1419,1874,1895]" pageId="14" pageNumber="445" pagination="11 - 59" refId="ref17188" refString="Fox RC. 1989. The Wounded Knee local fauna and mammalian evolution near the K / T boundary, Saskatchewan, Canada. Palaeontographica Abt A 208: 11 - 59." type="journal article" year="1989">Fox, 1989</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBDD903FF1BA9F8DEDCFF34" author="Archibald JD &amp; Zhang Y &amp; Harper T &amp; Cifelli RL" box="[163,406,197,219]" pageId="15" pageNumber="446" pagination="153 - 161" refId="ref16085" refString="Archibald JD, Zhang Y, Harper T, Cifelli RL. 2011. Protungulatum, confirmed Cretaceous occurrence of an otherwise Paleocene eutherian (placental?) mammal. Journal of Mammalian Evolution 18: 153 - 161." type="journal article" year="2011">
Archibald
<emphasis id="B96BEA9AFFBDD903FEA2A9FBDE18FF34" box="[282,338,197,219]" italics="true" pageId="15" pageNumber="446">et al.</emphasis>
, 2011
</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBDD903FE1AA9F8DDCFFF35" author="Redman CM &amp; Gardner JD &amp; Scott CS &amp; Braman DR" box="[418,645,197,219]" pageId="15" pageNumber="446" pagination="846 - 862" refId="ref19768" refString="Redman CM, Gardner JD, Scott CS, Braman DR. 2015. Geological setting of vertebrate microfossil localities across the Cretaceous - Paleogene boundary in southwestern Saskatchewan, Canada. Canadian Journal of Earth Sciences 52: 846 - 862." type="journal article" year="2015">
Redman
<emphasis id="B96BEA9AFFBDD903FDB1A9FBDD0BFF34" box="[521,577,197,219]" italics="true" pageId="15" pageNumber="446">et al.</emphasis>
, 2015
</bibRefCitation>
) but is best known from the Palaeocene of
<collectingRegion id="49DBF86AFFBDD903FDBCA9D9DD26FF17" box="[516,620,228,249]" country="United States of America" name="Montana" pageId="15" pageNumber="446">Montana</collectingRegion>
(
<bibRefCitation id="EF8E4B79FFBDD903FDC3A9D9DE66FEF6" author="Sloan RE &amp; Van Valen L" pageId="15" pageNumber="446" pagination="220 - 227" refId="ref20919" refString="Sloan RE, Van Valen L. 1965. Cretaceous Mammals from Montana. Science (New York, N. Y.) 148: 220 - 227." type="journal article" year="1965">Sloan &amp; Van Valen, 1965</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBDD903FE81A83FDEB9FEF6" author="Archibald JD" box="[313,499,258,280]" pageId="15" pageNumber="446" pagination="1 - 286" refId="ref15810" refString="Archibald JD. 1982. A study of Mammalia and geology across the Cretaceous - Tertiary boundary in Garfield County, Montana. University of California Publications in Geological Sciences 122: 1 - 286." type="journal article" year="1982">Archibald, 1982</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBDD903FDBAA83FDDEEFEF6" author="Lofgren DL" box="[514,676,258,280]" pageId="15" pageNumber="446" pagination="1 - 185" refId="ref18782" refString="Lofgren DL. 1995. The Bug Creek problem and the Cretaceous - Tertiary transition at McGuire Creek, Montana. University of California Publications in Geological Sciences 140: 1 - 185." type="journal article" year="1995">Lofgren, 1995</bibRefCitation>
). Dental characters for
<taxonomicName id="4C1F4D0BFFBDD903FEF6A81CDD4BFED8" authorityName="Sloan &amp; Van Valen" authorityYear="1965" box="[334,513,289,310]" class="Mammalia" family="Tupaiidae" genus="Protungulatum" higherTaxonomySource="GBIF" kingdom="Animalia" order="Insectivora" pageId="15" pageNumber="446" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBDD903FEF6A81CDD4BFED8" box="[334,513,289,310]" italics="true" pageId="15" pageNumber="446">Protungulatum</emphasis>
</taxonomicName>
were scored using the
<typeStatus id="54A4882AFFBDD903FF1BA87DDF9FFEBB" box="[163,213,320,341]" pageId="15" pageNumber="446">type</typeStatus>
species,
<taxonomicName id="4C1F4D0BFFBDD903FE83A87DDD9BFEB8" authority="Sloan &amp; Van Valen, 1965" authorityName="Sloan &amp; Van Valen" authorityYear="1965" box="[315,721,320,342]" class="Mammalia" family="Tupaiidae" genus="Protungulatum" higherTaxonomySource="GBIF" kingdom="Animalia" order="Insectivora" pageId="15" pageNumber="446" phylum="Chordata" rank="species" species="donnae">
<emphasis id="B96BEA9AFFBDD903FE83A87DDE01FEBB" box="[315,331,320,341]" italics="true" pageId="15" pageNumber="446">P</emphasis>
.
<emphasis id="B96BEA9AFFBDD903FEE1A87DDEE7FEBB" box="[345,429,320,341]" italics="true" pageId="15" pageNumber="446">donnae</emphasis>
<bibRefCitation id="EF8E4B79FFBDD903FE0CA87DDD9BFEB8" author="Sloan RE &amp; Van Valen L" box="[436,721,320,342]" pageId="15" pageNumber="446" pagination="220 - 227" refId="ref20919" refString="Sloan RE, Van Valen L. 1965. Cretaceous Mammals from Montana. Science (New York, N. Y.) 148: 220 - 227." type="journal article" year="1965">Sloan &amp; Van Valen, 1965</bibRefCitation>
</taxonomicName>
, and an as-yet unnamed new species from the Puercan of
<collectingRegion id="49DBF86AFFBDD903FF1BA840DFB4FE7D" box="[163,254,381,403]" country="Canada" name="Alberta" pageId="15" pageNumber="446">Alberta</collectingRegion>
.
<taxonomicName id="4C1F4D0BFFBDD903FEB1A840DEC4FE7C" authorityName="Cope" authorityYear="1885" box="[265,398,381,402]" class="Mammalia" family="Arctocyonidae" genus="Loxolophus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Procreodi" pageId="15" pageNumber="446" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBDD903FEB1A840DEC4FE7C" box="[265,398,381,402]" italics="true" pageId="15" pageNumber="446">Loxolophus</emphasis>
</taxonomicName>
, a potentially non-monophyletic ‘condylarth’ (see
<bibRefCitation id="EF8E4B79FFBDD903FECFA8A1DD2EFE5F" author="Hunter JP &amp; Hartman JH &amp; Krause DW" box="[375,612,412,433]" pageId="15" pageNumber="446" pagination="61 - 114" refId="ref18233" refString="Hunter JP, Hartman JH, Krause DW. 1997. Mammals and mollusks across the CretaceousTertiary boundary from Makoshika State Park and vicinity (Williston basin), Montana. University of Wyoming Contributions to Geology 32: 61 - 114." type="journal article" year="1997">
Hunter
<emphasis id="B96BEA9AFFBDD903FE62A8A1DD52FE5F" box="[474,536,412,433]" italics="true" pageId="15" pageNumber="446">et al.</emphasis>
, 1997
</bibRefCitation>
) known from Puercan and Torrejonian deposits in several areas of the North American Western Interior, was included as a derived arctocyonid; dental characters were scored for
<taxonomicName id="4C1F4D0BFFBDD903FF71AB2BDD1BFDC2" authority="Cope, 1885" authorityName="Cope" authorityYear="1885" box="[201,593,534,556]" class="Mammalia" family="Arctocyonidae" genus="Loxolophus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Procreodi" pageId="15" pageNumber="446" phylum="Chordata" rank="species" species="hyattianus">
<emphasis id="B96BEA9AFFBDD903FF71AB2BDE81FDC5" box="[201,459,534,555]" italics="true" pageId="15" pageNumber="446">Loxolophus hyattianus</emphasis>
<bibRefCitation id="EF8E4B79FFBDD903FE6AAB2ADD1BFDC2" author="Cope ED" box="[466,593,534,556]" pageId="15" pageNumber="446" pagination="385 - 386" refId="ref16852" refString="Cope ED. 1885. The oldest Tertiary Mammalia. American Naturalist 19: 385 - 386." type="journal article" year="1885">Cope, 1885</bibRefCitation>
</taxonomicName>
.
</paragraph>
<paragraph id="8BA03688FFBDD903FF1BAB6DDE06FAC9" blockId="15.[163,780,592,1319]" pageId="15" pageNumber="446">
<emphasis id="B96BEA9AFFBDD903FF1BAB6DDE7DFD8B" box="[163,311,592,613]" italics="true" pageId="15" pageNumber="446">
<taxonomicName id="4C1F4D0BFFBDD903FF1BAB6DDE7AFD8B" box="[163,304,592,613]" class="Mammalia" family="Zhelestidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Proteutheria" pageId="15" pageNumber="446" phylum="Chordata" rank="family">Zhelestidae</taxonomicName>
:
</emphasis>
The
<taxonomicName id="4C1F4D0BFFBDD903FEC0AB6DDD4CFD88" box="[376,518,592,614]" class="Mammalia" family="Zhelestidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Proteutheria" pageId="15" pageNumber="446" phylum="Chordata" rank="family">Zhelestidae</taxonomicName>
are a family of Late Cretaceous, primarily Central Asian eutherians that have previously been linked to crown group ungulates (e.g.
<bibRefCitation id="EF8E4B79FFBDD903FF62AB91DEDEFD2F" author="Archibald JD" box="[218,404,684,706]" pageId="15" pageNumber="446" pagination="292 - 331" refId="ref15846" refString="Archibald JD. 1998. Archaic ungulates (' Condylarthra'). In: Janis CM, Scott KM, Jacobs LL, eds. Evolution of tertiary mammals of North America, Vol. 1: terrestrial carnivores, ungulates, and ungulatelike mammals. Cambridge: Cambridge University Press, 292 - 331." type="book chapter" year="1998">Archibald, 1998</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBDD903FE1BAB91DDCBFD2F" author="Nessov LA &amp; Archibald JD &amp; Kielan-Jaworowska Z" box="[419,641,684,705]" pageId="15" pageNumber="446" pagination="40 - 88" refId="ref19374" refString="Nessov LA, Archibald JD, Kielan-Jaworowska Z. 1998. Ungulate-like mammals from the Late Cretaceous of Uzbekistan and a phylogenetic analysis of Ungulatomorpha. Bulletin of Carnegie Museum of Natural History 34: 40 - 88." type="journal article" year="1998">
Nessov
<emphasis id="B96BEA9AFFBDD903FE47AB91DD70FD2F" box="[511,570,684,705]" italics="true" pageId="15" pageNumber="446">et al.</emphasis>
, 1998
</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBDD903FD2FAB91DE69FD0E" author="Archibald JD &amp; Averianov AO &amp; Ekdale EG" pageId="15" pageNumber="446" pagination="62 - 65" refId="ref16053" refString="Archibald JD, Averianov AO, Ekdale EG. 2001. Late Cretaceous relatives of rabbits, rodents, and other extant eutherian mammals. Nature 414: 62 - 65." type="journal article" year="2001">
Archibald
<emphasis id="B96BEA9AFFBDD903FF1BABF6DF97FD31" box="[163,221,714,736]" italics="true" pageId="15" pageNumber="446">et al.</emphasis>
, 2001
</bibRefCitation>
), but more recently have been considered stem eutherians (e.g.
<bibRefCitation id="EF8E4B79FFBDD903FE25ABD4DD3FFD10" author="Ekdale EG &amp; Archibald JD &amp; Averianov AO" box="[413,629,745,767]" pageId="15" pageNumber="446" pagination="161 - 176" refId="ref17084" refString="Ekdale EG, Archibald JD, Averianov AO. 2004. Petrosal bones of placental mammals from the Late Cretaceous of Uzbekistan. Acta Palaeontologica Polonica 49: 161 - 176." type="journal article" year="2004">
Ekdale
<emphasis id="B96BEA9AFFBDD903FE4EABD7DD7AFD10" box="[502,560,745,767]" italics="true" pageId="15" pageNumber="446">et al.</emphasis>
, 2004
</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBDD903FD3AABD4DF97FCF3" author="Wible JR &amp; Rougier GW &amp; Novacek MJ &amp; Asher RJ" pageId="15" pageNumber="446" pagination="1003 - 1006" refId="ref21474" refString="Wible JR, Rougier GW, Novacek MJ, Asher RJ. 2007. Cretaceous eutherians and Laurasian origin for placental mammals near the K / T boundary. Nature 447: 1003 - 1006." type="journal article" year="2007">
Wible
<emphasis id="B96BEA9AFFBDD903FD73ABD7DC4FFD10" box="[715,773,745,767]" italics="true" pageId="15" pageNumber="446">et al.</emphasis>
, 2007
</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBDD903FF55AA35DE95FCF3" author="Chester SGB &amp; Sargis EJ &amp; Szalay FS &amp; Archibald JD &amp; Averianov AO" box="[237,479,776,798]" pageId="15" pageNumber="446" pagination="199 - 211" refId="ref16336" refString="Chester SGB, Sargis EJ, Szalay FS, Archibald JD, Averianov AO. 2010. Mammalian distal humeri from the Late Cretaceous of Uzbekistan. Acta Palaeontologica Polonica 55: 199 - 211." type="journal article" year="2010">
Chester
<emphasis id="B96BEA9AFFBDD903FEEEAA35DEDEFCF3" box="[342,404,776,797]" italics="true" pageId="15" pageNumber="446">et al.</emphasis>
, 2010
</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBDD903FE56AA35DF96FCD2" author="Archibald JD &amp; Averianov AO" pageId="15" pageNumber="446" pagination="361 - 426" refId="ref15931" refString="Archibald JD, Averianov AO. 2012. Phylogenetic analysis, taxonomic revision and dental ontogeny of the Cretaceous Zhelestidae (Mammalia: Eutheria). Zoological Journal of the Linnean Society 164: 361 - 426." type="journal article" year="2012">Archibald &amp; Averianov, 2012</bibRefCitation>
). Scott
<emphasis id="B96BEA9AFFBDD903FE83AA1ADE32FCD5" box="[315,376,806,828]" italics="true" pageId="15" pageNumber="446">et al.</emphasis>
(2006) noted several similarities between zhelestids and
<taxonomicName id="4C1F4D0BFFBDD903FE01AA78DD06FCB4" authorityName="Scott, Fox &amp; Webb" authorityYear="2006" box="[441,588,837,858]" class="Mammalia" family="Horolodectidae" genus="Horolodectes" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="446" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBDD903FE01AA78DD06FCB4" box="[441,588,837,858]" italics="true" pageId="15" pageNumber="446">Horolodectes</emphasis>
</taxonomicName>
, including teeth with low crowns and swollen cusps, molars with narrow stylar shelves, large conules, and lower molars with wide talonids and a closely approximated entoconid and hypoconulid. The dentition of
<taxonomicName id="4C1F4D0BFFBDD903FDE8AAFDDC41FC3B" authorityName="Scott" authorityYear="2019" box="[592,779,960,981]" class="Mammalia" family="Zhelestidae" genus="Ferrequitherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Proteutheria" pageId="15" pageNumber="446" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBDD903FDE8AAFDDC41FC3B" box="[592,779,960,981]" italics="true" pageId="15" pageNumber="446">Ferrequitherium</emphasis>
</taxonomicName>
resembles that of zhelestids in these features as well, but with additional similarities in the premolars, particularly the development of a metaconid on p4 (= p5 of zhelestids). Zhelestid characters were primarily scored for
<taxonomicName id="4C1F4D0BFFBDD903FEA3AD64DD33FB81" authority="Nessov, 1985" authorityName="Nessov" authorityYear="1985" box="[283,633,1113,1135]" class="Mammalia" family="Aegialodontidae" genus="Aspanlestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Aegialodontia" pageId="15" pageNumber="446" phylum="Chordata" rank="species" species="atap">
<emphasis id="B96BEA9AFFBDD903FEA3AD64DE96FB81" box="[283,476,1113,1135]" italics="true" pageId="15" pageNumber="446">Aspanlestes atap</emphasis>
<bibRefCitation id="EF8E4B79FFBDD903FE5CAD64DD33FB81" author="Nessov LA" box="[484,633,1113,1135]" pageId="15" pageNumber="446" pagination="199 - 219" refId="ref19333" refString="Nessov LA. 1985. Rare bony fishes, terrestrial lizards and mammals from the zone of estuaries and coastal plains of the Cretaceous of Kizylkum. Ezhegodnik Vsesoyuznogo Palaeontologicheskogo Obshchestva 28: 199 - 219 [in Russian]." type="journal article" year="1985">Nessov, 1985</bibRefCitation>
</taxonomicName>
;
<taxonomicName id="4C1F4D0BFFBDD903FD3DAD64DC41FB80" authorityName="Nessov" authorityYear="1985" box="[645,779,1113,1134]" class="Mammalia" family="Aegialodontidae" genus="Aspanlestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Aegialodontia" pageId="15" pageNumber="446" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBDD903FD3DAD64DC41FB80" box="[645,779,1113,1134]" italics="true" pageId="15" pageNumber="446">Aspanlestes</emphasis>
</taxonomicName>
is among the best-known zhelestids, with significant parts of the dentition, dermatocranium and basicranium now known (
<bibRefCitation id="EF8E4B79FFBDD903FE1EAD88DDB1FB24" author="Archibald JD &amp; Averianov AO" box="[422,763,1205,1227]" pageId="15" pageNumber="446" pagination="361 - 426" refId="ref15931" refString="Archibald JD, Averianov AO. 2012. Phylogenetic analysis, taxonomic revision and dental ontogeny of the Cretaceous Zhelestidae (Mammalia: Eutheria). Zoological Journal of the Linnean Society 164: 361 - 426." type="journal article" year="2012">Archibald &amp; Averianov, 2012</bibRefCitation>
). Because M3 is undiscovered for
<taxonomicName id="4C1F4D0BFFBDD903FD88ADE9DDF7FB07" authorityName="Nessov" authorityYear="1985" box="[560,701,1236,1257]" class="Mammalia" family="Aegialodontidae" genus="Aspanlestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Aegialodontia" pageId="15" pageNumber="446" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBDD903FD88ADE9DDF7FB07" box="[560,701,1236,1257]" italics="true" pageId="15" pageNumber="446">Aspanlestes</emphasis>
</taxonomicName>
, character 29 was scored for
<taxonomicName id="4C1F4D0BFFBDD903FE00ADCFDE02FAC9" authority="(Nessov, 1985)" baseAuthorityName="Nessov" baseAuthorityYear="1985" class="Mammalia" family="Gypsonictopidae" genus="Parazhelestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pilosa" pageId="15" pageNumber="446" phylum="Chordata" rank="species" species="mynbulakensis">
<emphasis id="B96BEA9AFFBDD903FE00ADCFDC40FAE9" box="[440,778,1266,1287]" italics="true" pageId="15" pageNumber="446">Parazhelestes mynbulakensis</emphasis>
(
<bibRefCitation id="EF8E4B79FFBDD903FF12AC2CDE75FAC9" author="Nessov LA" box="[170,319,1297,1319]" pageId="15" pageNumber="446" pagination="199 - 219" refId="ref19333" refString="Nessov LA. 1985. Rare bony fishes, terrestrial lizards and mammals from the zone of estuaries and coastal plains of the Cretaceous of Kizylkum. Ezhegodnik Vsesoyuznogo Palaeontologicheskogo Obshchestva 28: 199 - 219 [in Russian]." type="journal article" year="1985">Nessov, 1985</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
<paragraph id="8BA03688FFBDD903FF1BAC78DB2AFD0A" blockId="15.[163,780,1349,1892]" lastBlockId="15.[827,1444,197,740]" pageId="15" pageNumber="446">
<taxonomicName id="4C1F4D0BFFBDD903FF1BAC78DE9CFAB5" authority="Haeckel, 1866" authorityName="Haeckel" authorityYear="1866" box="[163,470,1349,1371]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lipotyphla" pageId="15" pageNumber="446" phylum="Chordata" rank="order">
<emphasis id="B96BEA9AFFBDD903FF1BAC78DE69FAB4" box="[163,291,1349,1370]" italics="true" pageId="15" pageNumber="446">Lipotyphla</emphasis>
<bibRefCitation id="EF8E4B79FFBDD903FE94AC78DE9CFAB5" author="Haeckel E" box="[300,470,1349,1371]" pageId="15" pageNumber="446" refId="ref17976" refString="Haeckel E. 1866. Generelle Morphologie der Organismen: allgemeine Grundzuge der organischen Formen-Wissenschaft, mechanisch begrundet durch die von Charles Darwin reformierte Descendenz-Theorie, v. 1 - 2: Berlin: Verlag von Georg Reimer." type="book" year="1866">Haeckel, 1866</bibRefCitation>
</taxonomicName>
: Whereas the dentition of
<taxonomicName id="4C1F4D0BFFBDD903FF1BAC59DE72FA97" authorityName="Scott, Fox &amp; Webb" authorityYear="2006" box="[163,312,1380,1401]" class="Mammalia" family="Horolodectidae" genus="Horolodectes" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="15" pageNumber="446" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBDD903FF1BAC59DE72FA97" box="[163,312,1380,1401]" italics="true" pageId="15" pageNumber="446">Horolodectes</emphasis>
</taxonomicName>
provides little indication of a potential relationship with lipotyphlans, the less specialized dentition of
<taxonomicName id="4C1F4D0BFFBDD903FE96AC9CDEA0FA58" authorityName="Scott" authorityYear="2019" box="[302,490,1441,1462]" class="Mammalia" family="Zhelestidae" genus="Ferrequitherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Proteutheria" pageId="15" pageNumber="446" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBDD903FE96AC9CDEA0FA58" box="[302,490,1441,1462]" italics="true" pageId="15" pageNumber="446">Ferrequitherium</emphasis>
</taxonomicName>
shows broad similarities with those of various putative lipotyphlans from the Palaeocene, particularly those that have been classified in the
<taxonomicName id="4C1F4D0BFFBDD903FE83ACC0DEB5F9FD" box="[315,511,1533,1555]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Erinaceomorpha" pageId="15" pageNumber="446" phylum="Chordata" rank="order">Erinaceomorpha</taxonomicName>
(hedgehog-like lipotyphlans;
<emphasis id="B96BEA9AFFBDD903FEB8AF21DE0AF9DF" box="[256,320,1564,1585]" italics="true" pageId="15" pageNumber="446">sensu</emphasis>
<bibRefCitation id="EF8E4B79FFBDD903FEF0AF21DD7EF9DF" author="Novacek MJ &amp; Bown TM &amp; Schankler DM" box="[328,564,1563,1586]" pageId="15" pageNumber="446" pagination="1 - 22" refId="ref19469" refString="Novacek MJ, Bown TM, Schankler DM. 1985. On the classification of Early Tertiary Erinaceomorpha (Insectivora, Mammalia). American Museum Novitates 2813: 1 - 22." type="journal article" year="1985">
Novacek
<emphasis id="B96BEA9AFFBDD903FE0BAF21DEA7F9DF" box="[435,493,1564,1585]" italics="true" pageId="15" pageNumber="446">et al.</emphasis>
, 1985
</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBDD903FDFAAF21DF96F9A1" author="McKenna MC &amp; Bell SK" pageId="15" pageNumber="446" refId="ref19265" refString="McKenna MC, Bell SK. 1997. Classification of mammals above the species level. New York: Columbia University Press." type="book" year="1997">McKenna &amp; Bell, 1997</bibRefCitation>
; but see the contrasting opinion of:
<bibRefCitation id="EF8E4B79FFBDD903FD28AF07DE6BF981" author="Penkrot TA &amp; Zack SP" pageId="15" pageNumber="446" pagination="1212059" refId="ref19622" refString="Penkrot TA, Zack SP. 2016. Tarsals of Sespedectinae (? Lipotyphla) from the Middle Eocene of southern California, and the affinities of Eocene ' erinaceomorphs'. Journal of Vertebrate Palaeontology 36: e 1212059." type="journal article" year="2016">Penkrot &amp; Zack, 2016</bibRefCitation>
). The teeth of
<taxonomicName id="4C1F4D0BFFBDD903FE76AF64DC41F980" authority="Gingerich, 1983" authorityName="Gingerich" authorityYear="1983" box="[462,779,1625,1647]" class="Mammalia" genus="Litocherus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Macroscelidea" pageId="15" pageNumber="446" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBDD903FE76AF64DD03F980" box="[462,585,1625,1646]" italics="true" pageId="15" pageNumber="446">Litocherus</emphasis>
Gingerich, 1983
</taxonomicName>
and
<taxonomicName id="4C1F4D0BFFBDD903FF61AF45DEBBF963" authority="Jepsen, 1930" authorityName="Jepsen" authorityYear="1930" box="[217,497,1656,1677]" class="Mammalia" family="Erinaceidae" genus="Litolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Erinaceomorpha" pageId="15" pageNumber="446" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBDD903FF61AF45DE00F963" box="[217,330,1656,1677]" italics="true" pageId="15" pageNumber="446">Litolestes</emphasis>
<bibRefCitation id="EF8E4B79FFBDD903FEECAF45DEBBF963" author="Jepsen GL" box="[340,497,1656,1677]" pageId="15" pageNumber="446" pagination="117 - 131" refId="ref18298" refString="Jepsen GL. 1930. New vertebrate fossils from the Lower Eocene of the Bighorn Basin, Wyoming. Proceedings of the American Philosophical Society 69: 117 - 131." type="journal article" year="1930">Jepsen, 1930</bibRefCitation>
</taxonomicName>
, particularly the lower fourth premolar, resemble those of
<taxonomicName id="4C1F4D0BFFBDD903FDFBAFABDC4EF945" authorityName="Scott" authorityYear="2019" box="[579,772,1686,1707]" class="Mammalia" family="Zhelestidae" genus="Ferrequitherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Proteutheria" pageId="15" pageNumber="446" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBDD903FDFBAFABDC4EF945" box="[579,772,1686,1707]" italics="true" pageId="15" pageNumber="446">Ferrequitherium</emphasis>
</taxonomicName>
, and the two former genera were included on that account, with
<taxonomicName id="4C1F4D0BFFBDD903FEFDAFE9DEF4F907" authorityName="Gingerich" authorityYear="1983" box="[325,446,1748,1769]" class="Mammalia" genus="Litocherus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Macroscelidea" pageId="15" pageNumber="446" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBDD903FEFDAFE9DEF4F907" box="[325,446,1748,1769]" italics="true" pageId="15" pageNumber="446">Litocherus</emphasis>
</taxonomicName>
being potentially among the basalmost erinaceomorphs, and
<taxonomicName id="4C1F4D0BFFBDD903FD9DAFCFDDDEF8E9" authorityName="Jepsen" authorityYear="1930" box="[549,660,1778,1799]" class="Mammalia" family="Erinaceidae" genus="Litolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Erinaceomorpha" pageId="15" pageNumber="446" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBDD903FD9DAFCFDDDEF8E9" box="[549,660,1778,1799]" italics="true" pageId="15" pageNumber="446">Litolestes</emphasis>
</taxonomicName>
being potentially the earliest erinaceid (see
<bibRefCitation id="EF8E4B79FFBDD903FDE0AE2CDF90F8AB" author="Novacek MJ &amp; Bown TM &amp; Schankler DM" pageId="15" pageNumber="446" pagination="1 - 22" refId="ref19469" refString="Novacek MJ, Bown TM, Schankler DM. 1985. On the classification of Early Tertiary Erinaceomorpha (Insectivora, Mammalia). American Museum Novitates 2813: 1 - 22." type="journal article" year="1985">
Novacek
<emphasis id="B96BEA9AFFBDD903FD7FAE2FDC4EF8C8" box="[711,772,1809,1831]" italics="true" pageId="15" pageNumber="446">et al.</emphasis>
, 1985
</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBDD903FF5EAE0DDE8EF8AB" author="O'Leary MA &amp; Bloch JI &amp; Flynn JJ &amp; Gaudin TJ &amp; Giallombardo A &amp; Giannini NP &amp; Goldberg SL &amp; Kraatz BP &amp; Luo ZX &amp; Meng J &amp; Ni X &amp; Novacek MJ &amp; Perini FA &amp; Randall ZS &amp; Rougier GW &amp; Sargis EJ &amp; Silcox MT &amp; Simmons NB &amp; Spaulding M &amp; Velazco PM &amp; Weksler M &amp; Wible JR &amp; Cirranello AL" box="[230,452,1839,1861]" pageId="15" pageNumber="446" pagination="662 - 667" refId="ref19502" refString="O'Leary MA, Bloch JI, Flynn JJ, Gaudin TJ, Giallombardo A, Giannini NP, Goldberg SL, Kraatz BP, Luo ZX, Meng J, Ni X, Novacek MJ, Perini FA, Randall ZS, Rougier GW, Sargis EJ, Silcox MT, Simmons NB, Spaulding M, Velazco PM, Weksler M, Wible JR, Cirranello AL. 2013. The placental mammal ancestor and the post-K-Pg radiation of placentals. Science (New York, N. Y.) 339: 662 - 667." type="journal article" year="2013">
O’Leary
<emphasis id="B96BEA9AFFBDD903FEF0AE0DDECAF8AA" box="[328,384,1839,1861]" italics="true" pageId="15" pageNumber="446">et al.</emphasis>
, 2013
</bibRefCitation>
). Characters were scored for
<taxonomicName id="4C1F4D0BFFBDD903FF1BAE73DD29F88A" authority="(Simpson, 1936)" baseAuthorityName="Simpson" baseAuthorityYear="1936" box="[163,611,1870,1892]" class="Mammalia" genus="Litocherus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Macroscelidea" pageId="15" pageNumber="446" phylum="Chordata" rank="species" species="notissimus">
<emphasis id="B96BEA9AFFBDD903FF1BAE73DEEBF88A" box="[163,417,1870,1892]" italics="true" pageId="15" pageNumber="446">Litocherus notissimus</emphasis>
(
<bibRefCitation id="EF8E4B79FFBDD903FE08AE73DD11F88A" author="Simpson GG" box="[432,603,1870,1892]" pageId="15" pageNumber="446" pagination="1 - 27" refId="ref20774" refString="Simpson GG. 1936. A new fauna from the Fort Union of Montana. American Museum Novitates 873: 1 - 27." type="journal article" year="1936">Simpson, 1936</bibRefCitation>
)
</taxonomicName>
and
<taxonomicName id="4C1F4D0BFFBDD903FD26AE73DB7CFF34" authority="Jepsen, 1930" authorityName="Jepsen" authorityYear="1930" box="[670,1078,197,1891]" class="Mammalia" family="Erinaceidae" genus="Litolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Erinaceomorpha" pageId="15" pageNumber="446" phylum="Chordata" rank="species" species="ignotus">
<emphasis id="B96BEA9AFFBDD903FD26AE73DCD8FF34" box="[670,914,197,1891]" italics="true" pageId="15" pageNumber="446">Litolestes ignotus</emphasis>
<bibRefCitation id="EF8E4B79FFBDD903FC23A9F8DB7CFF34" author="Jepsen GL" box="[923,1078,197,218]" pageId="15" pageNumber="446" pagination="117 - 131" refId="ref18298" refString="Jepsen GL. 1930. New vertebrate fossils from the Lower Eocene of the Bighorn Basin, Wyoming. Proceedings of the American Philosophical Society 69: 117 - 131." type="journal article" year="1930">Jepsen, 1930</bibRefCitation>
</taxonomicName>
. In addition to
<taxonomicName id="4C1F4D0BFFBDD903FB49A9F8DA27FF34" authorityName="Gingerich" authorityYear="1983" box="[1265,1389,197,218]" class="Mammalia" genus="Litocherus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Macroscelidea" pageId="15" pageNumber="446" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBDD903FB49A9F8DA27FF34" box="[1265,1389,197,218]" italics="true" pageId="15" pageNumber="446">Litocherus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C1F4D0BFFBDD903FC83A9DEDCECFF16" authorityName="Jepsen" authorityYear="1930" box="[827,934,227,248]" class="Mammalia" family="Erinaceidae" genus="Litolestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Erinaceomorpha" pageId="15" pageNumber="446" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBDD903FC83A9DEDCECFF16" box="[827,934,227,248]" italics="true" pageId="15" pageNumber="446">Litolestes</emphasis>
</taxonomicName>
, the genus
<taxonomicName id="4C1F4D0BFFBDD903FB91A9DEDA3AFF17" authority="Gingerich, 1983" authorityName="Gingerich" authorityYear="1983" box="[1065,1392,227,249]" class="Mammalia" genus="Diacocherus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Macroscelidea" pageId="15" pageNumber="446" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBDD903FB91A9DEDBFEFF16" box="[1065,1204,227,248]" italics="true" pageId="15" pageNumber="446">Diacocherus</emphasis>
Gingerich, 1983
</taxonomicName>
was also included in the analysis:
<taxonomicName id="4C1F4D0BFFBDD903FBC7A83FDA42FEF9" authorityName="Gingerich" authorityYear="1983" box="[1151,1288,258,279]" class="Mammalia" genus="Diacocherus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Macroscelidea" pageId="15" pageNumber="446" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBDD903FBC7A83FDA42FEF9" box="[1151,1288,258,279]" italics="true" pageId="15" pageNumber="446">Diacocherus</emphasis>
</taxonomicName>
has been considered a basal erinaceomorph (e.g.
<bibRefCitation id="EF8E4B79FFBDD903FB69A81CDAD7FED9" author="Krishtalka L" box="[1233,1437,289,311]" pageId="15" pageNumber="446" pagination="1 - 40" refId="ref18579" refString="Krishtalka L. 1976 a. Early Tertiary Adapisoricidae and Erinaceidae (Mammalia: Insectivora) of North America. Bulletin of Carnegie Museum of Natural History 1: 1 - 40." type="journal article" year="1976">Krishtalka, 1976a</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBDD903FC83A802DCA1FEBB" author="Novacek MJ" box="[827,1003,319,341]" pageId="15" pageNumber="446" pagination="135 - 149" refId="ref19413" refString="Novacek MJ. 1982. Diacodon alticuspis, an erinaceomorph insectivore from the Early Eocene of northern New Mexico. University of Wyoming Contributions to Geology 20: 135 - 149." type="journal article" year="1982">Novacek, 1982</bibRefCitation>
, 1985;
<bibRefCitation id="EF8E4B79FFBDD903FBFFA802DA75FEBB" author="Novacek MJ &amp; Bown TM &amp; Schankler DM" box="[1095,1343,319,341]" pageId="15" pageNumber="446" pagination="1 - 22" refId="ref19469" refString="Novacek MJ, Bown TM, Schankler DM. 1985. On the classification of Early Tertiary Erinaceomorpha (Insectivora, Mammalia). American Museum Novitates 2813: 1 - 22." type="journal article" year="1985">
Novacek
<emphasis id="B96BEA9AFFBDD903FB00A87DDBBFFEBA" box="[1208,1269,319,341]" italics="true" pageId="15" pageNumber="446">et al.</emphasis>
, 1985
</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBDD903FAE9A802DC3BFE9D" author="Butler PM" pageId="15" pageNumber="446" pagination="117 - 141" refId="ref16267" refString="Butler PM. 1988. Phylogeny of the insectivores. In: Benton M, ed. The phylogeny and classification of the tetrapods. Oxford: Clarendon Press, 117 - 141." type="book chapter" year="1988">Butler, 1988</bibRefCitation>
), although its status, both as a genus distinct from
<taxonomicName id="4C1F4D0BFFBDD903FC83A840DB01FE7D" authority="Russell, 1964" authorityName="Russell" authorityYear="1964" box="[827,1099,381,403]" class="Mammalia" genus="Adunator" higherTaxonomySource="GBIF" kingdom="Animalia" order="Macroscelidea" pageId="15" pageNumber="446" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBDD903FC83A840DCE3FE7C" box="[827,937,381,402]" italics="true" pageId="15" pageNumber="446">Adunator</emphasis>
<bibRefCitation id="EF8E4B79FFBDD903FC08A840DB01FE7D" author="Russell DE" box="[944,1099,381,403]" pageId="15" pageNumber="446" pagination="1 - 321" refId="ref20012" refString="Russell DE. 1964. Les Mammiferes paleocenes d'Europe. Memoirs du Museum National d'Histoire Naturelle (France), Nouvelle Serie, Serie C 13: 1 - 321." type="journal article" year="1964">Russell, 1964</bibRefCitation>
</taxonomicName>
, and as a lipotyphlan, continues to be debated (
<bibRefCitation id="EF8E4B79FFBDD903FBA8A8A6DB96FE5F" author="Krishtalka L" box="[1040,1244,411,433]" pageId="15" pageNumber="446" pagination="1 - 40" refId="ref18579" refString="Krishtalka L. 1976 a. Early Tertiary Adapisoricidae and Erinaceidae (Mammalia: Insectivora) of North America. Bulletin of Carnegie Museum of Natural History 1: 1 - 40." type="journal article" year="1976">Krishtalka, 1976a</bibRefCitation>
; Gingerich, 1983;
<bibRefCitation id="EF8E4B79FFBDD903FC83A887DC86FE21" author="Secord R" box="[827,972,442,464]" pageId="15" pageNumber="446" pagination="1 - 92" refId="ref20613" refString="Secord R. 2008. The Tiffanian land-mammal age (Middle and Late Paleocene) in the northern Bighorn Basin, Wyoming. University of Michigan Papers on Paleontology 35: 1 - 92." type="journal article" year="2008">Secord, 2008</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBDD903FC62A887DBA1FE21" author="Hooker JJ &amp; Russell DE" box="[986,1259,442,464]" pageId="15" pageNumber="446" pagination="856 - 936" refId="ref18197" refString="Hooker JJ, Russell DE. 2012. Early Paleogene Louisinidae (Macroscelidea, Mammalia), their relationships and north European diversity. Zoological Journal of the Linnean Society 164: 856 - 936." type="journal article" year="2012">Hooker &amp; Russell, 2012</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBDD903FB40A886DCBDFE01" author="Rose KD &amp; Storch G &amp; Krohmann K" pageId="15" pageNumber="446" pagination="95 - 124" refId="ref19982" refString="Rose KD, Storch G, Krohmann K. 2015. Small-mammal postcrania from the Middle Paleocene of Walbeck, Germany. Palaontologische Zeitschrift 89: 95 - 124." type="journal article" year="2015">Rose, Storch &amp; Krohmann, 2015</bibRefCitation>
). Dental characters were scored for
<taxonomicName id="4C1F4D0BFFBDD903FA32A8E4DB0CFDE3" authority="Gingerich, 1983" authorityName="Gingerich" authorityYear="1983" class="Mammalia" genus="Diacocherus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Macroscelidea" pageId="15" pageNumber="446" phylum="Chordata" rank="species" species="meizon">
<emphasis id="B96BEA9AFFBDD903FA32A8E4DAD7FE00" box="[1418,1437,473,494]" italics="true" pageId="15" pageNumber="446">D</emphasis>
.
<emphasis id="B96BEA9AFFBDD903FC83A8C5DCC0FDE3" box="[827,906,504,525]" italics="true" pageId="15" pageNumber="446">meizon</emphasis>
Gingerich, 1983
</taxonomicName>
. Finally, the genus
<taxonomicName id="4C1F4D0BFFBDD903FA99A8CADB15FDC5" authority="Matthew &amp; Granger, 1921" authorityName="Matthew &amp; Granger" authorityYear="1921" class="Mammalia" family="Adapisoricidae" genus="Leptacodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Erinaceomorpha" pageId="15" pageNumber="446" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBDD903FA99A8CADAE8FDE2" box="[1313,1442,503,524]" italics="true" pageId="15" pageNumber="446">Leptacodon</emphasis>
<bibRefCitation id="EF8E4B79FFBDD903FC83AB2BDB15FDC5" author="Matthew WD &amp; Granger W" box="[827,1119,534,556]" pageId="15" pageNumber="446" pagination="1 - 7" refId="ref19167" refString="Matthew WD, Granger W. 1921. New genera of Paleocene mammals: American Museum Novitates, Vol. 13, 1 - 7." type="journal article" year="1921">Matthew &amp; Granger, 1921</bibRefCitation>
</taxonomicName>
was included in order to test for potential broader lipotyphlan affinities, exclusive of
<taxonomicName id="4C1F4D0BFFBDD903FC83AB6EDCB3FD87" box="[827,1017,595,617]" class="Mammalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Erinaceomorpha" pageId="15" pageNumber="446" phylum="Chordata" rank="order">Erinaceomorpha</taxonomicName>
. Scoring decisions for
<taxonomicName id="4C1F4D0BFFBDD903FB5FAB6EDA22FD86" authorityName="Matthew &amp; Granger" authorityYear="1921" box="[1255,1384,595,616]" class="Mammalia" family="Adapisoricidae" genus="Leptacodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Erinaceomorpha" pageId="15" pageNumber="446" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBDD903FB5FAB6EDA22FD86" box="[1255,1384,595,616]" italics="true" pageId="15" pageNumber="446">Leptacodon</emphasis>
</taxonomicName>
were based on
<taxonomicName id="4C1F4D0BFFBDD903FC11AB4EDA58FD66" authority="Simpson, 1935" authorityName="Simpson" authorityYear="1935" box="[937,1298,626,648]" class="Mammalia" family="Adapisoricidae" genus="Leptacodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Erinaceomorpha" pageId="15" pageNumber="446" phylum="Chordata" rank="species" species="munusculum">
<emphasis id="B96BEA9AFFBDD903FC11AB4EDCF3FD66" box="[937,953,627,648]" italics="true" pageId="15" pageNumber="446">L</emphasis>
.
<emphasis id="B96BEA9AFFBDD903FC7FAB4FDB15FD69" box="[967,1119,626,647]" italics="true" pageId="15" pageNumber="446">munusculum</emphasis>
<bibRefCitation id="EF8E4B79FFBDD903FBDFAB4FDA58FD66" author="Simpson GG" box="[1127,1298,626,648]" pageId="15" pageNumber="446" pagination="221 - 244" refId="ref20746" refString="Simpson GG. 1935. New Paleocene mammals from the Fort Union of Montana. Proceedings of the United States National Museum 83: 221 - 244." type="journal article" year="1935">Simpson, 1935</bibRefCitation>
</taxonomicName>
, a relatively well-known species that is considered to be among the basalmost members of
<taxonomicName id="4C1F4D0BFFBDD903FBF0AB92DAE8FD2B" authority="Simpson, 1928" authorityName="Simpson" authorityYear="1928" box="[1096,1442,687,709]" class="Mammalia" family="Nyctitheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Insectivora" pageId="15" pageNumber="446" phylum="Chordata" rank="family">
Nyctitheriidae
<bibRefCitation id="EF8E4B79FFBDD903FB41AB92DAE8FD2B" author="Simpson GG" box="[1273,1442,687,709]" pageId="15" pageNumber="446" pagination="1 - 15" refId="ref20720" refString="Simpson GG. 1928. A new mammalian fauna from the Fort Union of southern Montana. American Museum Novitates 297: 1 - 15." type="journal article" year="1928">Simpson, 1928</bibRefCitation>
</taxonomicName>
(see
<bibRefCitation id="EF8E4B79FFBDD903FCD5ABF2DB1AFD0A" author="Manz CL &amp; Bloch JI" box="[877,1104,718,740]" pageId="15" pageNumber="446" pagination="307 - 342" refId="ref18958" refString="Manz CL, Bloch JI. 2015. Systematics and phyl- ogeny of Paleocene - Eocene Nyctitheriidae (Mammalia, Eulipotyphla?) with description of a new species from the Late Paleocene of the Clarks Fork Basin, Wyoming, USA. Journal of Mammalian Evolution 22: 307 - 342." type="journal article" year="2015">Manz &amp; Bloch, 2015</bibRefCitation>
).
</paragraph>
<paragraph id="8BA03688FFBDD903FC83ABC6DB68FCFD" blockId="15.[827,1444,763,1891]" box="[827,1058,763,787]" pageId="15" pageNumber="446">
<emphasis id="B96BEA9AFFBDD903FC83ABC6DB68FCFD" box="[827,1058,763,787]" italics="true" pageId="15" pageNumber="446">Analytical protocol</emphasis>
</paragraph>
<paragraph id="8BA03688FFBDD91CFC83AA21DDF3FDA5" blockId="15.[827,1444,763,1891]" lastBlockId="16.[145,761,197,587]" lastPageId="16" lastPageNumber="447" pageId="15" pageNumber="446">
Four heuristic parsimony analyses were performed in PAUP4.0a (
<bibRefCitation id="EF8E4B79FFBDD903FC06AA07DB2CFCBE" author="Swofford DL" box="[958,1126,826,848]" pageId="15" pageNumber="446" refId="ref21084" refString="Swofford DL. 2002. PAUP *. Phylogenetic Analysis Using Parsimony (* and Other Methods). Version 4. Sunderland: Sinauer Associates." type="book" year="2002">Swofford, 2002</bibRefCitation>
) using the random addition sequence option; default conditions were used for all other heuristic search options. Multistate characters were unordered and multistate terminals were considered polymorphic. PAUP4.0a was also used to perform bootstrap analyses (default conditions) and to calculate Bremer support. Uncertainty with respect to the nearest relative of the ingroup and conflicting character states among potential outgroups prompted the decision to run separate analyses, with the resulting trees examined for any congruence in topology. The inclusion of
<taxonomicName id="4C1F4D0BFFBDD903FC32ADB1DB44FB4C" box="[906,1038,1164,1186]" class="Mammalia" family="Zhelestidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Proteutheria" pageId="15" pageNumber="446" phylum="Chordata" rank="family">Zhelestidae</taxonomicName>
as part of the ingroup poses several difficulties for outgroup choice: zhelestids were once considered near-relatives of ungulates (e.g.
<bibRefCitation id="EF8E4B79FFBDD903FA94ADF4DC39FB13" author="Archibald JD" pageId="15" pageNumber="446" pagination="292 - 331" refId="ref15846" refString="Archibald JD. 1998. Archaic ungulates (' Condylarthra'). In: Janis CM, Scott KM, Jacobs LL, eds. Evolution of tertiary mammals of North America, Vol. 1: terrestrial carnivores, ungulates, and ungulatelike mammals. Cambridge: Cambridge University Press, 292 - 331." type="book chapter" year="1998">Archibald, 1998</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBDD903FC39ADD5DB16FB13" author="Nessov LA &amp; Archibald JD &amp; Kielan-Jaworowska Z" box="[897,1116,1256,1277]" pageId="15" pageNumber="446" pagination="40 - 88" refId="ref19374" refString="Nessov LA, Archibald JD, Kielan-Jaworowska Z. 1998. Ungulate-like mammals from the Late Cretaceous of Uzbekistan and a phylogenetic analysis of Ungulatomorpha. Bulletin of Carnegie Museum of Natural History 34: 40 - 88." type="journal article" year="1998">
Nessov
<emphasis id="B96BEA9AFFBDD903FC64ADD5DB5CFB13" box="[988,1046,1256,1277]" italics="true" pageId="15" pageNumber="446">et al.</emphasis>
, 1998
</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBDD903FBD2ADD5DA2FFB13" author="Archibald JD &amp; Averianov AO &amp; Ekdale EG" box="[1130,1381,1256,1278]" pageId="15" pageNumber="446" pagination="62 - 65" refId="ref16053" refString="Archibald JD, Averianov AO, Ekdale EG. 2001. Late Cretaceous relatives of rabbits, rodents, and other extant eutherian mammals. Nature 414: 62 - 65." type="journal article" year="2001">
Archibald
<emphasis id="B96BEA9AFFBDD903FB5CADD5DA54FB13" box="[1252,1310,1256,1277]" italics="true" pageId="15" pageNumber="446">et al.</emphasis>
, 2001
</bibRefCitation>
), but subsequent analyses have positioned the family outside of Placentalia and, in some cases, basal to several stem eutherians, including
<taxonomicName id="4C1F4D0BFFBDD903FB89AC79DC38FA96" authority="Kielan-Jaworowska, 1969" authorityName="Kielan-Jaworowska" authorityYear="1969" class="Mammalia" family="Gypsonictopidae" genus="Kennalestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pilosa" pageId="15" pageNumber="446" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBDD903FB89AC79DBFDFAB7" box="[1073,1207,1348,1369]" italics="true" pageId="15" pageNumber="446">Kennalestes</emphasis>
<bibRefCitation id="EF8E4B79FFBDD903FB04AC79DC38FA96" author="Kielan-Jaworowska Z" pageId="15" pageNumber="446" pagination="171 - 191" refId="ref18385" refString="Kielan-Jaworowska Z. 1969. Preliminary data on the Upper Cretaceous eutherian mammals from Bayn Dzak, Gobi Desert. Palaeontologia Polonica 19: 171 - 191." type="journal article" year="1969">Kielan-Jaworowska, 1969</bibRefCitation>
</taxonomicName>
and
<taxonomicName id="4C1F4D0BFFBDD903FC14AC5FDC39FA78" authority="Wible, Novacek, Rougier &amp; Asher, 2007" authorityName="Wible, Novacek, Rougier &amp; Asher" authorityYear="2007" class="Mammalia" family="Cimolestidae" genus="Maelestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="15" pageNumber="446" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBDD903FC14AC5FDB50FA99" box="[940,1050,1378,1399]" italics="true" pageId="15" pageNumber="446">Maelestes</emphasis>
<bibRefCitation id="EF8E4B79FFBDD903FB99AC5FDC39FA78" author="Wible JR &amp; Rougier GW &amp; Novacek MJ &amp; Asher RJ" pageId="15" pageNumber="446" pagination="1003 - 1006" refId="ref21474" refString="Wible JR, Rougier GW, Novacek MJ, Asher RJ. 2007. Cretaceous eutherians and Laurasian origin for placental mammals near the K / T boundary. Nature 447: 1003 - 1006." type="journal article" year="2007">Wible, Novacek, Rougier &amp; Asher, 2007</bibRefCitation>
</taxonomicName>
(e.g.
<bibRefCitation id="EF8E4B79FFBDD903FC0BACBCDBCBFA78" author="Wible JR &amp; Rougier GW &amp; Novacek MJ &amp; Asher RJ" box="[947,1153,1409,1431]" pageId="15" pageNumber="446" pagination="1 - 123" refId="ref21510" refString="Wible JR, Rougier GW, Novacek MJ, Asher RJ. 2009. The eutherian mammal Maelestes gobiensis from the Late Cretaceous of Mongolia and the phylogeny of Cretaceous Eutheria. Bulletin of the American Museum of Natural History 327: 1 - 123." type="journal article" year="2009">
Wible
<emphasis id="B96BEA9AFFBDD903FC47ACBFDB70FA78" box="[1023,1082,1409,1431]" italics="true" pageId="15" pageNumber="446">et al.</emphasis>
, 2009
</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBDD903FB28ACBCDC38FA5B" author="Archibald JD &amp; Averianov AO" pageId="15" pageNumber="446" pagination="361 - 426" refId="ref15931" refString="Archibald JD, Averianov AO. 2012. Phylogenetic analysis, taxonomic revision and dental ontogeny of the Cretaceous Zhelestidae (Mammalia: Eutheria). Zoological Journal of the Linnean Society 164: 361 - 426." type="journal article" year="2012">Archibald &amp; Averianov, 2012</bibRefCitation>
). On the contrary, more recent large-scale analyses of many of the relevant taxa included in this analysis recover
<taxonomicName id="4C1F4D0BFFBDD903FC61ACE0DB28FA1D" box="[985,1122,1501,1523]" class="Mammalia" family="Zhelestidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Proteutheria" pageId="15" pageNumber="446" phylum="Chordata" rank="family">Zhelestidae</taxonomicName>
crownward of
<taxonomicName id="4C1F4D0BFFBDD903FAAEACE0DAE8FA1C" authorityName="Kielan-Jaworowska" authorityYear="1969" box="[1302,1442,1501,1522]" class="Mammalia" family="Gypsonictopidae" genus="Kennalestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pilosa" pageId="15" pageNumber="446" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBDD903FAAEACE0DAE8FA1C" box="[1302,1442,1501,1522]" italics="true" pageId="15" pageNumber="446">Kennalestes</emphasis>
</taxonomicName>
and
<taxonomicName id="4C1F4D0BFFBDD903FCD6ACC1DC94F9FF" authorityName="Wible, Novacek, Rougier &amp; Asher" authorityYear="2007" box="[878,990,1532,1553]" class="Mammalia" family="Cimolestidae" genus="Maelestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="15" pageNumber="446" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBDD903FCD6ACC1DC94F9FF" box="[878,990,1532,1553]" italics="true" pageId="15" pageNumber="446">Maelestes</emphasis>
</taxonomicName>
(
<bibRefCitation id="EF8E4B79FFBDD903FC57ACC1DBCDF9FF" author="Hooker JJ" box="[1007,1159,1532,1554]" pageId="15" pageNumber="446" pagination="1 - 79" refId="ref18163" refString="Hooker JJ. 2014. New postcranial bones of the extinct mammalian family Nyctitheriidae (Paleogene, UK): primitive euarchontans with scansorial locomotion. Palaeontologia Electronica 17: 1 - 79." type="journal article" year="2014">Hooker, 2014</bibRefCitation>
;
<bibRefCitation id="EF8E4B79FFBDD903FB2EACC1DADEF9FC" author="Penkrot TA &amp; Zack SP" box="[1174,1428,1532,1554]" pageId="15" pageNumber="446" pagination="1212059" refId="ref19622" refString="Penkrot TA, Zack SP. 2016. Tarsals of Sespedectinae (? Lipotyphla) from the Middle Eocene of southern California, and the affinities of Eocene ' erinaceomorphs'. Journal of Vertebrate Palaeontology 36: e 1212059." type="journal article" year="2016">Penkrot &amp; Zack, 2016</bibRefCitation>
). Given these uncertainties, three analyses were run, each with a different outgroup. In the first analysis, the Late Cretaceous therian
<taxonomicName id="4C1F4D0BFFBDD903FB3FAF65DA48F965" authority="Kielan-Jaworowska &amp; Dashzeveg, 1989" authorityName="Kielan-Jaworowska &amp; Dashzeveg" authorityYear="1989" class="Mammalia" family="Gypsonictopidae" genus="Prokennalestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pilosa" pageId="15" pageNumber="446" phylum="Chordata" rank="species" species="trofimovi">
<emphasis id="B96BEA9AFFBDD903FB3FAF65DAE9F983" box="[1159,1443,1624,1645]" italics="true" pageId="15" pageNumber="446">Prokennalestes trofimovi</emphasis>
<bibRefCitation id="EF8E4B79FFBDD903FC83AF4BDA48F965" author="Kielan-Jaworowska Z &amp; Dashzeveg D" box="[827,1282,1654,1676]" pageId="15" pageNumber="446" pagination="347 - 355" refId="ref18413" refString="Kielan-Jaworowska Z, Dashzeveg D. 1989. Eutherian mammals from the Early Cretaceous of Mongolia. Zoologica Scripta 18: 347 - 355." type="journal article" year="1989">Kielan-Jaworowska &amp; Dashzeveg, 1989</bibRefCitation>
</taxonomicName>
served as the outgroup;
<taxonomicName id="4C1F4D0BFFBDD903FC0FAFA8DB21F944" box="[951,1131,1685,1706]" class="Mammalia" family="Gypsonictopidae" genus="Prokennalestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pilosa" pageId="15" pageNumber="446" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFBDD903FC0FAFA8DB21F944" box="[951,1131,1685,1706]" italics="true" pageId="15" pageNumber="446">Prokennalestes</emphasis>
</taxonomicName>
has in previous analyses been consistently recovered stemward of the ingroup taxa included here, including
<taxonomicName id="4C1F4D0BFFBDD903FB22AFEFDA6BF906" box="[1178,1313,1746,1768]" class="Mammalia" family="Zhelestidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Proteutheria" pageId="15" pageNumber="446" phylum="Chordata" rank="family">Zhelestidae</taxonomicName>
(e.g.
<bibRefCitation id="EF8E4B79FFBDD903FAD8AFEFDCFBF8E8" author="Wible JR &amp; Rougier GW &amp; Novacek MJ &amp; Asher RJ" pageId="15" pageNumber="446" pagination="1 - 123" refId="ref21510" refString="Wible JR, Rougier GW, Novacek MJ, Asher RJ. 2009. The eutherian mammal Maelestes gobiensis from the Late Cretaceous of Mongolia and the phylogeny of Cretaceous Eutheria. Bulletin of the American Museum of Natural History 327: 1 - 123." type="journal article" year="2009">
Wible
<emphasis id="B96BEA9AFFBDD903FC83AFCFDC3AF8E8" box="[827,880,1777,1799]" italics="true" pageId="15" pageNumber="446">et al.</emphasis>
, 2009
</bibRefCitation>
). The second and third analyses included the Late Cretaceous therian
<taxonomicName id="4C1F4D0BFFBDD903FBEBAE2DDA0DF8CB" authorityName="Kielan-Jaworowska" authorityYear="1968" box="[1107,1351,1808,1829]" class="Mammalia" family="Gypsonictopidae" genus="Kennalestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pilosa" pageId="15" pageNumber="446" phylum="Chordata" rank="species" species="gobiensis">
<emphasis id="B96BEA9AFFBDD903FBEBAE2DDA0DF8CB" box="[1107,1351,1808,1829]" italics="true" pageId="15" pageNumber="446">Kennalestes gobiensis</emphasis>
</taxonomicName>
and the Late Cretaceous therian
<taxonomicName id="4C1F4D0BFFBDD903FBE3AE13DA70F8AD" box="[1115,1338,1838,1859]" class="Mammalia" family="Cimolestidae" genus="Maelestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="15" pageNumber="446" phylum="Chordata" rank="species" species="gobiensis">
<emphasis id="B96BEA9AFFBDD903FBE3AE13DA70F8AD" box="[1115,1338,1838,1859]" italics="true" pageId="15" pageNumber="446">Maelestes gobiensis</emphasis>
</taxonomicName>
, respectively, as outgroups. Given the
<taxonomicName id="4C1F4D0BFFBDD903FB20AE70DA42F88D" box="[1176,1288,1869,1891]" order="Uncertain" pageId="15" pageNumber="434" rank="order">uncertain</taxonomicName>
phylogenetic position of
<taxonomicName id="4C1F4D0BFFA2D91CFEA1A9F8DEEEFF35" box="[281,420,197,219]" class="Mammalia" family="Zhelestidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Proteutheria" pageId="16" pageNumber="447" phylum="Chordata" rank="family">Zhelestidae</taxonomicName>
relative to
<taxonomicName id="4C1F4D0BFFA2D91CFD8DA9F8DD89FF34" authorityName="Kielan-Jaworowska" authorityYear="1969" box="[565,707,197,218]" class="Mammalia" family="Gypsonictopidae" genus="Kennalestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pilosa" pageId="16" pageNumber="447" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFA2D91CFD8DA9F8DD89FF34" box="[565,707,197,218]" italics="true" pageId="16" pageNumber="447">Kennalestes</emphasis>
</taxonomicName>
and
<taxonomicName id="4C1F4D0BFFA2D91CFF29A9D9DFB4FF17" authorityName="Wible, Novacek, Rougier &amp; Asher" authorityYear="2007" box="[145,254,228,249]" class="Mammalia" family="Cimolestidae" genus="Maelestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="16" pageNumber="447" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFA2D91CFF29A9D9DFB4FF17" box="[145,254,228,249]" italics="true" pageId="16" pageNumber="447">Maelestes</emphasis>
</taxonomicName>
, two subanalyses were performed with each outgroup in order to examine the effects of including or excluding
<taxonomicName id="4C1F4D0BFFA2D91CFE9DA81CDEE0FED9" box="[293,426,289,311]" class="Mammalia" family="Zhelestidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Proteutheria" pageId="16" pageNumber="447" phylum="Chordata" rank="family">Zhelestidae</taxonomicName>
from the ingroup. Four characters (78–81) were excluded from the subanalyses where
<taxonomicName id="4C1F4D0BFFA2D91CFF58A863DE2DFE9A" box="[224,359,350,372]" class="Mammalia" family="Zhelestidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Proteutheria" pageId="16" pageNumber="447" phylum="Chordata" rank="family">Zhelestidae</taxonomicName>
were excluded, as states for these characters were rendered either invariable for all taxa, or invariable in the ingroup. A fourth analysis was performed using
<taxonomicName id="4C1F4D0BFFA2D91CFE30A887DD78FE21" box="[392,562,442,463]" class="Mammalia" family="Gypsonictopidae" genus="Prokennalestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pilosa" pageId="16" pageNumber="447" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFA2D91CFE30A887DD78FE21" box="[392,562,442,463]" italics="true" pageId="16" pageNumber="447">Prokennalestes</emphasis>
</taxonomicName>
,
<taxonomicName id="4C1F4D0BFFA2D91CFD87A887DD8CFE21" authorityName="Kielan-Jaworowska" authorityYear="1969" box="[575,710,442,463]" class="Mammalia" family="Gypsonictopidae" genus="Kennalestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pilosa" pageId="16" pageNumber="447" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFA2D91CFD87A887DD8CFE21" box="[575,710,442,463]" italics="true" pageId="16" pageNumber="447">Kennalestes</emphasis>
</taxonomicName>
and
<taxonomicName id="4C1F4D0BFFA2D91CFF29A8E4DE4BFE00" authorityName="Wible, Novacek, Rougier &amp; Asher" authorityYear="2007" box="[145,257,473,494]" class="Mammalia" family="Cimolestidae" genus="Maelestes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cimolesta" pageId="16" pageNumber="447" phylum="Chordata" rank="genus">
<emphasis id="B96BEA9AFFA2D91CFF29A8E4DE4BFE00" box="[145,257,473,494]" italics="true" pageId="16" pageNumber="447">Maelestes</emphasis>
</taxonomicName>
together in order to examine the effects of multiple outgroups on the resulting topology. Character states were optimized using the DELTRAN option, a criterion that favours convergence over reversal.
</paragraph>
</subSubSection>
</treatment>
</document>
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