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<document ID-DOI="10.1038/s42003-021-02170-6" ID-GBIF-Dataset="3ab537ff-a436-454f-8ff4-320888ae57ae" ID-PMC="PMC8211792" ID-PubMed="34140631" ID-Zenodo-Dep="5013642" checkinTime="1624372779646" checkinUser="carolina" docAuthor="Deng, Tao, Lu, Xiaokang, Wang, Shiqi, Flynn, Lawrence J., Sun, Danhui, He, Wen &amp; Chen, Shanqin" docDate="2021" docId="301787BAFFFBFFF3FF3CFA92FD6B8AF2" docLanguage="en" docName="CommunBiol.639.1-10.pdf" docOrigin="Communications Biology 639 (1)" docSource="http://dx.doi.org/10.1038/s42003-021-02170-6" docTitle="Paraceratherium linxiaense Deng &amp; Lu &amp; Wang &amp; Flynn &amp; Sun &amp; He &amp; Chen 2021, sp. nov." docType="treatment" docVersion="13" lastPageNumber="3" masterDocId="CC2EFFC2FFFAFFF1FFB2FFB8FFCC8C5D" masterDocTitle="An Oligocene giant rhino provides insights into Paraceratherium evolution" masterLastPageNumber="10" masterPageNumber="1" pageNumber="2" updateTime="1668126836188" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0" zenodo-license-figures="CC-BY-4.0">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>An Oligocene giant rhino provides insights into Paraceratherium evolution</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Deng, Tao</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Lu, Xiaokang</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Wang, Shiqi</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Flynn, Lawrence J.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Sun, Danhui</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>He, Wen</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Chen, Shanqin</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
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<mods:titleInfo>
<mods:title>Communications Biology</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2021</mods:date>
<mods:detail type="pubDate">
<mods:number>2021-06-17</mods:number>
</mods:detail>
<mods:detail type="volume">
<mods:number>639</mods:number>
</mods:detail>
<mods:detail type="issue">
<mods:number>1</mods:number>
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<mods:url>http://dx.doi.org/10.1038/s42003-021-02170-6</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">10.1038/s42003-021-02170-6</mods:identifier>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.5041058" ID-GBIF-Taxon="183122940" ID-Zenodo-Dep="5041058" LSID="urn:lsid:plazi:treatment:301787BAFFFBFFF3FF3CFA92FD6B8AF2" httpUri="http://treatment.plazi.org/id/301787BAFFFBFFF3FF3CFA92FD6B8AF2" lastPageId="2" lastPageNumber="3" pageId="1" pageNumber="2">
<subSubSection box="[142,506,1322,1343]" pageId="1" pageNumber="2" type="nomenclature">
<paragraph blockId="1.[116,659,1238,1343]" box="[142,506,1322,1343]" pageId="1" pageNumber="2">
<taxonomicName authority=" Deng &amp; Lu &amp; Wang &amp; Flynn &amp; Sun &amp; He &amp; Chen, 2021" authorityName=" Deng &amp; Lu &amp; Wang &amp; Flynn &amp; Sun &amp; He &amp; Chen" authorityYear="2021" box="[142,417,1322,1343]" class="Mammalia" family="Hyracodontidae" genus="Paraceratherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Perissodactyla" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="linxiaense" status="sp. nov.">
<emphasis box="[142,417,1322,1343]" italics="true" pageId="1" pageNumber="2">Paraceratherium linxiaense</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[426,506,1322,1343]" pageId="1" pageNumber="2" rank="species">sp. nov.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection pageId="1" pageNumber="2" type="materials_examined">
<paragraph blockId="1.[116,778,1392,1972]" pageId="1" pageNumber="2">
<emphasis bold="true" box="[116,284,1392,1414]" pageId="1" pageNumber="2">
<typeStatus box="[116,168,1392,1413]" pageId="1" pageNumber="2">Type</typeStatus>
specimens
</emphasis>
.
<materialsCitation ID-GBIF-Occurrence="3312307301" collectionCode="HMV" country="China" elevation="1983" latitude="35.584766" location="Linxia Basin" longLatPrecision="1" longitude="103.31418" pageId="1" pageNumber="2" specimenCode="HMV 2006" specimenCount="1" specimenCount-adult="1" stateProvince="Linxia Basin" typeStatus="holotype">
Acomplete skull and mandible with the associated atlas (
<typeStatus box="[246,337,1420,1442]" pageId="1" pageNumber="2">holotype</typeStatus>
,
<specimenCode box="[351,469,1420,1442]" pageId="1" pageNumber="2">HMV 2006</specimenCode>
,
<figureCitation box="[482,541,1420,1442]" captionStart="Fig" captionStartId="2.[116,146,937,956]" captionTargetBox="[155,1434,136,912]" captionTargetId="figure-625@2.[154,1435,135,913]" captionTargetPageId="2" captionText="Fig. 1 Holotype (HMV 2006) of Paraceratherium linxiaense sp. nov. Skull: a lateral view; b ventral view; c dorsal view; d anterior view; e occipital view. Mandible: f, h lateral view and medial view of left ramus, respectively; g occlusal view. Skull and mandible share the scale bar, but both anterior and nuchal views have an independent scale bar." figureDoi="http://doi.org/10.5281/zenodo.5013644" httpUri="https://zenodo.org/record/5013644/files/figure.png" pageId="1" pageNumber="2">Fig. 1</figureCitation>
), and an axis and two thoracic vertebrae of another individual (paratype, HMV 2007,
<figureCitation box="[116,179,1475,1497]" captionStart="Fig" captionStartId="3.[116,146,1378,1397]" captionTargetBox="[362,1224,137,1350]" captionTargetId="figure-393@3.[361,1226,135,1353]" captionTargetPageId="3" captionText="Fig. 2 Vertebrae of Paraceratherium linxiaense. a atlas (HMV 2006); b axis (HMV 2007); c 4th5th articulated thoracic vertebrae (HMV 2007). 1, right view; 2, posterior view; 3, left view; 4, dorsal view; 5, anterior view; 6, ventral view; 7, postero-ventral view. The articular facets on 4th-5th thoracic vertebrae are marked with the red semitransparent outline whatever it is intact or has been damaged." figureDoi="http://doi.org/10.5281/zenodo.5013648" httpUri="https://zenodo.org/record/5013648/files/figure.png" pageId="1" pageNumber="2">Fig. 2</figureCitation>
), which are preserved at the Hezheng Paleozoological Museum in Hezheng County, Gansu Province, China. HMV 2006 represents a full
<specimenCount box="[292,343,1531,1553]" pageId="1" pageNumber="2" type="adult">adult</specimenCount>
individual. The specific name,
<taxonomicName authorityName="evolution &amp; Deng &amp; Lu &amp; Wang &amp; Flynn &amp; Sun &amp; He &amp; Chen" authorityYear="2021" box="[672,774,1531,1553]" class="Mammalia" family="Hyracodontidae" genus="Paraceratherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Perissodactyla" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="linxiaense">linxiaense</taxonomicName>
, refers to the geographical location of the discovery in the
<collectingRegion pageId="1" pageNumber="2">Linxia Basin</collectingRegion>
(
<figureCitation box="[187,246,1587,1609]" captionStart="Fig" captionStartId="4.[116,146,836,855]" captionTargetBox="[361,1226,135,812]" captionTargetId="figure-319@4.[361,1226,135,812]" captionTargetPageId="4" captionText="Fig. 3 Type locality of Paraceratherium linxiaense. Map showing the giant rhino fossil locality of the Linxia Basin in Wangjiachuan Village, Dongxiang County, Gansu Province, China." figureDoi="http://doi.org/10.5281/zenodo.5013650" httpUri="https://zenodo.org/record/5013650/files/figure.png" pageId="1" pageNumber="2">Fig. 3</figureCitation>
).
</materialsCitation>
</paragraph>
<paragraph blockId="1.[116,778,1392,1972]" pageId="1" pageNumber="2">
<emphasis bold="true" box="[116,384,1657,1679]" pageId="1" pageNumber="2">Type locality and horizon</emphasis>
. IVPP locality LX1808 (
<geoCoordinate box="[626,772,1656,1679]" degrees="35" direction="north" minutes="35" orientation="latitude" pageId="1" pageNumber="2" precision="1" seconds="05.16" value="35.584766">N35°3505.16”</geoCoordinate>
,
<geoCoordinate box="[116,269,1684,1707]" degrees="103" direction="east" minutes="18" orientation="longitude" pageId="1" pageNumber="2" precision="1" seconds="51.02" value="103.31418">E103°1851.02”</geoCoordinate>
;
<quantity box="[282,357,1685,1706]" metricMagnitude="3" metricUnit="m" metricValue="1.983" pageId="1" pageNumber="2" unit="m" value="1983.0">
<elevation box="[282,357,1685,1706]" metricMagnitude="3" metricUnit="m" metricValue="1.983" pageId="1" pageNumber="2" unit="m" value="1983.0">1983 m</elevation>
</quantity>
above sea level,
<figureCitation box="[527,584,1685,1707]" captionStart="Fig" captionStartId="4.[116,146,1421,1440]" captionTargetBox="[362,1226,929,1396]" captionTargetId="figure-347@4.[362,1226,929,1396]" captionTargetPageId="4" captionText="Fig. 4 Type horizon of Paraceratherium linxiaense. Exposures of fluvial and lacustrine sediments of the Linxia Basin (LX 1808) at the type locality of this new species (HMV 2006, yellow square) and where the axis and thoracic vertebrae were also found in Wangjiachuan Village, Dongxiang County, Gansu Province, China." figureDoi="http://doi.org/10.5281/zenodo.5013652" httpUri="https://zenodo.org/record/5013652/files/figure.png" pageId="1" pageNumber="2">Fig. 4</figureCitation>
)
<locationDeviation pageId="1" pageNumber="2">is near the village of Wangjiachuan</locationDeviation>
,
<locationDeviation pageId="1" pageNumber="2">
<quantity box="[307,387,1713,1735]" metricMagnitude="4" metricUnit="m" metricValue="1.08" pageId="1" pageNumber="2" unit="km" value="10.8">10.8 km</quantity>
southwest of the town of Dongxiang County
</locationDeviation>
, Linxia Hui Autonomous Prefecture,
<collectingRegion box="[605,774,1740,1762]" country="China" name="Gansu" pageId="1" pageNumber="2">Gansu Province</collectingRegion>
,
<collectingCountry box="[116,177,1768,1790]" name="China" pageId="1" pageNumber="2">China</collectingCountry>
(
<figureCitation box="[193,252,1768,1790]" captionStart="Fig" captionStartId="4.[116,146,836,855]" captionTargetBox="[361,1226,135,812]" captionTargetId="figure-319@4.[361,1226,135,812]" captionTargetPageId="4" captionText="Fig. 3 Type locality of Paraceratherium linxiaense. Map showing the giant rhino fossil locality of the Linxia Basin in Wangjiachuan Village, Dongxiang County, Gansu Province, China." figureDoi="http://doi.org/10.5281/zenodo.5013650" httpUri="https://zenodo.org/record/5013650/files/figure.png" pageId="1" pageNumber="2">Fig. 3</figureCitation>
). HMV 2006 and 2007 are from the sandstones in the lower part of the Jiaozigou Formation (
<figureCitation box="[555,613,1796,1818]" captionStart="Fig" captionStartId="5.[116,146,1600,1619]" captionTargetBox="[363,1225,136,1574]" captionTargetId="figure-196@5.[362,1226,135,1576]" captionTargetPageId="5" captionText="Fig. 5 Cenozoic section of the Linxia Basin. Deposits with correlation to ATNTS 201249 (left) for the magnetostratigraphic results50 (middle) and the Wangjiachuan section bearing the fossils of Paraceratherium linxiaense sp. nov. (right)." figureDoi="http://doi.org/10.5281/zenodo.5013654" httpUri="https://zenodo.org/record/5013654/files/figure.png" pageId="1" pageNumber="2">Fig. 5</figureCitation>
, Supplementary Note 1).
</paragraph>
<paragraph blockId="1.[116,778,1392,1972]" pageId="1" pageNumber="2">
<emphasis bold="true" box="[116,157,1894,1916]" pageId="1" pageNumber="2">Age</emphasis>
. IVPP locality LX1808 is faunally and paleomagnetically dated to the middle of chron C8r with an estimated age of 26.5 million years ago (Ma) in the late Oligocene (
<figureCitation box="[589,647,1949,1972]" captionStart="Fig" captionStartId="5.[116,146,1600,1619]" captionTargetBox="[363,1225,136,1574]" captionTargetId="figure-196@5.[362,1226,135,1576]" captionTargetPageId="5" captionText="Fig. 5 Cenozoic section of the Linxia Basin. Deposits with correlation to ATNTS 201249 (left) for the magnetostratigraphic results50 (middle) and the Wangjiachuan section bearing the fossils of Paraceratherium linxiaense sp. nov. (right)." figureDoi="http://doi.org/10.5281/zenodo.5013654" httpUri="https://zenodo.org/record/5013654/files/figure.png" pageId="1" pageNumber="2">Fig. 5</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="1" pageNumber="2" type="diagnosis">
<paragraph blockId="1.[810,1472,136,1972]" pageId="1" pageNumber="2">
<emphasis bold="true" box="[810,914,136,158]" pageId="1" pageNumber="2">Diagnosis</emphasis>
.
<taxonomicName authorityName="evolution &amp; Deng &amp; Lu &amp; Wang &amp; Flynn &amp; Sun &amp; He &amp; Chen" authorityYear="2021" box="[934,1216,136,157]" class="Mammalia" family="Hyracodontidae" genus="Paraceratherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Perissodactyla" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="linxiaense">
<emphasis box="[934,1216,136,157]" italics="true" pageId="1" pageNumber="2">Paraceratherium linxiaense</emphasis>
</taxonomicName>
possesses features that characterize the genus, such as a giant body size, long premaxillae with anterior ends extending downward, separated parietal crests, high condyle compared to the height of nuchal surface, lower inferior border of the posttympanic process than the condyle, roughly horizontal anterior part of symphysis, and downward turning cone-shaped
<date box="[1035,1061,304,325]" pageId="1" pageNumber="2">I1.</date>
It is more derived than other species within this genus in having a larger body size, deeper nasal notch above M2, much higher occipital part and posterior end of zygomatic arch, and smaller upper incisor
<date box="[1252,1277,388,409]" pageId="1" pageNumber="2">I1.</date>
The lower margin of the horizontal mandibular ramus is concave under the diastema, and small
<date box="[1006,1024,443,465]" pageId="1" pageNumber="2">i1</date>
extends anteriorly and horizontally. The dental formula is 1.0.3.3/1.0.3.3. P2 is semimolarized, while P3 and P4 are submolarized. The metaconule connects with the ectoloph and the anterior point of the hypocone in moderate wear; the antecrochet is moderate; the lingual border of the protocone is rounded on molars; and the ecto-posterior corner of the protolophid is angular on p3 and p4. The atlas has an expanded transverse foramen and a dumb-bell shaped vertebral fossa.
</paragraph>
</subSubSection>
<subSubSection lastPageId="2" lastPageNumber="3" pageId="1" pageNumber="2" type="description">
<paragraph blockId="1.[810,1472,136,1972]" pageId="1" pageNumber="2">
<emphasis bold="true" box="[810,1076,750,772]" pageId="1" pageNumber="2">Comparative description</emphasis>
. The new species differs from other species of
<taxonomicName authorityName="Forster-Cooper" authorityYear="1911" box="[932,1100,778,799]" class="Mammalia" family="Hyracodontidae" genus="Paraceratherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Perissodactyla" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
<emphasis box="[932,1100,778,799]" italics="true" pageId="1" pageNumber="2">Paraceratherium</emphasis>
</taxonomicName>
(
<emphasis box="[1126,1244,779,800]" italics="true" pageId="1" pageNumber="2">P. grangeri</emphasis>
,
<emphasis box="[1266,1429,778,800]" italics="true" pageId="1" pageNumber="2">P. huangheense</emphasis>
,
<emphasis italics="true" pageId="1" pageNumber="2">P. asiaticum</emphasis>
,
<emphasis box="[920,1039,806,828]" italics="true" pageId="1" pageNumber="2">P. bugtiense</emphasis>
, and
<emphasis box="[1098,1207,806,828]" italics="true" pageId="1" pageNumber="2">P. lepidum</emphasis>
) in having a deeper nasal notch whose bottom is located above the middle of M2, proportionally larger height of the condyle (43.9%) compared to height of the occipital surface (Supplementary Table 2), short muzzle bones and diastema anterior to cheek teeth, highly raised occiput, and high zygomatic arch with a prominent posterior end (
<figureCitation box="[817,876,973,995]" captionStart="Fig" captionStartId="2.[116,146,937,956]" captionTargetBox="[155,1434,136,912]" captionTargetId="figure-625@2.[154,1435,135,913]" captionTargetPageId="2" captionText="Fig. 1 Holotype (HMV 2006) of Paraceratherium linxiaense sp. nov. Skull: a lateral view; b ventral view; c dorsal view; d anterior view; e occipital view. Mandible: f, h lateral view and medial view of left ramus, respectively; g occlusal view. Skull and mandible share the scale bar, but both anterior and nuchal views have an independent scale bar." figureDoi="http://doi.org/10.5281/zenodo.5013644" httpUri="https://zenodo.org/record/5013644/files/figure.png" pageId="1" pageNumber="2">Fig. 1</figureCitation>
).
</paragraph>
<paragraph blockId="1.[810,1472,136,1972]" pageId="1" pageNumber="2">
The nasals of
<taxonomicName authorityName="evolution &amp; Deng &amp; Lu &amp; Wang &amp; Flynn &amp; Sun &amp; He &amp; Chen" authorityYear="2021" box="[981,1108,1001,1023]" class="Mammalia" family="Hyracodontidae" genus="Paraceratherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Perissodactyla" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="linxiaense">
<emphasis box="[981,1108,1001,1023]" italics="true" pageId="1" pageNumber="2">P. linxiaense</emphasis>
</taxonomicName>
are flat and straight, and the nasal notch is very deep above the middle of M2 with a short distance from the orbit (15.3% of the basal cranial length) (Supplementary Table 2), much deeper than those of other species of
<taxonomicName authorityName="Forster-Cooper" authorityYear="1911" box="[810,978,1113,1134]" class="Mammalia" family="Hyracodontidae" genus="Paraceratherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Perissodactyla" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
<emphasis box="[810,978,1113,1134]" italics="true" pageId="1" pageNumber="2">Paraceratherium</emphasis>
</taxonomicName>
except
<emphasis box="[1058,1167,1113,1135]" italics="true" pageId="1" pageNumber="2">P. lepidum</emphasis>
, indicating a short prehensile nose trunk. The dorsal surface of the skull is shallowly depressed, different from the domed skull of
<emphasis box="[1161,1270,1169,1190]" italics="true" pageId="1" pageNumber="2">P. grangeri</emphasis>
13 or the flat one of
<emphasis box="[810,920,1196,1218]" italics="true" pageId="1" pageNumber="2">P. lepidum</emphasis>
7. The distance between the parietal crests is narrow and smaller than that of
<emphasis box="[1069,1178,1224,1246]" italics="true" pageId="1" pageNumber="2">P. lepidum</emphasis>
. The infraorbital foramen is situated above the P4/M1 boundary, and the anterior margin of the orbit is located above the M2/M3 boundary. Both characters are similar to those of
<emphasis box="[1035,1142,1308,1330]" italics="true" pageId="1" pageNumber="2">P. lepidum</emphasis>
and more posteriorly positioned than those of
<emphasis box="[966,1080,1337,1358]" italics="true" pageId="1" pageNumber="2">P. grangeri</emphasis>
above the P3/P4 boundary and the middle of M2 respectively. The position of the zygomatic arch is high, posteriorly flush with the upper margin of the orbit like those of
<emphasis box="[896,1004,1420,1441]" italics="true" pageId="1" pageNumber="2">P. grangeri</emphasis>
and
<emphasis box="[1055,1162,1420,1441]" italics="true" pageId="1" pageNumber="2">P. lepidum</emphasis>
, and much higher than that of
<emphasis box="[810,937,1448,1469]" italics="true" pageId="1" pageNumber="2">P. bugtiense</emphasis>
14. The postorbital process is absent. The space between the posttympanic and postglenoid processes is moderate as in
<emphasis box="[866,974,1503,1525]" italics="true" pageId="1" pageNumber="2">P. lepidum</emphasis>
, wider than in
<emphasis box="[1134,1243,1504,1525]" italics="true" pageId="1" pageNumber="2">P. grangeri</emphasis>
and narrower than in
<emphasis box="[810,940,1531,1553]" italics="true" pageId="1" pageNumber="2">P. bugtiense</emphasis>
. The posttympanic process has no transverse expansion, and its lateral margin is almost flush with the postglenoid process. The posttympanic and paraoccipital processes fuse to become a wide and thick plate.
</paragraph>
<paragraph blockId="1.[810,1472,136,1972]" pageId="1" pageNumber="2">
For
<taxonomicName authorityName="evolution &amp; Deng &amp; Lu &amp; Wang &amp; Flynn &amp; Sun &amp; He &amp; Chen" authorityYear="2021" box="[883,1020,1643,1664]" class="Mammalia" family="Hyracodontidae" genus="Paraceratherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Perissodactyla" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="linxiaense">
<emphasis box="[883,1020,1643,1664]" italics="true" pageId="1" pageNumber="2">P. linxiaense</emphasis>
</taxonomicName>
, the posterior border of the mandibular symphysis is situated at the p4/m1 boundary, and the posterior margin of the ascending ramus is anteriorly inclined, different from vertical situation in
<emphasis box="[1101,1230,1726,1748]" italics="true" pageId="1" pageNumber="2">P. bugtiense</emphasis>
and
<emphasis box="[1295,1412,1726,1748]" italics="true" pageId="1" pageNumber="2">P. lepidum</emphasis>
. The vascular notch of lower margin of the mandible is deep but more anterior than that of
<emphasis box="[1146,1268,1782,1804]" italics="true" pageId="1" pageNumber="2">P. lepidum</emphasis>
. The mandibular diastema has a straight and slowly declining upper margin like in
<emphasis box="[844,963,1839,1860]" italics="true" pageId="1" pageNumber="2">P. grangeri</emphasis>
, while strongly declined in
<emphasis box="[1285,1419,1839,1860]" italics="true" pageId="1" pageNumber="2">P. asiaticum</emphasis>
and
<emphasis box="[810,920,1866,1888]" italics="true" pageId="1" pageNumber="2">P. lepidum</emphasis>
, and convex in
<emphasis box="[1085,1243,1866,1888]" italics="true" pageId="1" pageNumber="2">P. huangheense</emphasis>
and
<emphasis box="[1296,1418,1866,1888]" italics="true" pageId="1" pageNumber="2">P. bugtiense</emphasis>
. The mental foramen is situated under the p3/p4 boundary, more posteriorly than in
<emphasis box="[1020,1151,1922,1943]" italics="true" pageId="1" pageNumber="2">P. asiaticum</emphasis>
,
<emphasis box="[1167,1292,1922,1943]" italics="true" pageId="1" pageNumber="2">P. bugtiense</emphasis>
, and
<emphasis box="[1358,1471,1922,1943]" italics="true" pageId="1" pageNumber="2">P. lepidum</emphasis>
where is under p2.
</paragraph>
<footnote box="[685,1379,2007,2023]" pageId="1" pageNumber="2">
<paragraph blockId="1.[685,1379,2007,2023]" box="[685,1379,2007,2023]" pageId="1" pageNumber="2">(2021) 4:639 | https://doi.org/10.1038/s42003-021-02170-6 | www.nature.com/commsbio</paragraph>
</footnote>
<caption ID-DOI="http://doi.org/10.5281/zenodo.5013644" ID-Zenodo-Dep="5013644" httpUri="https://zenodo.org/record/5013644/files/figure.png" pageId="2" pageNumber="3" startId="2.[116,146,937,956]" targetBox="[155,1434,136,912]" targetPageId="2">
<paragraph blockId="2.[116,1473,937,1012]" pageId="2" pageNumber="3">
<emphasis bold="true" box="[116,732,937,956]" pageId="2" pageNumber="3">
Fig. 1 Holotype (HMV 2006) of
<taxonomicName authority="evolution &amp; Deng &amp; Lu &amp; Wang &amp; Flynn &amp; Sun &amp; He &amp; Chen, 2021" authorityName="evolution &amp; Deng &amp; Lu &amp; Wang &amp; Flynn &amp; Sun &amp; He &amp; Chen" authorityYear="2021" box="[413,654,937,956]" class="Mammalia" family="Hyracodontidae" genus="Paraceratherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Perissodactyla" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="linxiaense" status="sp. nov.">
<emphasis bold="true" box="[413,654,937,956]" italics="true" pageId="2" pageNumber="3">Paraceratherium linxiaense</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[660,732,937,956]" pageId="2" pageNumber="3" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
Skull:
<emphasis bold="true" box="[790,801,937,956]" pageId="2" pageNumber="3">a</emphasis>
lateral view;
<emphasis bold="true" box="[920,931,937,956]" pageId="2" pageNumber="3">b</emphasis>
ventral view;
<emphasis bold="true" box="[1057,1067,937,956]" pageId="2" pageNumber="3">c</emphasis>
dorsal view;
<emphasis bold="true" box="[1184,1195,937,956]" pageId="2" pageNumber="3">d</emphasis>
anterior view;
<emphasis bold="true" box="[1329,1340,937,956]" pageId="2" pageNumber="3">e</emphasis>
occipital view. Mandible:
<emphasis bold="true" box="[209,216,965,984]" pageId="2" pageNumber="3">f</emphasis>
,
<emphasis bold="true" box="[226,237,965,984]" pageId="2" pageNumber="3">h</emphasis>
lateral view and medial view of left ramus, respectively;
<emphasis bold="true" box="[732,743,965,984]" pageId="2" pageNumber="3">g</emphasis>
occlusal view. Skull and mandible share the scale bar, but both anterior and nuchal views have an independent scale bar.
</paragraph>
</caption>
<paragraph blockId="2.[116,779,1047,1711]" pageId="2" pageNumber="3">
The distance between the anterior margins of
<date box="[608,629,1048,1069]" pageId="2" pageNumber="3">I1</date>
and P2 is
<quantity metricMagnitude="-1" metricUnit="m" metricValue="1.64" pageId="2" pageNumber="3" unit="mm" value="164.0">164 mm</quantity>
. DP1 is absent like in most species of
<taxonomicName authorityName="Forster-Cooper" authorityYear="1911" box="[562,730,1075,1096]" class="Mammalia" family="Hyracodontidae" genus="Paraceratherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Perissodactyla" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis box="[562,730,1075,1096]" italics="true" pageId="2" pageNumber="3">Paraceratherium</emphasis>
</taxonomicName>
, but present in
<emphasis box="[224,333,1104,1125]" italics="true" pageId="2" pageNumber="3">P. grangeri</emphasis>
. The paracone rib is absent from P2 to M1, weak in M2, and marked in M3, which is the common character of the genus
<taxonomicName authorityName="Forster-Cooper" authorityYear="1911" box="[259,427,1159,1180]" class="Mammalia" family="Hyracodontidae" genus="Paraceratherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Perissodactyla" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis box="[259,427,1159,1180]" italics="true" pageId="2" pageNumber="3">Paraceratherium</emphasis>
</taxonomicName>
, much different from the strong paracone rib of the primitive giant rhino
<taxonomicName authorityName="Wood" authorityYear="1938" box="[551,704,1188,1209]" class="Mammalia" family="Hyracodontidae" genus="Forstercooperia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Perissodactyla" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis box="[551,704,1188,1209]" italics="true" pageId="2" pageNumber="3">Forstercooperia</emphasis>
</taxonomicName>
15. The occlusal surface of P2 is triangular in
<taxonomicName authorityName="evolution &amp; Deng &amp; Lu &amp; Wang &amp; Flynn &amp; Sun &amp; He &amp; Chen" authorityYear="2021" box="[497,623,1215,1237]" class="Mammalia" family="Hyracodontidae" genus="Paraceratherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Perissodactyla" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="linxiaense">
<emphasis box="[497,623,1215,1237]" italics="true" pageId="2" pageNumber="3">P. linxiaense</emphasis>
</taxonomicName>
, different from the trapezium outline in
<emphasis box="[368,477,1243,1265]" italics="true" pageId="2" pageNumber="3">P. grangeri</emphasis>
and
<emphasis box="[528,652,1243,1265]" italics="true" pageId="2" pageNumber="3">P. asiaticum</emphasis>
. There is an obvious separating groove between the protocone and hypocone of P3 and P4. The hypocone of P4 is situated behind the metaconule as in
<emphasis box="[299,409,1327,1348]" italics="true" pageId="2" pageNumber="3">P. grangeri</emphasis>
and
<emphasis box="[464,573,1326,1348]" italics="true" pageId="2" pageNumber="3">P. lepidum</emphasis>
, and it is expanded and rounded, while that of
<emphasis box="[396,504,1354,1376]" italics="true" pageId="2" pageNumber="3">P. lepidum</emphasis>
is square. The antecrochet on upper molars is larger than those of
<emphasis box="[576,692,1383,1404]" italics="true" pageId="2" pageNumber="3">P. grangeri</emphasis>
and
<emphasis italics="true" pageId="2" pageNumber="3">P. asiaticum</emphasis>
. M3 is triangular in occlusal outline, similar to those of
<emphasis box="[116,238,1438,1460]" italics="true" pageId="2" pageNumber="3">P. bugtiense</emphasis>
and
<emphasis box="[296,407,1438,1460]" italics="true" pageId="2" pageNumber="3">P. lepidum</emphasis>
, but different from the rectangular outline in
<emphasis box="[224,334,1467,1488]" italics="true" pageId="2" pageNumber="3">P. grangeri</emphasis>
and
<emphasis box="[389,514,1467,1488]" italics="true" pageId="2" pageNumber="3">P. asiaticum</emphasis>
.
</paragraph>
<paragraph blockId="2.[116,779,1047,1711]" pageId="2" pageNumber="3">
<taxonomicName authorityName="evolution &amp; Deng &amp; Lu &amp; Wang &amp; Flynn &amp; Sun &amp; He &amp; Chen" authorityYear="2021" box="[142,268,1494,1515]" class="Mammalia" family="Hyracodontidae" genus="Paraceratherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Perissodactyla" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="linxiaense">
<emphasis box="[142,268,1494,1515]" italics="true" pageId="2" pageNumber="3">P. linxiaense</emphasis>
</taxonomicName>
has more reduced
<date box="[467,491,1494,1516]" pageId="2" pageNumber="3">i1.</date>
The p2 is small and singlerooted, but large and double-rooted in
<emphasis box="[510,618,1522,1543]" italics="true" pageId="2" pageNumber="3">P. grangeri</emphasis>
. The p3 and p4 have a rudimentary entolophid, while the entoconid of
<emphasis box="[116,242,1578,1599]" italics="true" pageId="2" pageNumber="3">P. asiaticum</emphasis>
is an isolated cusp. The entolophid of m3 is nearly transverse.
</paragraph>
</subSubSection>
<subSubSection pageId="2" pageNumber="3" type="discussion">
<paragraph blockId="2.[116,779,1047,1711]" pageId="2" pageNumber="3">Additional descriptions, measurements and comparisons, such as information of the postcranial bones, are provided in Supplementary Note 2 and Supplementary Tables 16.</paragraph>
</subSubSection>
</treatment>
</document>