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<document ID-DOI="10.1093/zoolinnean/zlac019" ID-ISSN="0024-4082" ID-Zenodo-Dep="7381112" approvalRequired="1" checkinTime="1669807628251" checkinUser="plazi" docAuthor="Corrie, Joshua E &amp; Fordyce, R Ewan" docDate="2022" docId="03F3AE42FF96FFE6FC6902D0FB7E37CA" docLanguage="en" docName="zlac019.pdf" docOrigin="Zoological Journal of the Linnean Society 196 (4)" docSource="https://academic.oup.com/zoolinnean/article/196/4/1637/6598844" docStyle="DocumentStyle:36B3BD6A90C22AB4F7F465C853188CC8.7:ZoolJLinnSoc.2017-.journal_article" docStyleId="36B3BD6A90C22AB4F7F465C853188CC8" docStyleName="ZoolJLinnSoc.2017-.journal_article" docStyleVersion="7" docTitle="Kekenodon onamata Beds (McKay 1882" docType="treatment" docVersion="3" lastPageNumber="1661" masterDocId="FFCAD63AFF85FFFEFFA00408FFDF361A" masterDocTitle="A redescription and re-evaluation of Kekenodon onamata (Mammalia: Cetacea), a late-surviving archaeocete from the Late Oligocene of New Zealand" masterLastPageNumber="1670" masterPageNumber="1637" pageNumber="1656" updateTime="1669905974674" updateUser="diego">
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<mods:titleInfo>
<mods:title>A redescription and re-evaluation of Kekenodon onamata (Mammalia: Cetacea), a late-surviving archaeocete from the Late Oligocene of New Zealand</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Corrie, Joshua E</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Fordyce, R Ewan</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>Zoological Journal of the Linnean Society</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2022</mods:date>
<mods:detail type="pubDate">
<mods:number>2022-12-01</mods:number>
</mods:detail>
<mods:detail type="volume">
<mods:number>196</mods:number>
</mods:detail>
<mods:detail type="issue">
<mods:number>4</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>1637</mods:start>
<mods:end>1670</mods:end>
</mods:extent>
</mods:part>
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<mods:location>
<mods:url>https://academic.oup.com/zoolinnean/article/196/4/1637/6598844</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">10.1093/zoolinnean/zlac019</mods:identifier>
<mods:identifier type="ISSN">0024-4082</mods:identifier>
<mods:identifier type="Zenodo-Dep">7381112</mods:identifier>
</mods:mods>
<treatment LSID="urn:lsid:plazi:treatment:03F3AE42FF96FFE6FC6902D0FB7E37CA" httpUri="http://treatment.plazi.org/id/03F3AE42FF96FFE6FC6902D0FB7E37CA" lastPageId="24" lastPageNumber="1661" pageId="19" pageNumber="1656">
<subSubSection lastPageId="20" lastPageNumber="1657" pageId="19" pageNumber="1656" type="nomenclature">
<paragraph blockId="19.[969,1300,1751,1776]" box="[969,1300,1751,1776]" pageId="19" pageNumber="1656">
<heading box="[969,1300,1751,1776]" centered="true" fontSize="9" level="2" pageId="19" pageNumber="1656" reason="2">
<taxonomicName authorityName="Beds (McKay" authorityYear="1882" box="[969,1300,1751,1776]" class="Mammalia" family="Basilosauridae" genus="Kekenodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="19" pageNumber="1666" phylum="Chordata" rank="genus">
<emphasis box="[969,1113,1752,1775]" italics="true" pageId="19" pageNumber="1656">KEKENODON</emphasis>
CF.
<emphasis box="[1161,1180,1752,1775]" italics="true" pageId="19" pageNumber="1656">K</emphasis>
.
<emphasis box="[1193,1300,1755,1775]" italics="true" pageId="19" pageNumber="1656">ONAMATA</emphasis>
</taxonomicName>
</heading>
</paragraph>
<paragraph blockId="19.[827,1442,1824,1907]" lastBlockId="20.[145,760,1593,1738]" lastPageId="20" lastPageNumber="1657" pageId="19" pageNumber="1656">
<emphasis box="[827,1049,1824,1845]" italics="true" pageId="19" pageNumber="1656">Referred specimen:</emphasis>
<materialsCitation collectionCode="AUGD" collectorName="Oruawharo Road" country="New Zealand" location="Port Albert" municipality="North Island" pageId="19" pageNumber="1656" specimenCode="AUGD 2469" specimenCount="1">
<specimenCode box="[1062,1207,1824,1846]" collectionCode="AUGD" country="0" httpUri="http://grbio.org/cool/2hr6-2fnn" name="Department of Geology and Petroleum Geology" pageId="19" pageNumber="1656">AUGD 2469</specimenCode>
, partial denticulate tooth crown, from
<collectorName box="[1048,1254,1854,1876]" pageId="19" pageNumber="1656">Oruawharo Road</collectorName>
,
<emphasis box="[1268,1285,1855,1876]" italics="true" pageId="19" pageNumber="1656">c.</emphasis>
<quantity box="[1294,1354,1854,1876]" metricMagnitude="3" metricUnit="m" metricValue="5.0" pageId="19" pageNumber="1656" unit="km" value="5.0">5 km</quantity>
east of
<location LSID="urn:lsid:plazi:treatment:03F3AE42FF96FFE6FC6902D0FB7E37CA:8E85498FFF96FFEDFC9B0356FC603169" box="[827,959,1885,1907]" country="New Zealand" municipality="North Island" name="Port Albert" pageId="19" pageNumber="1656">Port Albert</location>
,
<collectingMunicipality box="[972,1125,1885,1907]" pageId="19" pageNumber="1656">North Island</collectingMunicipality>
,
<collectingCountry box="[1137,1294,1885,1907]" name="New Zealand" pageId="19" pageNumber="1656">New Zealand</collectingCountry>
</materialsCitation>
,
<materialsCitation collectingDate="1969-07" collectionCode="AUGD" collectorName="M. Dowson" country="New Zealand" lastPageId="20" lastPageNumber="1657" location="New Zealand" pageId="19" pageNumber="1656" specimenCount="1">
collected by
<collectorName box="[145,275,1593,1614]" pageId="20" pageNumber="1657">M. Dowson</collectorName>
in
<date box="[314,432,1593,1615]" pageId="20" pageNumber="1657" value="1969-07">
<collectingDate box="[314,432,1593,1615]" pageId="20" pageNumber="1657" value="1969-07">July 1969</collectingDate>
</date>
.
<collectingCountry box="[443,598,1593,1615]" name="New Zealand" pageId="20" pageNumber="1657">New Zealand</collectingCountry>
</materialsCitation>
<materialsCitation collectionCode="AUGD" collectorName="Map Series" country="New Zealand" latitude="-36.2637" longLatPrecision="5" longitude="174.46725" pageId="20" pageNumber="1657" specimenCode="NZGD49" specimenCount="1">
<collectorName box="[606,739,1593,1615]" pageId="20" pageNumber="1657">Map Series</collectorName>
1 N28:985316, equivalent to
<geoCoordinate box="[476,597,1624,1646]" degrees="36.2637" direction="south" orientation="latitude" pageId="20" pageNumber="1657" precision="5" value="-36.2637">36.2637ºS</geoCoordinate>
,
<geoCoordinate box="[611,760,1623,1645]" degrees="174.46725" direction="east" orientation="longitude" pageId="20" pageNumber="1657" precision="1" value="174.46725">174.46725ºE</geoCoordinate>
(
<specimenCode box="[152,260,1654,1675]" collectionCode="NZGD" pageId="20" pageNumber="1657">NZGD49</specimenCode>
)
</materialsCitation>
.
<materialsCitation collectionCode="NZGD" collectorName="Fossil" country="New Zealand" location="Geosciences Society" pageId="20" pageNumber="1657" specimenCount="1">
<collectorName box="[279,346,1654,1676]" pageId="20" pageNumber="1657">Fossil</collectorName>
record number Q09/f7902 (original number N28/f0902;
<collectingCountry box="[388,548,1685,1707]" name="New Zealand" pageId="20" pageNumber="1657">New Zealand</collectingCountry>
fossil record file,
<location LSID="urn:lsid:plazi:treatment:03F3AE42FF96FFE6FC6902D0FB7E37CA:8E85498FFF91FFEAFF3102BCFEAD30D0" box="[145,370,1716,1738]" country="New Zealand" name="Geosciences Society" pageId="20" pageNumber="1657">Geosciences Society</location>
of
<collectingCountry box="[404,558,1716,1738]" name="New Zealand" pageId="20" pageNumber="1657">New Zealand</collectingCountry>
)
</materialsCitation>
.
</paragraph>
</subSubSection>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7381132" ID-Zenodo-Dep="7381132" httpUri="https://zenodo.org/record/7381132/files/figure.png" pageId="20" pageNumber="1657" startId="20.[145,226,1503,1525]" targetBox="[147,1419,198,1461]" targetPageId="20">
<paragraph blockId="20.[145,1424,1503,1555]" pageId="20" pageNumber="1657">
<emphasis bold="true" box="[145,253,1503,1525]" pageId="20" pageNumber="1657">Figure 9.</emphasis>
Atlas of
<taxonomicName authorityName=", Hector" authorityYear="1881" box="[357,577,1503,1525]" class="Mammalia" family="Basilosauridae" genus="Kekenodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="20" pageNumber="1657" phylum="Chordata" rank="species" species="onamata">
<emphasis box="[357,577,1503,1525]" italics="true" pageId="20" pageNumber="1657">Kekenodon onamata</emphasis>
</taxonomicName>
(
<emphasis box="[592,756,1503,1525]" italics="true" pageId="20" pageNumber="1657">NMNZ Ma 306</emphasis>
). A, anterior view. B, posterior view. Specimen whitened with ammonium chloride.
</paragraph>
</caption>
<subSubSection lastPageId="24" lastPageNumber="1661" pageId="20" pageNumber="1657" type="description">
<paragraph blockId="20.[145,759,1778,1892]" lastBlockId="20.[809,1426,1593,1891]" pageId="20" pageNumber="1657">
<emphasis box="[145,423,1778,1800]" italics="true" pageId="20" pageNumber="1657">Stratigraphy and age:</emphasis>
AUGD 2469 was collected near, and is presumably from, the Mahurangi Limestone, underlying the Miocene Waitemata Group. Matrix from the Mahurangi Limestone consists of whitish grey, fine-grained, sparsely macrofossiliferous and heavily bioturbated rock. Matrix collected from the limestone did not yield the lower Whaingaroan planktonic foraminifera
<taxonomicName authority="Hornibrook, 1965" authorityName="Hornibrook" authorityYear="1965" class="Globothalamea" family="Globigerinidae" genus="Globigerina" kingdom="Chromista" order="Rotaliida" pageId="20" pageNumber="1657" phylum="Foraminifera" rank="species" species="angiporoides" subGenus="Subbotina">
<emphasis box="[809,953,1716,1737]" italics="true" pageId="20" pageNumber="1657">Globigerina</emphasis>
(
<emphasis box="[971,1096,1716,1737]" italics="true" pageId="20" pageNumber="1657">Subbotina</emphasis>
)
<emphasis box="[1114,1270,1716,1737]" italics="true" pageId="20" pageNumber="1657">angiporoides</emphasis>
Hornibrook, 1965
</taxonomicName>
. The presence of
<taxonomicName authority="Jenkins, 1960" authorityName="Jenkins" authorityYear="1960" class="Globothalamea" family="Globigerinidae" genus="Globigerina" kingdom="Chromista" order="Rotaliida" pageId="20" pageNumber="1657" phylum="Foraminifera" rank="species" species="euapertura">
<emphasis box="[1114,1425,1746,1768]" italics="true" pageId="20" pageNumber="1657">Globigerina euapertura</emphasis>
Jenkins, 1960
</taxonomicName>
indicates an upper Whaingaroan- Waitakian (
<emphasis box="[974,985,1808,1829]" italics="true" pageId="20" pageNumber="1657">c</emphasis>
. 28.1 21.7 Mya;
<bibRefCitation author="Carter L" box="[1233,1406,1808,1830]" pageId="20" pageNumber="1657" pagination="104 - 118" refId="ref22766" refString="Carter L. 1969. The Mahurangi limestone rom Puketotara Peninsula, Northland, New Zealand. New Zealand Journal of Geology and Geophysics 12: 104 - 118." type="journal article" year="1969">Carter, 1969</bibRefCitation>
), contemporaneous with the
<taxonomicName authority="Beds" authorityName="Beds (McKay" authorityYear="1882" box="[1156,1362,1838,1860]" class="Mammalia" family="Basilosauridae" genus="Kekenodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="20" pageNumber="1657" phylum="Chordata" rank="genus">
<emphasis box="[1156,1291,1838,1859]" italics="true" pageId="20" pageNumber="1657">Kekenodon</emphasis>
Beds
</taxonomicName>
that produced
<emphasis box="[926,1106,1869,1891]" italics="true" pageId="20" pageNumber="1657">NMNZ Ma 306</emphasis>
.
</paragraph>
<paragraph blockId="21.[163,303,198,221]" box="[163,303,198,221]" pageId="21" pageNumber="1658">
<emphasis box="[163,303,198,221]" italics="true" pageId="21" pageNumber="1658">Description</emphasis>
</paragraph>
<paragraph blockId="21.[163,779,237,811]" pageId="21" pageNumber="1658">
<emphasis box="[163,289,238,259]" italics="true" pageId="21" pageNumber="1658">Dentition:</emphasis>
A single, denticulate tooth crown is preserved (
<figureCitation box="[294,373,268,290]" captionStart="Figure 10" captionStartId="21.[160,238,1803,1825]" captionTargetBox="[162,785,1435,1763]" captionTargetId="figure-626@21.[165,773,1462,1764]" captionTargetPageId="21" captionText="Figure 10. Partial crown of a possible left p2 of Kekenodon cf. K. onamata (AUGD 2469) in presumed labial (left) and lingual (right) view. Specimen whitened with ammonium chloride." figureDoi="http://doi.org/10.5281/zenodo.7381134" httpUri="https://zenodo.org/record/7381134/files/figure.png" pageId="21" pageNumber="1658">Fig. 10</figureCitation>
). The apex of the primary denticle is missing and crushing and cracks are prolific on all surfaces of the crown, indicating post-mortem deformation. The crown has a tall, triangular profile that is vertically oriented. A prominent anterior carina is present on the primary denticle, which is markedly larger than all preserved accessory denticles. The lower-half of the anterior surface of the crown is eroded, although the profile of the tooth suggests at least one accessory denticle was present just above the enamelodentin junction.Two large and robust posterior accessory denticles are preserved with possible additional denticles (now lost) located more basally. There is no indication of an ecto- or entocingulum. The enamel on one surface of the crown displays a slight wrinkled ornament, which appears natural, possibly indicating the lingual surface of the crown. In contrast, enamel on the inferred labial surface is smooth. When
</paragraph>
<paragraph blockId="21.[827,1443,197,710]" pageId="21" pageNumber="1658">
compared with basilosaurids and
<emphasis box="[1217,1392,197,219]" italics="true" pageId="21" pageNumber="1658">NMNZ Ma 306</emphasis>
, the presence of accessory denticles combined with the tall, triangular profile of the crown is suggestive of a premolar. Moreover, the arrangement of accessory denticles is characteristic of a posterior premolar or anterior molar. There is no evidence of a re-entrant groove on the anterior surface of the crown to suggest the tooth is a lower molar, as in basilosaurids. However, re-entrant grooves are largely absent from the molars of heterodont cetaceans in the Oligocene, making its use here in tooth identification dubious. The crown lacks the lingual curvature of the upper denticulate teeth of
<emphasis box="[1054,1226,565,587]" italics="true" pageId="21" pageNumber="1658">NMNZ Ma 306</emphasis>
, indicating a lower tooth. Despite roots being absent, the tooth was almost certainly double-rooted; there is no indication of a third lingual root. Given the preserved characteristics, the tooth is provisionally identified as a left p2.
</paragraph>
<paragraph blockId="21.[827,1444,750,1723]" pageId="21" pageNumber="1658">
<emphasis box="[827,934,750,771]" italics="true" pageId="21" pageNumber="1658">Remarks:</emphasis>
AUGD2469resembles preserved denticulate teeth belonging to the
<typeStatus box="[1081,1178,781,803]" pageId="21" pageNumber="1658">holotype</typeStatus>
of
<taxonomicName authorityName=", Hector" authorityYear="1881" box="[1212,1442,781,803]" class="Mammalia" family="Basilosauridae" genus="Kekenodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="21" pageNumber="1658" phylum="Chordata" rank="species" species="onamata">
<emphasis box="[1212,1442,781,803]" italics="true" pageId="21" pageNumber="1658">Kekenodon onamata</emphasis>
</taxonomicName>
(
<emphasis box="[835,1009,812,833]" italics="true" pageId="21" pageNumber="1658">NMNZ Ma 306</emphasis>
). The size of the crown is larger than all preserved teeth of
<emphasis box="[1084,1261,842,864]" italics="true" pageId="21" pageNumber="1658">NMNZ Ma 306</emphasis>
, but the profile and spacing of the accessory denticles is similar to
<taxonomicName authorityName=", Hector" authorityYear="1881" box="[827,966,904,925]" class="Mammalia" family="Basilosauridae" genus="Kekenodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="21" pageNumber="1658" phylum="Chordata" rank="species" species="onamata">
<emphasis box="[827,845,904,925]" italics="true" pageId="21" pageNumber="1658">K</emphasis>
.
<emphasis box="[861,966,904,925]" italics="true" pageId="21" pageNumber="1658">onamata</emphasis>
</taxonomicName>
. Alternatively, the tooth may belong to a toothed mysticete similar to what is known from the Oligocene of South Carolina, such as
<taxonomicName class="Mammalia" genus="Coronodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="21" pageNumber="1658" phylum="Chordata" rank="species" species="havensteini">
<emphasis italics="true" pageId="21" pageNumber="1658">Coronodon havensteini</emphasis>
</taxonomicName>
and the unpublished specimens ChM PV 5720 and ChM PV 4745 (
<bibRefCitation author="Geisler JH &amp; Boessenecker RW &amp; Brown M &amp; Beatty BL" box="[1108,1318,1026,1048]" pageId="21" pageNumber="1658" pagination="20361 - 2042" refId="ref24197" refString="Geisler JH, Boessenecker RW, Brown M, Beatty BL. 2017. The origin of filter feeding in whales. Current Biology 27: 20361 - 2042. e 2." type="journal article" year="2017">
Geisler
<emphasis box="[1196,1251,1026,1048]" italics="true" pageId="21" pageNumber="1658">et al.</emphasis>
, 2017
</bibRefCitation>
). However, there is a current lack of fossil evidence to indicate that large body-sized (&gt;
<quantity box="[1136,1186,1087,1109]" metricMagnitude="0" metricUnit="m" metricValue="5.0" pageId="21" pageNumber="1658" unit="m" value="5.0">5 m</quantity>
), toothed mysticetes with relatively large teeth were present in Oligocene
<collectingCountry box="[827,978,1149,1171]" name="New Zealand" pageId="21" pageNumber="1658">New Zealand</collectingCountry>
marine environments. Affinities with
<taxonomicName class="Mammalia" genus="Coronodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="21" pageNumber="1658" phylum="Chordata" rank="species" species="havensteini">
<emphasis box="[1419,1437,1149,1170]" italics="true" pageId="21" pageNumber="1658">C</emphasis>
.
<emphasis box="[827,962,1180,1201]" italics="true" pageId="21" pageNumber="1658">havensteini</emphasis>
</taxonomicName>
are further unlikely given the complete absence of enamel ornament on its cheek teeth. In contrast, small body-sized toothed mysticetes (e.g.
<taxonomicName authority="Fordyce &amp; Marx, 2016" authorityName="Fordyce &amp; Marx" authorityYear="2016" box="[827,1393,1271,1294]" class="Mammalia" family="Squalodontidae" genus="Mammalodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="21" pageNumber="1658" phylum="Chordata" rank="species" species="hakataramea">
<emphasis box="[827,1142,1272,1293]" italics="true" pageId="21" pageNumber="1658">Mammalodon hakataramea</emphasis>
<bibRefCitation author="Fordyce RE &amp; Marx FG" box="[1147,1393,1271,1294]" pageId="21" pageNumber="1658" pagination="107 - 116" refId="ref23893" refString="Fordyce RE, Marx FG. 2016. Mysticetes baring their teeth: new fossil whale, Mammalodon hakataramea, from the southwest Pacific. Memoirs of the Museum of Victoria 74: 107 - 116." type="journal article" year="2016">Fordyce &amp; Marx, 2016</bibRefCitation>
</taxonomicName>
) are known from Late Oligocene
<collectingCountry box="[1155,1308,1302,1324]" name="New Zealand" pageId="21" pageNumber="1658">New Zealand</collectingCountry>
(
<bibRefCitation author="Fordyce RE &amp; Marx FG" pageId="21" pageNumber="1658" pagination="107 - 116" refId="ref23893" refString="Fordyce RE, Marx FG. 2016. Mysticetes baring their teeth: new fossil whale, Mammalodon hakataramea, from the southwest Pacific. Memoirs of the Museum of Victoria 74: 107 - 116." type="journal article" year="2016">Fordyce &amp; Marx, 2016</bibRefCitation>
). Affinities with
<taxonomicName baseAuthorityName="Delfortrie" baseAuthorityYear="1873" box="[1148,1392,1333,1355]" class="Mammalia" genus="Phococetus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="21" pageNumber="1658" phylum="Chordata" rank="species" species="vasconum">
<emphasis box="[1148,1392,1333,1355]" italics="true" pageId="21" pageNumber="1658">Phococetus vasconum</emphasis>
</taxonomicName>
and cf.
<taxonomicName box="[860,1028,1363,1385]" class="Mammalia" genus="Phococetus" kingdom="Animalia" order="Cetacea" pageId="21" pageNumber="1658" phylum="Chordata" rank="species" species="undetermined">
<emphasis box="[860,987,1363,1384]" italics="true" pageId="21" pageNumber="1658">Phococetus</emphasis>
sp.
</taxonomicName>
, in addition to
<taxonomicName box="[1208,1391,1364,1385]" class="Mammalia" family="Inticetidae" genus="Inticetus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="21" pageNumber="1658" phylum="Chordata" rank="species" species="vertizi">
<emphasis box="[1208,1391,1364,1385]" italics="true" pageId="21" pageNumber="1658">Inticetus vertizi</emphasis>
</taxonomicName>
and
<taxonomicName box="[827,926,1395,1416]" class="Mammalia" family="Inticetidae" genus="Inticetus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="21" pageNumber="1658" phylum="Chordata" rank="genus">
<emphasis box="[827,926,1395,1416]" italics="true" pageId="21" pageNumber="1658">Inticetus</emphasis>
</taxonomicName>
-grade odontocetes, are unlikely for structural disparities previously discussed with
<taxonomicName authorityName=", Hector" authorityYear="1881" box="[1259,1392,1425,1447]" class="Mammalia" family="Basilosauridae" genus="Kekenodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="21" pageNumber="1658" phylum="Chordata" rank="species" species="onamata">
<emphasis box="[1259,1277,1426,1447]" italics="true" pageId="21" pageNumber="1658">K</emphasis>
.
<emphasis box="[1291,1392,1426,1447]" italics="true" pageId="21" pageNumber="1658">onamata</emphasis>
</taxonomicName>
and the absence of large body-sized odontocetes from the fossil record of Late Oligocene
<collectingCountry box="[1178,1332,1486,1508]" name="New Zealand" pageId="21" pageNumber="1658">New Zealand</collectingCountry>
. Thus far,
<taxonomicName authorityName=", Hector" authorityYear="1881" box="[827,963,1517,1538]" class="Mammalia" family="Basilosauridae" genus="Kekenodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="21" pageNumber="1658" phylum="Chordata" rank="species" species="onamata">
<emphasis box="[827,845,1517,1538]" italics="true" pageId="21" pageNumber="1658">K</emphasis>
.
<emphasis box="[860,963,1517,1538]" italics="true" pageId="21" pageNumber="1658">onamata</emphasis>
</taxonomicName>
possesses the largest teeth, and in turn one of the largest body sizes, in the cetacean fossil record from the Oligocene of
<collectingCountry box="[1188,1351,1578,1600]" name="New Zealand" pageId="21" pageNumber="1658">New Zealand</collectingCountry>
. AUGD 2469 is conservatively identified as
<taxonomicName authorityName="Beds (McKay" authorityYear="1882" box="[1250,1377,1609,1630]" class="Mammalia" family="Basilosauridae" genus="Kekenodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="21" pageNumber="1658" phylum="Chordata" rank="genus">
<emphasis box="[1250,1377,1609,1630]" italics="true" pageId="21" pageNumber="1658">Kekenodon</emphasis>
</taxonomicName>
cf.
<taxonomicName authorityName=", Hector" authorityYear="1881" class="Mammalia" family="Basilosauridae" genus="Kekenodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="21" pageNumber="1658" phylum="Chordata" rank="species" species="onamata">
<emphasis box="[1418,1436,1609,1630]" italics="true" pageId="21" pageNumber="1658">K</emphasis>
.
<emphasis box="[827,931,1640,1661]" italics="true" pageId="21" pageNumber="1658">onamata</emphasis>
</taxonomicName>
based on the structural similarities of the tooth and its contemporaneous stratigraphic position with the
<typeStatus box="[929,1026,1701,1723]" pageId="21" pageNumber="1658">holotype</typeStatus>
of
<taxonomicName authorityName=", Hector" authorityYear="1881" box="[1060,1291,1701,1722]" class="Mammalia" family="Basilosauridae" genus="Kekenodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="21" pageNumber="1658" phylum="Chordata" rank="species" species="onamata">
<emphasis box="[1060,1291,1701,1722]" italics="true" pageId="21" pageNumber="1658">Kekenodon onamata</emphasis>
</taxonomicName>
.
</paragraph>
<caption pageId="21" pageNumber="1658" startId="21.[161,225,866,887]" targetBox="[161,776,1041,1420]" targetIsTable="true" targetPageId="21">
<paragraph blockId="21.[161,746,866,1005]" pageId="21" pageNumber="1658">
<emphasis bold="true" box="[161,251,866,887]" pageId="21" pageNumber="1658">Table 5.</emphasis>
Measurements (in mm) of the holotype atlas of
<taxonomicName authorityName=", Hector" authorityYear="1881" box="[187,399,895,917]" class="Mammalia" family="Basilosauridae" genus="Kekenodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="21" pageNumber="1658" phylum="Chordata" rank="species" species="onamata">
<emphasis box="[187,399,895,917]" italics="true" pageId="21" pageNumber="1658">Kekenodon onamata</emphasis>
</taxonomicName>
(
<emphasis box="[413,571,895,917]" italics="true" pageId="21" pageNumber="1658">NMNZ Ma 306</emphasis>
). Measurements taken to the nearest tenth of a millimetre. Asterisk (*) denotes incomplete measurements as preserved; (e) denotes estimated measurements
</paragraph>
</caption>
<paragraph pageId="21" pageNumber="1658">
<table box="[161,776,1041,1420]" gridcols="2" gridrows="13" pageId="21" pageNumber="1658">
<tr box="[161,776,1041,1063]" gridrow="0" pageId="21" pageNumber="1658">
<th box="[161,615,1041,1063]" gridcol="0" gridrow="0" pageId="21" pageNumber="1658">Maximum dorsoventral depth</th>
<th box="[687,776,1041,1063]" gridcol="1" gridrow="0" pageId="21" pageNumber="1658">114.5*</th>
</tr>
<tr box="[161,776,1071,1093]" gridrow="1" pageId="21" pageNumber="1658">
<td box="[161,615,1071,1093]" gridcol="0" gridrow="1" pageId="21" pageNumber="1658">Maximum transverse width</td>
<td box="[687,776,1071,1093]" gridcol="1" gridrow="1" pageId="21" pageNumber="1658">181.5*</td>
</tr>
<tr box="[161,776,1101,1123]" gridrow="2" pageId="21" pageNumber="1658">
<td box="[161,615,1101,1123]" gridcol="0" gridrow="2" pageId="21" pageNumber="1658">Anteroposterior length of body</td>
<td box="[687,776,1101,1123]" gridcol="1" gridrow="2" pageId="21" pageNumber="1658">57*</td>
</tr>
<tr box="[161,776,1131,1153]" gridrow="3" pageId="21" pageNumber="1658">
<td box="[161,615,1131,1153]" gridcol="0" gridrow="3" pageId="21" pageNumber="1658">Dorsoventral depth of anterior articular</td>
<td box="[687,776,1131,1153]" gridcol="1" gridrow="3" pageId="21" pageNumber="1658">89.3*</td>
</tr>
<tr box="[161,776,1160,1182]" gridrow="4" pageId="21" pageNumber="1658" rowspan-1="1">
<td box="[161,615,1160,1182]" gridcol="0" gridrow="4" pageId="21" pageNumber="1658">surfaces</td>
</tr>
<tr box="[161,776,1190,1212]" gridrow="5" pageId="21" pageNumber="1658">
<td box="[161,615,1190,1212]" gridcol="0" gridrow="5" pageId="21" pageNumber="1658">Transverse width across anterior articular</td>
<td box="[687,776,1190,1212]" gridcol="1" gridrow="5" pageId="21" pageNumber="1658">152.1*</td>
</tr>
<tr box="[161,776,1220,1242]" gridrow="6" pageId="21" pageNumber="1658" rowspan-1="1">
<td box="[161,615,1220,1242]" gridcol="0" gridrow="6" pageId="21" pageNumber="1658">surfaces</td>
</tr>
<tr box="[161,776,1250,1272]" gridrow="7" pageId="21" pageNumber="1658">
<td box="[161,615,1250,1272]" gridcol="0" gridrow="7" pageId="21" pageNumber="1658">Dorsoventral depth of posterior articular</td>
<td box="[687,776,1250,1272]" gridcol="1" gridrow="7" pageId="21" pageNumber="1658">78</td>
</tr>
<tr box="[161,776,1279,1301]" gridrow="8" pageId="21" pageNumber="1658" rowspan-1="1">
<td box="[161,615,1279,1301]" gridcol="0" gridrow="8" pageId="21" pageNumber="1658">surfaces</td>
</tr>
<tr box="[161,776,1309,1331]" gridrow="9" pageId="21" pageNumber="1658">
<td box="[161,615,1309,1331]" gridcol="0" gridrow="9" pageId="21" pageNumber="1658">Transverse width across posterior articular</td>
<td box="[687,776,1309,1331]" gridcol="1" gridrow="9" pageId="21" pageNumber="1658">164</td>
</tr>
<tr box="[161,776,1338,1360]" gridrow="10" pageId="21" pageNumber="1658" rowspan-1="1">
<td box="[161,615,1338,1360]" gridcol="0" gridrow="10" pageId="21" pageNumber="1658">surfaces</td>
</tr>
<tr box="[161,776,1368,1390]" gridrow="11" pageId="21" pageNumber="1658">
<td box="[161,615,1368,1390]" gridcol="0" gridrow="11" pageId="21" pageNumber="1658">Dorsoventral depth of neural canal</td>
<td box="[687,776,1368,1390]" gridcol="1" gridrow="11" pageId="21" pageNumber="1658">59.8 (e)</td>
</tr>
<tr box="[161,776,1398,1420]" gridrow="12" pageId="21" pageNumber="1658">
<td box="[161,615,1398,1420]" gridcol="0" gridrow="12" pageId="21" pageNumber="1658">Transverse width of neural canal</td>
<td box="[687,776,1398,1420]" gridcol="1" gridrow="12" pageId="21" pageNumber="1658">67</td>
</tr>
</table>
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7381134" ID-Zenodo-Dep="7381134" httpUri="https://zenodo.org/record/7381134/files/figure.png" pageId="21" pageNumber="1658" startId="21.[160,238,1803,1825]" targetBox="[162,785,1435,1763]" targetPageId="21">
<paragraph blockId="21.[160,778,1803,1913]" pageId="21" pageNumber="1658">
<emphasis bold="true" box="[160,277,1803,1825]" pageId="21" pageNumber="1658">Figure 10.</emphasis>
Partial crown of a possible left p2 of
<taxonomicName authorityName="Beds (McKay" authorityYear="1882" box="[664,778,1803,1825]" class="Mammalia" family="Basilosauridae" genus="Kekenodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="21" pageNumber="1658" phylum="Chordata" rank="genus">
<emphasis box="[664,778,1803,1825]" italics="true" pageId="21" pageNumber="1658">Kekenodon</emphasis>
</taxonomicName>
cf.
<taxonomicName authority="(AUGD 2469)" baseAuthorityName="AUGD" baseAuthorityYear="2469" box="[191,465,1833,1855]" class="Mammalia" family="Basilosauridae" genus="Kekenodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="21" pageNumber="1658" phylum="Chordata" rank="species" species="onamata">
<emphasis box="[191,208,1833,1854]" italics="true" pageId="21" pageNumber="1658">K</emphasis>
.
<emphasis box="[221,315,1833,1854]" italics="true" pageId="21" pageNumber="1658">onamata</emphasis>
(AUGD 2469)
</taxonomicName>
in presumed labial (left) and lingual (right) view. Specimen whitened with ammonium chloride.
</paragraph>
</caption>
<paragraph blockId="21.[1050,1219,1791,1816]" box="[1050,1219,1791,1816]" pageId="21" pageNumber="1658">PHYLOGENY</paragraph>
<paragraph blockId="21.[826,1443,1832,1915]" lastBlockId="22.[145,762,197,802]" lastPageId="22" lastPageNumber="1659" pageId="21" pageNumber="1658">
The cladistic analysis under equal weighting recovered a single most-parsimonious tree (consistency index [CI]: 0.435; retention index [RI]: 0.654); tree length 786 steps;
<figureCitation box="[218,314,197,219]" captionStart="Figure 11" captionStartId="22.[146,226,1790,1812]" captionTargetBox="[146,1422,859,1749]" captionTargetId="figure-341@22.[145,1423,858,1750]" captionTargetPageId="22" captionText="Figure 11. Phylogenetic relationships of Kekenodon onamata. A, single most-parsimonious tree recovered under equal weighting. B, strict consensus of three equally most-parsimonious trees recovered under implied weighting (K = 3). OU 22100 is an undescribed basilosaurid from New Zealand designated Zygorhiza sp. Branch support shown as GC frequency values." figureDoi="http://doi.org/10.5281/zenodo.7381136" httpUri="https://zenodo.org/record/7381136/files/figure.png" pageId="22" pageNumber="1659">Fig. 11A</figureCitation>
) and three equally most-parsimonious trees under implied weighting (CI: 0.435; RI: 0.653; tree length 72 steps;
<figureCitation box="[381,475,258,280]" captionStart="Figure 11" captionStartId="22.[146,226,1790,1812]" captionTargetBox="[146,1422,859,1749]" captionTargetId="figure-341@22.[145,1423,858,1750]" captionTargetPageId="22" captionText="Figure 11. Phylogenetic relationships of Kekenodon onamata. A, single most-parsimonious tree recovered under equal weighting. B, strict consensus of three equally most-parsimonious trees recovered under implied weighting (K = 3). OU 22100 is an undescribed basilosaurid from New Zealand designated Zygorhiza sp. Branch support shown as GC frequency values." figureDoi="http://doi.org/10.5281/zenodo.7381136" httpUri="https://zenodo.org/record/7381136/files/figure.png" pageId="22" pageNumber="1659">Fig. 11B</figureCitation>
). The affinities of Eocene archaeocetes and within the Neoceti (excluding taxa previously identified as non-aetiocetid toothed mysticetes; see below) are well resolved, probably owing to the comprehensive nature of the taxa and morphological characters included in the cladistic data matrix to prevent bias toward a particular group or clade in determining the taxonomic affinities of
<taxonomicName baseAuthorityName="AUGD" baseAuthorityYear="2469" box="[145,375,504,525]" class="Mammalia" family="Basilosauridae" genus="Kekenodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="22" pageNumber="1659" phylum="Chordata" rank="species" species="onamata">
<emphasis box="[145,375,504,525]" italics="true" pageId="22" pageNumber="1659">Kekenodon onamata</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph blockId="22.[145,762,197,802]" lastBlockId="23.[163,780,197,1905]" lastPageId="23" lastPageNumber="1660" pageId="22" pageNumber="1659">
There are few topological differences between the equal and implied weighting analyses, namely occurring in Eocene archaeocetes and non-aetiocetid toothed mysticetes (
<figureCitation box="[381,461,626,648]" captionStart="Figure 11" captionStartId="22.[146,226,1790,1812]" captionTargetBox="[146,1422,859,1749]" captionTargetId="figure-341@22.[145,1423,858,1750]" captionTargetPageId="22" captionText="Figure 11. Phylogenetic relationships of Kekenodon onamata. A, single most-parsimonious tree recovered under equal weighting. B, strict consensus of three equally most-parsimonious trees recovered under implied weighting (K = 3). OU 22100 is an undescribed basilosaurid from New Zealand designated Zygorhiza sp. Branch support shown as GC frequency values." figureDoi="http://doi.org/10.5281/zenodo.7381136" httpUri="https://zenodo.org/record/7381136/files/figure.png" pageId="22" pageNumber="1659">Fig. 11</figureCitation>
). The affinities of Eocene archaeocetes are better resolved under equal weighting with a paraphyletic
<taxonomicName box="[379,524,688,710]" class="Mammalia" family="Protocetidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="22" pageNumber="1659" phylum="Chordata" rank="family">Protocetidae</taxonomicName>
and
<taxonomicName box="[582,756,688,710]" class="Mammalia" family="Basilosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="22" pageNumber="1659" phylum="Chordata" rank="family">Basilosauridae</taxonomicName>
, whereas dorudontine basilosaurids form a polytomy under implied weighting. A
<taxonomicName authorityName="Harlan" authorityYear="1834" box="[516,683,749,770]" class="Mammalia" family="Basilosauridae" genus="Basilosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="22" pageNumber="1659" phylum="Chordata" rank="genus">
<emphasis box="[516,683,749,770]" italics="true" pageId="22" pageNumber="1659">Basilosaurus</emphasis>
</taxonomicName>
clade composed of
<taxonomicName authorityName="Andrews" authorityYear="1904" box="[281,471,780,801]" class="Mammalia" family="Basilosauridae" genus="Basilosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="22" pageNumber="1659" phylum="Chordata" rank="species" species="isis">
<emphasis box="[281,471,780,801]" italics="true" pageId="22" pageNumber="1659">Basilosaurus isis</emphasis>
</taxonomicName>
and
<taxonomicName authority="Owen, 1839" authorityName="Owen" authorityYear="1839" box="[521,947,197,801]" class="Mammalia" family="Basilosauridae" genus="Basilosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="22" pageNumber="1659" phylum="Chordata" rank="species" species="cetoides">
<emphasis box="[521,761,780,801]" italics="true" pageId="22" pageNumber="1659">Basilosaurus cetoides</emphasis>
<bibRefCitation author="Owen R" box="[809,947,197,218]" pageId="22" pageNumber="1659" pagination="69 - 79" refId="ref26078" refString="Owen R. 1839. Observations on the Basilosaurus of Dr. Harlan (Zeuglodon cetoides, Owen). Transactions of the Geological Society of London 6: 69 - 79." type="journal article" year="1839">Owen, 1839</bibRefCitation>
</taxonomicName>
is formed under both equal and implied analyses with strong support (branch support = 85% and 87%, respectively) in addition to a poorly supported
<taxonomicName authority="Uhen, 2005" authorityName="Uhen" authorityYear="2005" box="[809,1256,289,311]" class="Mammalia" family="Basilosauridae" genus="Cynthiacetus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="22" pageNumber="1659" phylum="Chordata" rank="species" species="peruvianus">
<emphasis box="[809,1108,289,311]" italics="true" pageId="22" pageNumber="1659">Cynthiacetus peruvianus</emphasis>
<bibRefCitation author="Uhen MD" box="[1118,1256,289,311]" pageId="22" pageNumber="1659" pagination="157 - 172" refId="ref26803" refString="Uhen MD. 2005. A new genus and species of archaeocete whale from Mississippi. Southeastern Geology 43: 157 - 172." type="journal article" year="2005">Uhen, 2005</bibRefCitation>
</taxonomicName>
and
<taxonomicName baseAuthorityName="Uhen" baseAuthorityYear="2004" class="Mammalia" family="Basilosauridae" genus="Dorudon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="22" pageNumber="1659" phylum="Chordata" rank="species" species="atrox">
<emphasis italics="true" pageId="22" pageNumber="1659">Dorudon atrox</emphasis>
</taxonomicName>
clade. Neoceti monophyly is recovered under both analyses with identical topology, but with poor support. Odontocete monophyly is recovered under both equal and implied weighting with moderate support (branch support = 56% and 82%, respectively). A
<taxonomicName box="[834,994,473,495]" class="Mammalia" family="Xenorophidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="22" pageNumber="1659" phylum="Chordata" rank="family">Xenorophidae</taxonomicName>
clade is recovered with poor support under equal weighting and moderate support under implied weighting (branch support = 67%). Within
<taxonomicName box="[809,972,565,587]" class="Mammalia" family="Xenorophidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="22" pageNumber="1659" phylum="Chordata" rank="family">Xenorophidae</taxonomicName>
,
<taxonomicName authority="Uhen, 2008" authorityName="Uhen" authorityYear="2008" box="[983,1424,565,587]" class="Mammalia" family="Xenorophidae" genus="Albertocetus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="22" pageNumber="1659" phylum="Chordata" rank="species" species="meffordorum">
<emphasis box="[983,1284,565,586]" italics="true" pageId="22" pageNumber="1659">Albertocetus meffordorum</emphasis>
Uhen, 2008
</taxonomicName>
and
<taxonomicName authority="Kellogg, 1923" authorityName="Kellogg" authorityYear="1923" box="[860,1245,596,618]" class="Mammalia" family="Xenorophidae" genus="Xenorophus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="22" pageNumber="1659" phylum="Chordata" rank="species" species="sloanii">
<emphasis box="[860,1081,596,617]" italics="true" pageId="22" pageNumber="1659">Xenorophus sloanii</emphasis>
<bibRefCitation author="Kellogg AR" box="[1089,1245,596,618]" pageId="22" pageNumber="1659" pagination="1 - 69" refId="ref24870" refString="Kellogg AR. 1923. Description of two squalodonts recently discovered in the Calvert Cliffs, Maryland; and notes on the shark-toothed dolphins. Proceedings of the United States National Museum 62: 1 - 69." type="journal article" year="1923">Kellogg, 1923</bibRefCitation>
</taxonomicName>
form a clade to the exclusion of stemward-positioned
<taxonomicName authority="Allen, 1921" authorityName="Allen" authorityYear="1921" class="Mammalia" family="Xenorophidae" genus="Archaeodelphis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="22" pageNumber="1659" phylum="Chordata" rank="species" species="patrius">
<emphasis italics="true" pageId="22" pageNumber="1659">Archaeodelphis patrius</emphasis>
Allen, 1921
</taxonomicName>
with moderate support under e q u a l w e i g h t i n g (b r a n c h s u p p o r t = 8 4%) a n d strong support under implied weighting (branch support = 91%). A second clade consists of the remaining odontocete taxa from this analysis, which includes
<taxonomicName authority="Dubrovo &amp; Sanders 2000" authorityName="Dubrovo &amp; Sanders" authorityYear="2000" class="Mammalia" family="Patriocetidae" genus="Patriocetus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="kazakhstanicus">
<emphasis box="[280,629,197,219]" italics="true" pageId="23" pageNumber="1660">Patriocetus kazakhstanicus</emphasis>
<bibRefCitation author="Dubrovo IA &amp; Sanders AE" pageId="23" pageNumber="1660" pagination="577 - 590" refId="ref23095" refString="Dubrovo IA, Sanders AE. 2000. A new species of Patriocetus (Mammalia, Cetacea) from the late Oligocene of Kazakhstan. Journal of Vertebrate Paleontology 20: 577 - 590." type="journal article" year="2000">Dubrovo &amp; Sanders 2000</bibRefCitation>
</taxonomicName>
,
<taxonomicName authority="(J. P. Muller, 1849)" baseAuthorityName="J. P. Muller" baseAuthorityYear="1849" box="[326,775,228,250]" class="Mammalia" family="Agorophiidae" genus="Agorophius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="pygmaeus">
<emphasis box="[326,573,228,250]" italics="true" pageId="23" pageNumber="1660">Agorophius pygmaeus</emphasis>
(J.P.Muller, 1849)
</taxonomicName>
,
<taxonomicName authority="Fordyce, 2002" authorityName="Fordyce" authorityYear="2002" box="[163,498,258,280]" class="Mammalia" family="Simocetidae" genus="Simocetus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="rayi">
<emphasis box="[163,332,259,280]" italics="true" pageId="23" pageNumber="1660">Simocetus rayi</emphasis>
Fordyce, 2002
</taxonomicName>
,
<taxonomicName authority="Fordyce, 1994" authorityName="Fordyce" authorityYear="1994" class="Mammalia" family="Waipatiidae" genus="Waipatia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="maerewhenua">
<emphasis box="[509,778,258,280]" italics="true" pageId="23" pageNumber="1660">Waipatia maerewhenua</emphasis>
<bibRefCitation author="Fordyce RE" box="[163,323,289,311]" pageId="23" pageNumber="1660" pagination="147 - 176" refId="ref23644" refString="Fordyce RE. 1994. Waipatia maerewhenua, a new genus and new species (Waipatiidae, new Family), an archaic Late Oligocene dolphin (Cetacea: Odontoceti: Platanistoidea) from New Zealand. In: Berta A, Demere TA, eds. Contributions in marine mammal paleontology honoring Frank C. Whitmore, Jr. Proceedings of the San Diego Society of Natural History 29: 147 - 176." type="journal article" year="1994">Fordyce, 1994</bibRefCitation>
</taxonomicName>
,
<taxonomicName authority="Kellogg, 1923" authorityName="Kellogg" authorityYear="1923" box="[338,773,289,311]" class="Mammalia" family="Squalodontidae" genus="Squalodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="calvertensis">
<emphasis box="[338,607,289,310]" italics="true" pageId="23" pageNumber="1660">Squalodon calvertensis</emphasis>
<bibRefCitation author="Kellogg AR" box="[615,773,289,311]" pageId="23" pageNumber="1660" pagination="1 - 69" refId="ref24870" refString="Kellogg AR. 1923. Description of two squalodonts recently discovered in the Calvert Cliffs, Maryland; and notes on the shark-toothed dolphins. Proceedings of the United States National Museum 62: 1 - 69." type="journal article" year="1923">Kellogg, 1923</bibRefCitation>
</taxonomicName>
,
<taxonomicName authority="(Gervais, 1855)" baseAuthorityName="Gervais" baseAuthorityYear="1855" box="[163,618,320,342]" class="Mammalia" family="Hyperoodontidae" genus="Mesoplodon" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="europaeus">
<emphasis box="[163,429,320,341]" italics="true" pageId="23" pageNumber="1660">Mesoplodon europaeus</emphasis>
(Gervais, 1855)
</taxonomicName>
and
<taxonomicName authorityName="Linnaeus" authorityYear="1758" class="Mammalia" family="Physeteridae" genus="Physeter" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="macrocephalus">
<emphasis italics="true" pageId="23" pageNumber="1660">Physeter macrocephalus</emphasis>
</taxonomicName>
with moderate support under equal and implied weighting (branch support = 53% and 78%, respectively). A separate clade composed of
<taxonomicName authorityName="Dubrovo &amp; Sanders" authorityYear="2000" class="Mammalia" family="Patriocetidae" genus="Patriocetus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="kazakhstanicus">
<emphasis box="[756,772,412,433]" italics="true" pageId="23" pageNumber="1660">P</emphasis>
.
<emphasis box="[163,339,442,463]" italics="true" pageId="23" pageNumber="1660">kazakhstanicus</emphasis>
</taxonomicName>
and
<taxonomicName baseAuthorityName="J. P. Muller" baseAuthorityYear="1849" box="[392,534,443,464]" class="Mammalia" family="Agorophiidae" genus="Agorophius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="pygmaeus">
<emphasis box="[392,409,443,464]" italics="true" pageId="23" pageNumber="1660">A</emphasis>
.
<emphasis box="[421,534,443,464]" italics="true" pageId="23" pageNumber="1660">pygmaeus</emphasis>
</taxonomicName>
is recovered, but with poor support. Crownward, the two extant odontocete taxa included in this analysis (
<taxonomicName baseAuthorityName="Gervais" baseAuthorityYear="1855" box="[536,694,504,525]" class="Mammalia" family="Hyperoodontidae" genus="Mesoplodon" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="europaeus">
<emphasis box="[536,559,504,525]" italics="true" pageId="23" pageNumber="1660">M</emphasis>
.
<emphasis box="[575,694,504,525]" italics="true" pageId="23" pageNumber="1660">europaeus</emphasis>
</taxonomicName>
and
<taxonomicName authority="Linnaeus, 1758" authorityName="Linnaeus" authorityYear="1758" class="Mammalia" family="Physeteridae" genus="Physeter" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="macrocephalus">
<emphasis box="[756,772,504,525]" italics="true" pageId="23" pageNumber="1660">P</emphasis>
.
<emphasis box="[163,344,534,555]" italics="true" pageId="23" pageNumber="1660">macrocephalus</emphasis>
<bibRefCitation author="Linnaeus C" box="[354,542,534,556]" pageId="23" pageNumber="1660" refId="ref25253" refString="Linnaeus C. 1758. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Impensis Direct. Stockholm: L. Salvius." type="book" year="1758">Linnaeus, 1758</bibRefCitation>
</taxonomicName>
) form a clade with strong support (branch support = 99% under both analyses). Mysticete monophyly (excluding previously recognized non-aetiocetid toothed mysticetes) is recovered with poor support under equal weighting and moderate support under implied weighting (branch support = 60%).
<taxonomicName authorityName="Emlong" authorityYear="1966" box="[476,631,718,740]" class="Mammalia" family="Aetiocetidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="family">Aetiocetidae</taxonomicName>
monophyly is recovered, but with poor support. Within the
<taxonomicName authorityName="Emlong" authorityYear="1966" box="[163,317,780,802]" class="Mammalia" family="Aetiocetidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="family">Aetiocetidae</taxonomicName>
, a clade composed of members of the genus
<taxonomicName authorityName="Emlong" authorityYear="1966" box="[246,371,811,832]" class="Mammalia" family="Aetiocetidae" genus="Aetiocetus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="genus">
<emphasis box="[246,371,811,832]" italics="true" pageId="23" pageNumber="1660">Aetiocetus</emphasis>
</taxonomicName>
(
<taxonomicName authority="Emlong, 1966" authorityName="Emlong" authorityYear="1966" class="Mammalia" family="Aetiocetidae" genus="Aetiocetus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="cotylalveus">
<emphasis box="[392,666,810,832]" italics="true" pageId="23" pageNumber="1660">Aetiocetus cotylalveus</emphasis>
<bibRefCitation author="Emlong D" pageId="23" pageNumber="1660" pagination="1 - 51" refId="ref23130" refString="Emlong D. 1966. A new archaic cetacean from the Oligocene of Northwest Oregon. Bulletin of the Museum of Natural History, University of Oregon 3: 1 - 51." type="journal article" year="1966">Emlong, 1966</bibRefCitation>
</taxonomicName>
,
<taxonomicName box="[233,508,841,862]" class="Mammalia" family="Aetiocetidae" genus="Aetiocetus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="polydentatus">
<emphasis box="[233,508,841,862]" italics="true" pageId="23" pageNumber="1660">Aetiocetus polydentatus</emphasis>
</taxonomicName>
and
<taxonomicName authority="Barnes et al., 1995" authorityName="Barnes" authorityYear="1995" class="Mammalia" family="Aetiocetidae" genus="Aetiocetus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="weltoni">
<emphasis box="[568,779,841,862]" italics="true" pageId="23" pageNumber="1660">Aetiocetus weltoni</emphasis>
<bibRefCitation author="Barnes LG &amp; Kimura M &amp; Furusawa H &amp; Sawamura H" box="[163,399,872,894]" pageId="23" pageNumber="1660" pagination="392 - 431" refId="ref22258" refString="Barnes LG, Kimura M, Furusawa H, Sawamura H. 1995. Classification and distribution of Oligocene Aetiocetidae (Mammalia; Cetacea; Mysticeti) from western North America and Japan. The Island Arc 3: 392 - 431." type="journal article" year="1995">
Barnes
<emphasis box="[260,323,872,893]" italics="true" pageId="23" pageNumber="1660">et al.</emphasis>
, 1995
</bibRefCitation>
</taxonomicName>
) is recovered to the exclusion of
<taxonomicName box="[195,445,902,924]" class="Mammalia" family="Aetiocetidae" genus="Fucaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="goedertorum">
<emphasis box="[195,445,902,924]" italics="true" pageId="23" pageNumber="1660">Fucaia goedertorum</emphasis>
</taxonomicName>
and
<emphasis box="[513,779,902,923]" italics="true" pageId="23" pageNumber="1660">Chonocetus sookensis</emphasis>
Russell, 1968 with moderate support under equal weighting (branch support = 54%) and poor support under implied weighting. A clade including
<taxonomicName box="[665,778,994,1016]" class="Mammalia" family="Aetiocetidae" genus="Aetiocetus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="weltoni">
<emphasis box="[665,682,995,1016]" italics="true" pageId="23" pageNumber="1660">A</emphasis>
.
<emphasis box="[695,778,994,1015]" italics="true" pageId="23" pageNumber="1660">weltoni</emphasis>
</taxonomicName>
and
<taxonomicName box="[217,407,1025,1046]" class="Mammalia" family="Aetiocetidae" genus="Aetiocetus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="polydentatus">
<emphasis box="[217,234,1025,1046]" italics="true" pageId="23" pageNumber="1660">A</emphasis>
.
<emphasis box="[252,407,1025,1046]" italics="true" pageId="23" pageNumber="1660">polydentatus</emphasis>
</taxonomicName>
is recovered in the
<taxonomicName authorityName="Emlong" authorityYear="1966" box="[657,779,1025,1046]" class="Mammalia" family="Aetiocetidae" genus="Aetiocetus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="genus">
<emphasis box="[657,779,1025,1046]" italics="true" pageId="23" pageNumber="1660">Aetiocetus</emphasis>
</taxonomicName>
clade with moderate support under both equal and implied weighting (branch support = 55% and 64%, respectively).
<taxonomicName box="[334,578,1117,1139]" class="Mammalia" family="Aetiocetidae" genus="Fucaia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="goedertorum">
<emphasis box="[334,578,1117,1139]" italics="true" pageId="23" pageNumber="1660">Fucaia goedertorum</emphasis>
</taxonomicName>
and
<emphasis italics="true" pageId="23" pageNumber="1660">Chonocetus sookensis</emphasis>
form a clade separate to
<taxonomicName authorityName="Emlong" authorityYear="1966" box="[551,665,1148,1169]" class="Mammalia" family="Aetiocetidae" genus="Aetiocetus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="genus">
<emphasis box="[551,665,1148,1169]" italics="true" pageId="23" pageNumber="1660">Aetiocetus</emphasis>
</taxonomicName>
with poor support under equal weighting and moderate support under implied weighting (branch support = 68%). A Chaeomysticeti clade is recovered with moderate support under equal and implied weighting (branch support = 78% and 50%, respectively), which also includes a clade of crown Mysticeti with strong support (branch support = 99% under both analyses). A complete list of synapomorphies for all clades is included in the Supporting Information (Appendix S1).
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7381136" ID-Zenodo-Dep="7381136" httpUri="https://zenodo.org/record/7381136/files/figure.png" pageId="22" pageNumber="1659" startId="22.[146,226,1790,1812]" targetBox="[146,1422,859,1749]" targetPageId="22">
<paragraph blockId="22.[145,1423,1790,1900]" pageId="22" pageNumber="1659">
<emphasis bold="true" box="[146,265,1790,1812]" pageId="22" pageNumber="1659">Figure 11.</emphasis>
Phylogenetic relationships of
<taxonomicName baseAuthorityName="AUGD" baseAuthorityYear="2469" box="[594,811,1790,1812]" class="Mammalia" family="Basilosauridae" genus="Kekenodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="22" pageNumber="1659" phylum="Chordata" rank="species" species="onamata">
<emphasis box="[594,811,1790,1812]" italics="true" pageId="22" pageNumber="1659">Kekenodon onamata</emphasis>
</taxonomicName>
. A, single most-parsimonious tree recovered under equal weighting. B, strict consensus of three equally most-parsimonious trees recovered under implied weighting (
<emphasis box="[1309,1326,1820,1841]" italics="true" pageId="22" pageNumber="1659">K</emphasis>
= 3). OU 22100 is an undescribed basilosaurid from New Zealand designated
<taxonomicName box="[860,1000,1849,1871]" class="Mammalia" family="Basilosauridae" genus="Zygorhiza" kingdom="Animalia" order="Cetacea" pageId="22" pageNumber="1659" phylum="Chordata" rank="species" species="undetermined">
<emphasis box="[860,965,1849,1871]" italics="true" pageId="22" pageNumber="1659">Zygorhiza</emphasis>
sp.
</taxonomicName>
Branch support shown as GC frequency values.
</paragraph>
</caption>
<paragraph blockId="23.[163,780,197,1905]" lastBlockId="23.[827,1444,197,1905]" pageId="23" pageNumber="1660">
A novel position for taxa previously recognized as non-aetiocetid toothed mysticetes as sister to the Neoceti is recovered under both equal and implied weighting with poor support. Among these taxa, a clade composed of
<taxonomicName authorityName="Mitchell" authorityYear="1989" box="[318,620,1577,1598]" class="Mammalia" family="Llanocetidae" genus="Llanocetus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="denticrenatus">
<emphasis box="[318,620,1577,1598]" italics="true" pageId="23" pageNumber="1660">Llanocetus denticrenatus</emphasis>
</taxonomicName>
,
<taxonomicName class="Mammalia" family="Mystacodontidae" genus="Mystacodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="selenesis">
<emphasis italics="true" pageId="23" pageNumber="1660">Mystacodon selenesis</emphasis>
</taxonomicName>
,
<taxonomicName box="[278,547,1608,1629]" class="Mammalia" genus="Coronodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="havensteini">
<emphasis box="[278,547,1608,1629]" italics="true" pageId="23" pageNumber="1660">Coronodon havensteini</emphasis>
</taxonomicName>
, ChM PV 4745 and ChM PV 5720 is recovered under equal weighting, but with poor support. Under implied weighting, these taxa are paraphyletic with a
<taxonomicName authorityName="Mitchell" authorityYear="1989" box="[586,779,1700,1721]" class="Mammalia" family="Llanocetidae" genus="Llanocetus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="denticrenatus">
<emphasis box="[586,602,1700,1721]" italics="true" pageId="23" pageNumber="1660">L</emphasis>
.
<emphasis box="[618,779,1700,1721]" italics="true" pageId="23" pageNumber="1660">denticrenatus</emphasis>
</taxonomicName>
and
<taxonomicName box="[220,371,1730,1752]" class="Mammalia" family="Mystacodontidae" genus="Mystacodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="selenesis">
<emphasis box="[220,243,1731,1752]" italics="true" pageId="23" pageNumber="1660">M</emphasis>
.
<emphasis box="[263,371,1730,1751]" italics="true" pageId="23" pageNumber="1660">selenesis</emphasis>
</taxonomicName>
clade representing the earliest diverging lineage, but with poor support. Identifying either clade as the
<taxonomicName authority="(Mitchell, 1989)" baseAuthorityName="Mitchell" baseAuthorityYear="1989" box="[397,749,1791,1813]" class="Mammalia" family="Llanocetidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="family">
Llanocetidae (
<bibRefCitation author="Mitchell ED" box="[569,740,1791,1813]" pageId="23" pageNumber="1660" pagination="2219 - 2235" refId="ref25867" refString="Mitchell ED. 1989. A new cetacean from the Late Eocene La Meseta Formation, Seymour Island, Antarctic Peninsula. Canadian Journal of Fisheries and Aquatic Sciences 46: 2219 - 2235." type="journal article" year="1989">Mitchell, 1989</bibRefCitation>
)
</taxonomicName>
is speculative given the differences in topology and poor support across both analyses. A clade comprised of
<taxonomicName box="[163,334,1883,1905]" class="Mammalia" genus="Coronodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="havensteini">
<emphasis box="[163,181,1884,1905]" italics="true" pageId="23" pageNumber="1660">C</emphasis>
.
<emphasis box="[197,334,1883,1904]" italics="true" pageId="23" pageNumber="1660">havensteini</emphasis>
</taxonomicName>
, ChM PV 4745 and ChM PV 5720 is recovered under both analyses with moderate support under implied weighting (branch support = 79%). A single synapomorphy common to both analyses supports this clade: tooth enamel smooth (character 34: 1). Specimens ChM PV 4745 and ChM PV 5720 form a clade under both equal and implied weighting with moderate support (branch support = 86% and 83%, respectively). Six synapomorphies common to both analyses supports this clade: upper-posterior cheek teeth are double-rooted with roots closely appressed for the entire length (character 23: 1); number of upper molars equalling three (character 26: 1); anteriormost point of the supraoccipital in line with the gap between the anterior edge of the zygomatic process of squamosal and the anteriormost point along the posterior edge of the supraorbital process of frontal (character 123: 2); apex of the anterior process of the periotic bears a tubercle (character 159: 1); anteroexternal sulcus absent (character 164: 1); and involucrum with groove or grooves absent (character 216: 0). A
<taxonomicName box="[1231,1443,779,801]" class="Mammalia" family="Mammalodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="family">Mammalodontidae</taxonomicName>
clade (
<taxonomicName authorityName="Fitzgerald" authorityYear="2006" box="[901,1120,810,832]" class="Mammalia" family="Mammalodontidae" genus="Janjucetus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="hunderi">
<emphasis box="[901,1120,810,832]" italics="true" pageId="23" pageNumber="1660">Janjucetus hunderi</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Pritchard" authorityYear="1939" box="[1178,1434,810,831]" class="Mammalia" family="Squalodontidae" genus="Mammalodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="colliveri">
<emphasis box="[1178,1434,810,831]" italics="true" pageId="23" pageNumber="1660">Mammalodon colliveri</emphasis>
</taxonomicName>
) positioned as the most crownward is recovered with moderate support under equal weighting (branch support = 58%) and strong support under implied weighing (branch support = 88%). Six synapomorphies common to both analyses support this clade: percentage of skull length relative to bicondylar breadth less than 700% (character 1: 0); preorbital process of frontal has a rounded off, anteriorly convex linguiform outline (character 56: 1); posteromedial edge of the ascending process of the maxilla is separated from the posterolateral edge of the nasal or premaxilla by a triangular wedge of frontal (character 84: 1); suture between right and left nasals and right and left frontals shifted toward the left side (character 98: 1); frontal shield breadth taken at mid-orbit relative to condylar breadth less than two (character 110: 0); and frontoparietal suture with skull in dorsal view, the suture is strongly V- or U-shaped with the vertex of the V or U pointed posteriorly such that the posteriormost edge of the frontal is posterior to the anteriormost edge of the parietal (character 119: 1).
</paragraph>
<paragraph blockId="23.[827,1444,197,1905]" lastBlockId="24.[145,762,197,1905]" lastPageId="24" lastPageNumber="1661" pageId="23" pageNumber="1660">
These results are interesting given the recent surge in studies on toothed mysticete phylogeny brought on by the discovery of new morphologically and taxonomically informative fossils (e.g.
<bibRefCitation author="Fitzgerald EMG" box="[1254,1437,1577,1599]" pageId="23" pageNumber="1660" pagination="2955 - 2963" refId="ref23347" refString="Fitzgerald EMG. 2006. A bizarre new toothed mysticete (Cetacea) from Australia and the early evolution of baleen whales. Proceedings of the Royal Society B: Biological Sciences 273: 2955 - 2963." type="journal article" year="2006">Fitzgerald, 2006</bibRefCitation>
; Marx
<emphasis box="[897,948,1607,1629]" italics="true" pageId="23" pageNumber="1660">et al</emphasis>
., 2015, 2016;
<bibRefCitation author="Geisler JH &amp; Boessenecker RW &amp; Brown M &amp; Beatty BL" box="[1109,1328,1607,1629]" pageId="23" pageNumber="1660" pagination="20361 - 2042" refId="ref24197" refString="Geisler JH, Boessenecker RW, Brown M, Beatty BL. 2017. The origin of filter feeding in whales. Current Biology 27: 20361 - 2042. e 2." type="journal article" year="2017">
Geisler
<emphasis box="[1200,1258,1607,1629]" italics="true" pageId="23" pageNumber="1660">et al.</emphasis>
, 2017
</bibRefCitation>
;
<bibRefCitation author="Lambert O &amp; Martinez-Caceres M &amp; Bianucci G &amp; Di Celma C &amp; Salas-Gismondi R &amp; Steurbaut E &amp; Urbina M &amp; de Muizon C" pageId="23" pageNumber="1660" pagination="1535 - 1541" refId="ref25109" refString="Lambert O, Martinez-Caceres M, Bianucci G, Di Celma C, Salas-Gismondi R, Steurbaut E, Urbina M, de Muizon C. 2017. Earliest mysticete from the late Eocene of Peru sheds new light on the origin of baleen whales. Current Biology 27: 1535 - 1541. e 2." type="journal article" year="2017">
Lambert
<emphasis box="[827,881,1638,1660]" italics="true" pageId="23" pageNumber="1660">et al.</emphasis>
, 2017
</bibRefCitation>
;
<bibRefCitation author="Fordyce RE &amp; Marx FG" box="[958,1206,1638,1660]" pageId="23" pageNumber="1660" pagination="1670 - 1676" refId="ref23930" refString="Fordyce RE, Marx FG. 2018. Gigantism precedes filter feeding in baleen whale evolution. Current Biology 28: 1670 - 1676. e 2." type="journal article" year="2018">Fordyce &amp; Marx, 2018</bibRefCitation>
;
<bibRefCitation author="de Muizon C &amp; Bianucci G &amp; Martinez-Caceres M &amp; Lambert O" box="[1217,1429,1638,1660]" pageId="23" pageNumber="1660" pagination="401 - 499" refId="ref26028" refString="de Muizon C, Bianucci G, Martinez-Caceres M, Lambert O. 2019. Mystacodon selenesis, the earliest known toothed mysticete (Cetacea, Mammalia) from the late Eocene of Peru, anatomy, phylogeny, and feeding adaptations. Geodiversitas 41: 401 - 499." type="journal article" year="2019">
Muizon
<emphasis box="[1308,1355,1638,1660]" italics="true" pageId="23" pageNumber="1660">et al</emphasis>
., 2019
</bibRefCitation>
), which, while having some topological differences, posit these taxa within Mysticeti. However, the position of these taxa in previous studies is seemingly vulnerable to change due to low branch support, namely when more archaeocete and odontocete taxa are included in phylogenetic analyses. Removing
<taxonomicName baseAuthorityName="AUGD" baseAuthorityYear="2469" box="[1211,1443,1822,1844]" class="Mammalia" family="Basilosauridae" genus="Kekenodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="onamata">
<emphasis box="[1211,1443,1822,1844]" italics="true" pageId="23" pageNumber="1660">Kekenodon onamata</emphasis>
</taxonomicName>
from the analyses yielded no differences in topology under equal weighting, while the
<taxonomicName authorityName="Mitchell" authorityYear="1989" box="[1245,1443,1883,1905]" class="Mammalia" family="Llanocetidae" genus="Llanocetus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="1660" phylum="Chordata" rank="species" species="denticrenatus">
<emphasis box="[1245,1261,1884,1905]" italics="true" pageId="23" pageNumber="1660">L</emphasis>
.
<emphasis box="[1278,1443,1883,1904]" italics="true" pageId="23" pageNumber="1660">denticrenatus</emphasis>
</taxonomicName>
and
<taxonomicName box="[199,343,197,219]" class="Mammalia" family="Mystacodontidae" genus="Mystacodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="24" pageNumber="1661" phylum="Chordata" rank="species" species="selenesis">
<emphasis box="[199,222,198,219]" italics="true" pageId="24" pageNumber="1661">M</emphasis>
.
<emphasis box="[239,343,197,218]" italics="true" pageId="24" pageNumber="1661">selenesis</emphasis>
</taxonomicName>
clade moved crownward to the
<taxonomicName class="Mammalia" genus="Coronodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="24" pageNumber="1661" phylum="Chordata" rank="species" species="havensteini">
<emphasis box="[736,754,198,219]" italics="true" pageId="24" pageNumber="1661">C</emphasis>
.
<emphasis box="[145,281,228,249]" italics="true" pageId="24" pageNumber="1661">havensteini</emphasis>
</taxonomicName>
, ChM PV 4745 and ChM PV 5720 clade under implied weighting. Critically, the position of these taxa as sister to Neoceti remained under equal and implied weighting with strong support (branch support = 98% and 99%, respectively). The results of analyses with
<taxonomicName baseAuthorityName="AUGD" baseAuthorityYear="2469" box="[355,496,382,403]" class="Mammalia" family="Basilosauridae" genus="Kekenodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="24" pageNumber="1661" phylum="Chordata" rank="species" species="onamata">
<emphasis box="[355,373,382,403]" italics="true" pageId="24" pageNumber="1661">K</emphasis>
.
<emphasis box="[390,496,382,403]" italics="true" pageId="24" pageNumber="1661">onamata</emphasis>
</taxonomicName>
removed can be seen in the Supporting Information (Appendix S1). Ten synapomorphies common to both analyses support mysticete monophyly: anterior-half of maxilla forms a highly acute angle with a flattened maxilla (character 4: 1); suture between maxilla and premaxilla on rostrum is unfused and marked by a deep groove (character 9: 3); nutrient foramina and sulci on palatal surface of maxilla present (character 19: 1); mandibular symphysis not sutured (character 47: 1); symphyseal surface smooth (character 48: 1); symphyseal groove present (character 49: 1); alveolar margin on the mandible is parallel to the ventral margin for its entire length (character 52: 1); lateral edges of nasals anterior to the level of the anterior edge of preorbital process of frontal diverge anteriorly (character 101: 2); inferior lamina of pterygoid present and floors most of sinus cavity (character 139: 1); and median furrow of tympanic bulla forms notch on posterior edge of bulla between medial and lateral prominences (character 212: 1). Given that nearly all of these characters relate to the feeding apparatus, the relatively more primitive rostral and mandibular structures of previously recognized non-aetiocetid toothed mysticetes may at least partially explain why these taxa are being posited in a more basal position outside of Neoceti.
</paragraph>
<paragraph blockId="24.[145,762,197,1905]" pageId="24" pageNumber="1661">
It is possible that excluding additional archaeocete and odontocete taxa may cause previously recognized non-aetiocetid toothed mysticetes to be posited in Mysticeti, which would suggest evidence of long-branch attraction. In most studies focused on the affinities of fossil Mysticeti, few archaeocete and odontocete taxa are included and are primarily relegated to the outgroup. Including additional archaeocete and odontocete taxa in previous studies on mysticete phylogeny may limit taxonomic bias and more accurately reflect the affinities of basal toothed mysticetes. The novel positioning of these taxa as sister to Neoceti requires future study to evaluate whether these results more accurately reflect the affinities of these archaic toothed cetaceans, including possible expansion of
<taxonomicName baseAuthorityName="Mitchell" baseAuthorityYear="1989" box="[521,673,1608,1630]" class="Mammalia" family="Llanocetidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="24" pageNumber="1661" phylum="Chordata" rank="family">Llanocetidae</taxonomicName>
, or if it is the result of long-branch attraction spurred on by increased inclusion of archaeocete and odontocete taxa in the phylogenetic analyses.
</paragraph>
<paragraph blockId="24.[145,762,197,1905]" lastBlockId="24.[809,1426,197,464]" pageId="24" pageNumber="1661">
The primary concern of the cladistic analysis was to discern the phylogenetic affinities of
<taxonomicName baseAuthorityName="AUGD" baseAuthorityYear="2469" box="[540,761,1761,1783]" class="Mammalia" family="Basilosauridae" genus="Kekenodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="24" pageNumber="1661" phylum="Chordata" rank="species" species="onamata">
<emphasis box="[540,761,1761,1783]" italics="true" pageId="24" pageNumber="1661">Kekenodon onamata</emphasis>
</taxonomicName>
within
<taxonomicName authorityName="BRISSON" authorityYear="1762" box="[221,310,1792,1813]" class="Insecta" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="24" pageNumber="1661" phylum="Arthropoda" rank="order">Cetacea</taxonomicName>
. The results under both equal and implied weighting posit
<taxonomicName baseAuthorityName="AUGD" baseAuthorityYear="2469" box="[342,585,1822,1844]" class="Mammalia" family="Basilosauridae" genus="Kekenodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="24" pageNumber="1661" phylum="Chordata" rank="species" species="onamata">
<emphasis box="[342,585,1822,1844]" italics="true" pageId="24" pageNumber="1661">Kekenodon onamata</emphasis>
</taxonomicName>
crownward to
<taxonomicName box="[145,309,1853,1875]" class="Mammalia" family="Basilosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="24" pageNumber="1661" phylum="Chordata" rank="family">Basilosauridae</taxonomicName>
as sister-taxon to Neoceti with moderate support (branch support = 64% and 81%, respectively), identifying
<taxonomicName baseAuthorityName="AUGD" baseAuthorityYear="2469" box="[930,1053,197,219]" class="Mammalia" family="Basilosauridae" genus="Kekenodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="24" pageNumber="1661" phylum="Chordata" rank="species" species="onamata">
<emphasis box="[930,948,198,219]" italics="true" pageId="24" pageNumber="1661">K</emphasis>
.
<emphasis box="[956,1053,198,219]" italics="true" pageId="24" pageNumber="1661">onamata</emphasis>
</taxonomicName>
as the latest-diverging archaeocete and expanding the temporal range of archaeocetes into the Late Oligocene. The inclusion of undescribed putative kekenodontids from the OU collections, including a wellpreserved specimen with a nearly complete skull (OU 22294), will further elucidate interfamilial relationships within the
<taxonomicName authorityName="Mitchell, 1989 sensu Fordyce" authorityYear="1992" box="[949,1146,381,403]" family="Kekenodontidae" pageId="24" pageNumber="1665" rank="family">Kekenodontidae</taxonomicName>
(e.g. monophyletic vs. paraphyletic) and evaluate the phylogenetic position of kekenodontids as sister to Neoceti.
</paragraph>
</subSubSection>
</treatment>
</document>