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<document ID-DOI="10.1093/zoolinnean/zlab009" ID-GBIF-Dataset="6f2fedbe-5f67-47ae-9e8a-de80ce9f2167" ID-ISSN="0024-4082" ID-Zenodo-Dep="6351896" checkinTime="1647255935986" checkinUser="plazi" docAuthor="Antoine, Pierre-Olivier, Reyes, Marian C, Amano, Noel, Bautista, Angel P, Chang, Chun-Hsiang, Claude, Julien, De Vos, John &amp; Ingicco, Thomas" docDate="2022" docId="1B53A12D1C52BF40FC6FC5BB3446983D" docLanguage="en" docName="zlab009.pdf" docOrigin="Zoological Journal of the Linnean Society 194 (2)" docSource="https://academic.oup.com/zoolinnean/article/194/2/416/6280384" docStyle="DocumentStyle:36B3BD6A90C22AB4F7F465C853188CC8.7:ZoolJLinnSoc.2017-.journal_article" docStyleId="36B3BD6A90C22AB4F7F465C853188CC8" docStyleName="ZoolJLinnSoc.2017-.journal_article" docStyleVersion="7" docTitle="Nesorhinus Antoine &amp; Reyes &amp; Amano &amp; Bautista &amp; Chang &amp; Claude &amp; De Vos &amp; Ingicco 2022, GEN. NOV." docType="treatment" docVersion="4" lastPageNumber="423" masterDocId="E76AD9551C57BF47FFA0C32930769A77" masterDocTitle="A new rhinoceros clade from the Pleistocene of Asia sheds light on mammal dispersals to the Philippines" masterLastPageNumber="430" masterPageNumber="416" pageNumber="421" updateTime="1674141033289" updateUser="diego">
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<mods:titleInfo>
<mods:title>A new rhinoceros clade from the Pleistocene of Asia sheds light on mammal dispersals to the Philippines</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Antoine, Pierre-Olivier</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Reyes, Marian C</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Amano, Noel</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Bautista, Angel P</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Chang, Chun-Hsiang</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Claude, Julien</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>De Vos, John</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Ingicco, Thomas</mods:namePart>
</mods:name>
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<mods:title>Zoological Journal of the Linnean Society</mods:title>
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<mods:part>
<mods:date>2022</mods:date>
<mods:detail type="pubDate">
<mods:number>2022-02-01</mods:number>
</mods:detail>
<mods:detail type="volume">
<mods:number>194</mods:number>
</mods:detail>
<mods:detail type="issue">
<mods:number>2</mods:number>
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<mods:start>416</mods:start>
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<mods:url>https://academic.oup.com/zoolinnean/article/194/2/416/6280384</mods:url>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">10.1093/zoolinnean/zlab009</mods:identifier>
<mods:identifier type="GBIF-Dataset">6f2fedbe-5f67-47ae-9e8a-de80ce9f2167</mods:identifier>
<mods:identifier type="ISSN">0024-4082</mods:identifier>
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<treatment ID-GBIF-Taxon="194018689" LSID="urn:lsid:plazi:treatment:1B53A12D1C52BF40FC6FC5BB3446983D" httpUri="http://treatment.plazi.org/id/1B53A12D1C52BF40FC6FC5BB3446983D" lastPageId="7" lastPageNumber="423" pageId="5" pageNumber="421">
<subSubSection box="[975,1259,1682,1706]" pageId="5" pageNumber="421" type="nomenclature">
<paragraph blockId="5.[975,1259,1682,1706]" box="[975,1259,1682,1706]" pageId="5" pageNumber="421">
<heading box="[975,1259,1682,1706]" centered="true" fontSize="9" level="2" pageId="5" pageNumber="421" reason="2">
<taxonomicName authority="Antoine &amp; Reyes &amp; Amano &amp; Bautista &amp; Chang &amp; Claude &amp; De Vos &amp; Ingicco, 2022" authorityName="Antoine &amp; Reyes &amp; Amano &amp; Bautista &amp; Chang &amp; Claude &amp; De Vos &amp; Ingicco" authorityYear="2022" box="[975,1133,1682,1705]" class="Mammalia" family="Rhinocerotidae" genus="Nesorhinus" kingdom="Animalia" order="Perissodactyla" pageId="5" pageNumber="421" phylum="Chordata" rank="genus" status="GEN. NOV.">NESORHINUS</taxonomicName>
<emphasis bold="true" box="[1140,1259,1682,1706]" pageId="5" pageNumber="421">
<taxonomicNameLabel box="[1140,1259,1682,1706]" pageId="5" pageNumber="421" rank="genus">GEN. NOV.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection pageId="5" pageNumber="421" type="description">
<paragraph blockId="5.[1074,1159,1721,1746]" box="[1074,1159,1721,1746]" pageId="5" pageNumber="421">
(
<figureCitation box="[1083,1151,1721,1746]" captionStart="Figure 3" captionStartId="6.[164,244,1179,1201]" captionTargetBox="[176,1426,211,1137]" captionTargetId="figure-257@6.[163,1443,197,1140]" captionTargetPageId="6" captionText="Figure 3. Dental and postcranial features characterizing the new genus Nesorhinus in the phylogenetic framework as depicted in Figure 2. Red circles denote synapomorphies of Nesorhinus, whilst green and blue circles correspond to diagnostic characters (autapomorphies) of N. philippinensis and of N. hayasakai, respectively. Nesorhinus philippinensis: A, left upper dental series (D1M1) in occlusal view (II-2014-J1-294, 095, 409, 427); B, left m23 in occlusal view (II-2014-J1-405); C, right p3m1 in labial view (II-2014-J1-451); D, left scapula in lateral view (II-2014-J1-291). Nesorhinus hayasakai: E, left upper dental series (D1M3) in occlusal view (DGNTU-FV11b; modified from Hayasaka, 1942); F, right m3 in occlusolabial view (HTR-55); G, left m3 in labial view (HTR-91); H, left fragmentary scapula (HTR-1). Completeness of the skeleton of N. philippinensis found at Kalinga (I): preserved elements appear in dark green. Tentative silhouettes of N philippinensis (J) and N. hayasakai (K) are drawn at a same scale, with a shoulder height of 1.26 m for N. philippinensis. See Supporting Information for further details on body size. Scale bar, 5 cm (AC, EG) and 10 cm (D, H)." figureDoi="http://doi.org/10.5281/zenodo.6351902" httpUri="https://zenodo.org/record/6351902/files/figure.png" pageId="5" pageNumber="421">FIG. 3</figureCitation>
)
</paragraph>
<paragraph blockId="5.[809,1426,1761,1814]" pageId="5" pageNumber="421">
<emphasis box="[809,1089,1761,1783]" italics="true" pageId="5" pageNumber="421">Zoobank registration:</emphasis>
<uri pageId="5" pageNumber="421">
urn:lsid:zoobank.org:act:
<uuid box="[809,1325,1792,1814]" pageId="5" pageNumber="421">CE90CF70-64F7-4665-AD9C-367B4D662372</uuid>
</uri>
.
</paragraph>
</subSubSection>
<subSubSection lastPageId="6" lastPageNumber="422" pageId="5" pageNumber="421" type="etymology">
<paragraph blockId="5.[809,1426,1854,1907]" lastBlockId="6.[163,777,1510,1562]" lastPageId="6" lastPageNumber="422" pageId="5" pageNumber="421">
<emphasis box="[809,943,1854,1875]" italics="true" pageId="5" pageNumber="421">Etymology:</emphasis>
From the ancient Greek
<emphasis box="[1266,1330,1854,1877]" italics="true" pageId="5" pageNumber="421">
n
<emphasis bold="true" box="[1282,1293,1854,1877]" italics="true" pageId="5" pageNumber="421"></emphasis>
sos
</emphasis>
(νῆσος, island) and the suffix -
<emphasis box="[1083,1161,1885,1906]" italics="true" pageId="5" pageNumber="421">rhinus</emphasis>
(from Greek ῥίς,
<emphasis box="[1371,1418,1885,1906]" italics="true" pageId="5" pageNumber="421">rhis</emphasis>
, nose), frequently used for designating rhinocerotid genera.
</paragraph>
</subSubSection>
<caption ID-DOI="http://doi.org/10.5281/zenodo.6351902" ID-Zenodo-Dep="6351902" httpUri="https://zenodo.org/record/6351902/files/figure.png" pageId="6" pageNumber="422" startId="6.[164,244,1179,1201]" targetBox="[176,1426,211,1137]" targetPageId="6">
<paragraph blockId="6.[163,1443,1179,1466]" pageId="6" pageNumber="422">
<emphasis bold="true" box="[164,271,1179,1201]" pageId="6" pageNumber="422">Figure 3.</emphasis>
Dental and postcranial features characterizing the new genus
<taxonomicName authorityName="Antoine &amp; Reyes &amp; Amano &amp; Bautista &amp; Chang &amp; Claude &amp; De Vos &amp; Ingicco" authorityYear="2022" box="[955,1077,1180,1202]" class="Mammalia" family="Rhinocerotidae" genus="Nesorhinus" kingdom="Animalia" order="Perissodactyla" pageId="6" pageNumber="422" phylum="Chordata" rank="genus">
<emphasis box="[955,1077,1180,1202]" italics="true" pageId="6" pageNumber="422">Nesorhinus</emphasis>
</taxonomicName>
in the phylogenetic framework as depicted in Figure 2. Red circles denote synapomorphies of
<taxonomicName authorityName="Antoine &amp; Reyes &amp; Amano &amp; Bautista &amp; Chang &amp; Claude &amp; De Vos &amp; Ingicco" authorityYear="2022" box="[766,885,1209,1231]" class="Mammalia" family="Rhinocerotidae" genus="Nesorhinus" kingdom="Animalia" order="Perissodactyla" pageId="6" pageNumber="422" phylum="Chordata" rank="genus">
<emphasis box="[766,885,1209,1231]" italics="true" pageId="6" pageNumber="422">Nesorhinus</emphasis>
</taxonomicName>
, whilst green and blue circles correspond to diagnostic characters (autapomorphies) of
<taxonomicName baseAuthorityName="von Koenigswald" baseAuthorityYear="1956" box="[494,672,1238,1260]" class="Mammalia" family="Rhinocerotidae" genus="Nesorhinus" kingdom="Animalia" order="Perissodactyla" pageId="6" pageNumber="422" phylum="Chordata" rank="species" species="philippinensis">
<emphasis box="[494,672,1238,1260]" italics="true" pageId="6" pageNumber="422">N. philippinensis</emphasis>
</taxonomicName>
and of
<taxonomicName authorityName="Antoine &amp; Reyes &amp; Amano &amp; Bautista &amp; Chang &amp; Claude &amp; De Vos &amp; Ingicco" authorityYear="2022" baseAuthorityName="Otsuka &amp; Lin" baseAuthorityYear="1984" box="[747,883,1238,1260]" class="Mammalia" family="Rhinocerotidae" genus="Nesorhinus" kingdom="Animalia" order="Perissodactyla" pageId="6" pageNumber="422" phylum="Chordata" rank="species" species="hayasakai">
<emphasis box="[747,883,1238,1260]" italics="true" pageId="6" pageNumber="422">N. hayasakai</emphasis>
</taxonomicName>
, respectively.
<taxonomicName baseAuthorityName="von Koenigswald" baseAuthorityYear="1956" box="[1028,1303,1238,1260]" class="Mammalia" family="Rhinocerotidae" genus="Nesorhinus" kingdom="Animalia" order="Perissodactyla" pageId="6" pageNumber="422" phylum="Chordata" rank="species" species="philippinensis">
<emphasis box="[1028,1303,1238,1260]" italics="true" pageId="6" pageNumber="422">Nesorhinus philippinensis</emphasis>
</taxonomicName>
: A, left upper dental series (D1M1) in occlusal view (II-2014-J1-294, 095, 409, 427); B, left m23 in occlusal view (II-2014-J1-405); C, right p3m1 in labial view (II-2014-J1-451); D, left scapula in lateral view (II-2014-J1-291).
<taxonomicName authorityName="Antoine &amp; Reyes &amp; Amano &amp; Bautista &amp; Chang &amp; Claude &amp; De Vos &amp; Ingicco" authorityYear="2022" baseAuthorityName="Otsuka &amp; Lin" baseAuthorityYear="1984" box="[1133,1367,1297,1319]" class="Mammalia" family="Rhinocerotidae" genus="Nesorhinus" kingdom="Animalia" order="Perissodactyla" pageId="6" pageNumber="422" phylum="Chordata" rank="species" species="hayasakai">
<emphasis box="[1133,1367,1297,1319]" italics="true" pageId="6" pageNumber="422">Nesorhinus hayasakai</emphasis>
</taxonomicName>
: E, left upper dental series (D1M3) in occlusal view (DGNTU-FV11b; modified from
<bibRefCitation author="Hayasaka" box="[974,1141,1326,1348]" firstAuthor="Hayasaka" pageId="6" pageNumber="422" pagination="55 - 109" refId="ref9647" refString="Hayasaka I. 1942. On the occurrence of mammalian remains in Taiwan. A preliminary summary. Taiwan Tigaku Kizi 13: 55 - 109." type="journal article" year="1942">Hayasaka, 1942</bibRefCitation>
); F, right m3 in occlusolabial view (HTR-55); G, left m3 in labial view (HTR-91); H, left fragmentary scapula (HTR-1). Completeness of the skeleton of
<taxonomicName baseAuthorityName="von Koenigswald" baseAuthorityYear="1956" box="[163,341,1385,1407]" class="Mammalia" family="Rhinocerotidae" genus="Nesorhinus" kingdom="Animalia" order="Perissodactyla" pageId="6" pageNumber="422" phylum="Chordata" rank="species" species="philippinensis">
<emphasis box="[163,341,1385,1407]" italics="true" pageId="6" pageNumber="422">N. philippinensis</emphasis>
</taxonomicName>
found at Kalinga (I): preserved elements appear in dark green. Tentative silhouettes of
<taxonomicName baseAuthorityName="von Koenigswald" baseAuthorityYear="1956" box="[1269,1443,1385,1407]" class="Mammalia" family="Rhinocerotidae" genus="Nesorhinus" kingdom="Animalia" order="Perissodactyla" pageId="6" pageNumber="422" phylum="Chordata" rank="species" species="philippinensis">
<emphasis box="[1269,1443,1385,1407]" italics="true" pageId="6" pageNumber="422">N philippinensis</emphasis>
</taxonomicName>
(J) and
<taxonomicName authorityName="Antoine &amp; Reyes &amp; Amano &amp; Bautista &amp; Chang &amp; Claude &amp; De Vos &amp; Ingicco" authorityYear="2022" baseAuthorityName="Otsuka &amp; Lin" baseAuthorityYear="1984" box="[243,379,1414,1436]" class="Mammalia" family="Rhinocerotidae" genus="Nesorhinus" kingdom="Animalia" order="Perissodactyla" pageId="6" pageNumber="422" phylum="Chordata" rank="species" species="hayasakai">
<emphasis box="[243,379,1414,1436]" italics="true" pageId="6" pageNumber="422">N. hayasakai</emphasis>
</taxonomicName>
(K) are drawn at a same scale, with a shoulder height of 1.26 m for
<taxonomicName baseAuthorityName="von Koenigswald" baseAuthorityYear="1956" box="[1092,1270,1414,1436]" class="Mammalia" family="Rhinocerotidae" genus="Nesorhinus" kingdom="Animalia" order="Perissodactyla" pageId="6" pageNumber="422" phylum="Chordata" rank="species" species="philippinensis">
<emphasis box="[1092,1270,1414,1436]" italics="true" pageId="6" pageNumber="422">N. philippinensis</emphasis>
</taxonomicName>
. See Supporting Information for further details on body size. Scale bar, 5 cm (AC, EG) and 10 cm (D, H).
</paragraph>
</caption>
<subSubSection pageId="6" pageNumber="422" type="type_taxon">
<paragraph blockId="6.[163,777,1595,1678]" pageId="6" pageNumber="422">
<emphasis box="[163,708,1595,1617]" italics="true" pageId="6" pageNumber="422">Type species: Nesorhinus philippinensis</emphasis>
(
<bibRefCitation author="von Koenigswald GHR" firstAuthor="von Koenigswald" pageId="6" pageNumber="422" pagination="1 - 14" refId="ref10231" refString="von Koenigswald GHR. 1956. Fossil mammals from the Philippines. Quezon City: National Research Council of the Philippines, 1 - 14." type="book chapter" year="1956">von Koenigswald, 1956</bibRefCitation>
) comb. nov. See Supporting Information for further details.
</paragraph>
</subSubSection>
<subSubSection pageId="6" pageNumber="422" type="synonymic_list">
<paragraph blockId="6.[163,779,1719,1772]" pageId="6" pageNumber="422">
<emphasis box="[163,352,1719,1741]" italics="true" pageId="6" pageNumber="422">Referred species:</emphasis>
<taxonomicName authority="(Otsuka &amp; Lin, 1984)" authorityName="Antoine &amp; Reyes &amp; Amano &amp; Bautista &amp; Chang &amp; Claude &amp; De Vos &amp; Ingicco" authorityYear="2022" baseAuthorityName="Otsuka &amp; Lin" baseAuthorityYear="1984" class="Mammalia" family="Rhinocerotidae" genus="Nesorhinus" kingdom="Animalia" order="Perissodactyla" pageId="6" pageNumber="422" phylum="Chordata" rank="species" species="hayasakai" status="comb. nov.">
<emphasis bold="true" box="[364,651,1719,1741]" italics="true" pageId="6" pageNumber="422">Nesorhinus hayasakai</emphasis>
(
<bibRefCitation author="Otsuka &amp; Lin" firstAuthor="Otsuka" pageId="6" pageNumber="422" pagination="1 - 35" refId="ref11145" refString="Otsuka H, Lin CC. 1984. Fossil rhinoceros from the T'ouk'oushan Group in Taiwan. Journal of Taiwan Museum 37: 1 - 35." type="journal article" year="1984">Otsuka &amp; Lin, 1984</bibRefCitation>
)
</taxonomicName>
<emphasis bold="true" box="[285,421,1750,1771]" pageId="6" pageNumber="422">
<taxonomicNameLabel box="[285,421,1750,1771]" pageId="6" pageNumber="422" rank="species">comb. nov.</taxonomicNameLabel>
</emphasis>
</paragraph>
</subSubSection>
<subSubSection lastPageId="7" lastPageNumber="423" pageId="6" pageNumber="422" type="diagnosis">
<paragraph blockId="6.[163,778,1815,1898]" lastBlockId="7.[145,760,197,250]" lastPageId="7" lastPageNumber="423" pageId="6" pageNumber="422">
<emphasis box="[163,316,1816,1837]" italics="true" pageId="6" pageNumber="422">Diagnosis:</emphasis>
Medium-sized rhinocerotines, characterized by roots fully isolated on upper cheek teeth, a crochet usually present on P2-4, a crista absent on P3, a protocone constriction always absent and a posterior half of the ectoloph concave on M1-2, a trigonid angular in occlusal view on lower cheek teeth and a posterior supraglenoid tubercle convex and salient on the scapula. Differing from representatives of both
<taxonomicName authorityName="Gloger" authorityYear="1841" box="[1104,1255,1663,1684]" class="Mammalia" family="Rhinocerotidae" genus="Dicerorhinus" kingdom="Animalia" order="Perissodactyla" pageId="6" pageNumber="422" phylum="Chordata" rank="genus">
<emphasis box="[1104,1255,1663,1684]" italics="true" pageId="6" pageNumber="422">Dicerorhinus</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Linnaeus" authorityYear="1758" box="[1315,1443,1663,1684]" class="Mammalia" family="Rhinocerotidae" genus="Rhinoceros" kingdom="Animalia" order="Perissodactyla" pageId="6" pageNumber="422" phylum="Chordata" rank="genus">
<emphasis box="[1315,1443,1663,1684]" italics="true" pageId="6" pageNumber="422">Rhinoceros</emphasis>
</taxonomicName>
in having no cement on cheek teeth, a protocone joined to the ectoloph on P2, a proximal border of the third metatarsal sigmoid in anterior view, and intermediate reliefs high and sharp on metapodials. Distinct from
<taxonomicName authorityName="Linnaeus" authorityYear="1758" box="[827,954,1816,1837]" class="Mammalia" family="Rhinocerotidae" genus="Rhinoceros" kingdom="Animalia" order="Perissodactyla" pageId="6" pageNumber="422" phylum="Chordata" rank="genus">
<emphasis box="[827,954,1816,1837]" italics="true" pageId="6" pageNumber="422">Rhinoceros</emphasis>
</taxonomicName>
in possessing a protocone and a hypocone equally developed on P2 and in having no anterior trochlear notch on the astragalus. Further differing from
<taxonomicName authorityName="Gloger" authorityYear="1841" box="[209,363,197,218]" class="Mammalia" family="Rhinocerotidae" genus="Dicerorhinus" kingdom="Animalia" order="Perissodactyla" pageId="7" pageNumber="423" phylum="Chordata" rank="genus">
<emphasis box="[209,363,197,218]" italics="true" pageId="7" pageNumber="423">Dicerorhinus</emphasis>
</taxonomicName>
in possessing a V-shaped lingual opening of the posterior valley on lower premolars.
</paragraph>
</subSubSection>
<subSubSection pageId="7" pageNumber="423" type="distribution">
<paragraph blockId="7.[145,761,290,404]" pageId="7" pageNumber="423">
<emphasis box="[145,617,290,312]" italics="true" pageId="7" pageNumber="423">Geographic and stratigraphic range:</emphasis>
Early and Middle Pleistocene of Luzon Island,
<collectingCountry box="[571,705,321,343]" name="Philippines" pageId="7" pageNumber="423">Philippines</collectingCountry>
, and of
<collectingCountry box="[173,342,352,374]" name="Taiwan Island" pageId="7" pageNumber="423">Taiwan Island</collectingCountry>
(
<bibRefCitation author="von Koenigswald" box="[360,631,352,374]" firstAuthor="von Koenigswald" pageId="7" pageNumber="423" pagination="1 - 14" refId="ref10231" refString="von Koenigswald GHR. 1956. Fossil mammals from the Philippines. Quezon City: National Research Council of the Philippines, 1 - 14." type="book chapter" year="1956">von Koenigswald, 1956</bibRefCitation>
;
<bibRefCitation author="Otsuka &amp; Lin" firstAuthor="Otsuka" pageId="7" pageNumber="423" pagination="1 - 35" refId="ref11145" refString="Otsuka H, Lin CC. 1984. Fossil rhinoceros from the T'ouk'oushan Group in Taiwan. Journal of Taiwan Museum 37: 1 - 35." type="journal article" year="1984">Otsuka &amp; Lin, 1984</bibRefCitation>
;
<bibRefCitation author="Ingicco" box="[264,475,382,404]" etAl="et al." firstAuthor="Ingicco" pageId="7" pageNumber="423" pagination="233 - 237" refId="ref9978" refString="Ingicco T, Van den Bergh GD, Jago-On C, Bahain JJ, Chacon MG, Amano N, Forestier H, King C, Manalo K, Nomade S, Pereira A, Reyes MC, Semah AM, Shao Q, Voinchet P, Falgueres C, Albers PCH, Lising M, Lyras G, Yurnaldi D, Rochette P, Bautista A, de Vos J. 2018. Earliest known hominin activity in the Philippines by 709 thousand years ago. Nature 557: 233 - 237." type="journal article" year="2018">
Ingicco
<emphasis box="[352,408,382,404]" italics="true" pageId="7" pageNumber="423">et al.</emphasis>
, 2018
</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="7" pageNumber="423" type="description">
<paragraph blockId="7.[145,762,445,1908]" pageId="7" pageNumber="423">
<emphasis box="[145,298,446,467]" italics="true" pageId="7" pageNumber="423">Description:</emphasis>
See morpho-anatomical characters in Supporting Information, Text S3. Even if nasal or frontal bones are not recognized in the current hypodigm of
<taxonomicName baseAuthorityName="von Koenigswald" baseAuthorityYear="1956" box="[309,523,537,559]" class="Mammalia" family="Rhinocerotidae" genus="Nesorhinus" kingdom="Animalia" order="Perissodactyla" pageId="7" pageNumber="423" phylum="Chordata" rank="species" species="philippinensis">
<emphasis box="[309,523,537,559]" italics="true" pageId="7" pageNumber="423">N. philippinensis</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Antoine &amp; Reyes &amp; Amano &amp; Bautista &amp; Chang &amp; Claude &amp; De Vos &amp; Ingicco" authorityYear="2022" baseAuthorityName="Otsuka &amp; Lin" baseAuthorityYear="1984" box="[591,753,537,559]" class="Mammalia" family="Rhinocerotidae" genus="Nesorhinus" kingdom="Animalia" order="Perissodactyla" pageId="7" pageNumber="423" phylum="Chordata" rank="species" species="hayasakai">
<emphasis box="[591,753,537,559]" italics="true" pageId="7" pageNumber="423">N. hayasakai</emphasis>
</taxonomicName>
,
<taxonomicName authorityName="Antoine &amp; Reyes &amp; Amano &amp; Bautista &amp; Chang &amp; Claude &amp; De Vos &amp; Ingicco" authorityYear="2022" box="[145,279,568,589]" class="Mammalia" family="Rhinocerotidae" genus="Nesorhinus" kingdom="Animalia" order="Perissodactyla" pageId="7" pageNumber="423" phylum="Chordata" rank="genus">
<emphasis box="[145,279,568,589]" italics="true" pageId="7" pageNumber="423">Nesorhinus</emphasis>
</taxonomicName>
was most probably one-horned, i.e. with a nasal horn and no frontal horn, as inferred by the topology of the consensus tree: this is the ancestral condition in
<taxonomicName authorityName="GRAY" authorityYear="1821" box="[291,455,660,682]" class="Mammalia" family="Rhinocerotidae" kingdom="Animalia" order="Perissodactyla" pageId="7" pageNumber="423" phylum="Chordata" rank="tribe" tribe="Rhinocerotina">Rhinocerotina</taxonomicName>
and Rhinoceroti, retained in
<taxonomicName authorityName="Linnaeus" authorityYear="1758" box="[178,309,690,711]" class="Mammalia" family="Rhinocerotidae" genus="Rhinoceros" kingdom="Animalia" order="Perissodactyla" pageId="7" pageNumber="423" phylum="Chordata" rank="genus">
<emphasis box="[178,309,690,711]" italics="true" pageId="7" pageNumber="423">Rhinoceros</emphasis>
</taxonomicName>
. According to the most-parsimonious topologies, a frontal horn was acquired independently in Diceroti and
<taxonomicName baseAuthorityName="Fischer" baseAuthorityYear="1814" box="[317,608,752,773]" class="Mammalia" family="Rhinocerotidae" genus="Dicerorhinus" kingdom="Animalia" order="Perissodactyla" pageId="7" pageNumber="423" phylum="Chordata" rank="species" species="sumatrensis">
<emphasis box="[317,608,752,773]" italics="true" pageId="7" pageNumber="423">Dicerorhinus sumatrensis</emphasis>
</taxonomicName>
(and perhaps also in
<taxonomicName box="[229,375,782,804]" class="Mammalia" family="Rhinocerotidae" genus="Rhinoceros" kingdom="Animalia" order="Perissodactyla" pageId="7" pageNumber="423" phylum="Chordata" rank="species" species="fusuiensis">
<emphasis box="[229,375,782,804]" italics="true" pageId="7" pageNumber="423">D. fusuiensis</emphasis>
</taxonomicName>
), which is in full agreement with the most recent genomic phylogenies.
</paragraph>
<paragraph blockId="7.[145,762,445,1908]" lastBlockId="7.[809,1425,197,587]" pageId="7" pageNumber="423">
Body mass, as predicted from regressions on upper and lower teeth, but also on limb bones, with consistent results, is estimated at
<quantity box="[427,577,905,927]" metricMagnitude="3" metricUnit="kg" metricValue="1.334" metricValueMax="1.67" metricValueMin="0.998" pageId="7" pageNumber="423" unit="kg" value="1334.0" valueMax="1670.0" valueMin="998.0">9981670 kg</quantity>
for
<taxonomicName authorityName="Antoine &amp; Reyes &amp; Amano &amp; Bautista &amp; Chang &amp; Claude &amp; De Vos &amp; Ingicco" authorityYear="2022" box="[627,761,905,926]" class="Mammalia" family="Rhinocerotidae" genus="Nesorhinus" kingdom="Animalia" order="Perissodactyla" pageId="7" pageNumber="423" phylum="Chordata" rank="genus">
<emphasis box="[627,761,905,926]" italics="true" pageId="7" pageNumber="423">Nesorhinus</emphasis>
</taxonomicName>
(Supporting Information, Table S5).
<taxonomicName baseAuthorityName="von Koenigswald" baseAuthorityYear="1956" class="Mammalia" family="Rhinocerotidae" genus="Nesorhinus" kingdom="Animalia" order="Perissodactyla" pageId="7" pageNumber="423" phylum="Chordata" rank="species" species="philippinensis">
<emphasis italics="true" pageId="7" pageNumber="423">Nesorhinus philippinensis</emphasis>
</taxonomicName>
, documented at the
<collectingRegion box="[551,643,966,988]" country="Philippines" name="Kalinga" pageId="7" pageNumber="423">Kalinga</collectingRegion>
site by at least two individuals similar in size, was the smallest and lightest species. Body weight ranged between 1025 and
<quantity box="[192,280,1058,1080]" metricMagnitude="3" metricUnit="kg" metricValue="1.185" pageId="7" pageNumber="423" unit="kg" value="1185.0">1185 kg</quantity>
(mean:
<quantity box="[366,454,1058,1080]" metricMagnitude="3" metricUnit="kg" metricValue="1.103" pageId="7" pageNumber="423" unit="kg" value="1103.0">1103 kg</quantity>
) based on dental predictors and
<quantity box="[194,337,1089,1111]" metricMagnitude="3" metricUnit="kg" metricValue="1.069" metricValueMax="1.14" metricValueMin="0.998" pageId="7" pageNumber="423" unit="kg" value="1069.0" valueMax="1140.0" valueMin="998.0">9981140 kg</quantity>
(mean:
<quantity box="[426,515,1089,1111]" metricMagnitude="3" metricUnit="kg" metricValue="1.069" pageId="7" pageNumber="423" unit="kg" value="1069.0">1069 kg</quantity>
) based on postcranial predictors, which falls between the known ranges of the Sumatran and Javan rhinos, the smallest of the extant rhinos.
<taxonomicName authorityName="Antoine &amp; Reyes &amp; Amano &amp; Bautista &amp; Chang &amp; Claude &amp; De Vos &amp; Ingicco" authorityYear="2022" baseAuthorityName="Otsuka &amp; Lin" baseAuthorityYear="1984" box="[318,580,1181,1202]" class="Mammalia" family="Rhinocerotidae" genus="Nesorhinus" kingdom="Animalia" order="Perissodactyla" pageId="7" pageNumber="423" phylum="Chordata" rank="species" species="hayasakai">
<emphasis box="[318,580,1181,1202]" italics="true" pageId="7" pageNumber="423">Nesorhinus hayasakai</emphasis>
</taxonomicName>
was somewhat heavier, with more variable weight estimates based on teeth (mean:
<quantity box="[291,381,1242,1264]" metricMagnitude="3" metricUnit="kg" metricValue="1.263" pageId="7" pageNumber="423" unit="kg" value="1263.0">1263 kg</quantity>
; range:
<quantity box="[466,624,1242,1264]" metricMagnitude="3" metricUnit="kg" metricValue="1.344" metricValueMax="1.67" metricValueMin="1.018" pageId="7" pageNumber="423" unit="kg" value="1344.0" valueMax="1670.0" valueMin="1018.0">10181670 kg</quantity>
; Supporting Information, Table S3), and a slightly heavier estimate based on radius (
<quantity box="[354,453,1304,1326]" metricMagnitude="3" metricUnit="kg" metricValue="1.3059999999999998" pageId="7" pageNumber="423" unit="kg" value="1306.0">1306 kg</quantity>
; Supporting Information, Table S5). A shoulder height of
<emphasis box="[532,543,1335,1356]" italics="true" pageId="7" pageNumber="423">c</emphasis>
.
<quantity box="[559,706,1334,1356]" metricMagnitude="0" metricUnit="m" metricValue="1.265" metricValueMax="1.3" metricValueMin="1.23" pageId="7" pageNumber="423" unit="m" value="1.265" valueMax="1.3" valueMin="1.23">1.231.30 m</quantity>
was estimated for
<taxonomicName baseAuthorityName="von Koenigswald" baseAuthorityYear="1956" box="[312,513,1365,1387]" class="Mammalia" family="Rhinocerotidae" genus="Nesorhinus" kingdom="Animalia" order="Perissodactyla" pageId="7" pageNumber="423" phylum="Chordata" rank="species" species="philippinensis">
<emphasis box="[312,513,1365,1387]" italics="true" pageId="7" pageNumber="423">N. philippinensis</emphasis>
</taxonomicName>
through comparison of forelimb dimensions with recent rhinos (mean:
<quantity metricMagnitude="0" metricUnit="m" metricValue="1.26" pageId="7" pageNumber="423" unit="m" value="1.26">1.26 m</quantity>
). This estimate is similar to the smallest Javanese rhino individuals and to average Sumatran rhino individuals (Supporting Information, Tables S6, S7). A marginally higher stature (
<emphasis box="[485,496,1519,1540]" italics="true" pageId="7" pageNumber="423">c</emphasis>
.
<quantity box="[509,588,1518,1540]" metricMagnitude="0" metricUnit="m" metricValue="1.31" pageId="7" pageNumber="423" unit="m" value="1.31">1.31 m</quantity>
) is inferred for
<taxonomicName authorityName="Antoine &amp; Reyes &amp; Amano &amp; Bautista &amp; Chang &amp; Claude &amp; De Vos &amp; Ingicco" authorityYear="2022" baseAuthorityName="Otsuka &amp; Lin" baseAuthorityYear="1984" box="[145,295,1549,1571]" class="Mammalia" family="Rhinocerotidae" genus="Nesorhinus" kingdom="Animalia" order="Perissodactyla" pageId="7" pageNumber="423" phylum="Chordata" rank="species" species="hayasakai">
<emphasis box="[145,295,1549,1571]" italics="true" pageId="7" pageNumber="423">N. hayasakai</emphasis>
</taxonomicName>
(see Supporting Information, Tables S6, S7). Nevertheless, comparison of skeletal proportions shows that
<taxonomicName baseAuthorityName="von Koenigswald" baseAuthorityYear="1956" box="[303,529,1610,1632]" class="Mammalia" family="Rhinocerotidae" genus="Nesorhinus" kingdom="Animalia" order="Perissodactyla" pageId="7" pageNumber="423" phylum="Chordata" rank="species" species="philippinensis">
<emphasis box="[303,529,1610,1632]" italics="true" pageId="7" pageNumber="423">N. philippinensis</emphasis>
</taxonomicName>
was particularly slender-limbed, with the notable exception of the scapula and the metapodials. Its gracility indices are closely similar to those of the most gracile living rhinoceros, i.e.
<taxonomicName baseAuthorityName="Fischer" baseAuthorityYear="1814" box="[319,615,1733,1755]" class="Mammalia" family="Rhinocerotidae" genus="Dicerorhinus" kingdom="Animalia" order="Perissodactyla" pageId="7" pageNumber="423" phylum="Chordata" rank="species" species="sumatrensis">
<emphasis box="[319,615,1733,1755]" italics="true" pageId="7" pageNumber="423">Dicerorhinus sumatrensis</emphasis>
</taxonomicName>
(Supporting Information, Table S8; Figs S5, S6). Within
<taxonomicName authorityName="Antoine &amp; Reyes &amp; Amano &amp; Bautista &amp; Chang &amp; Claude &amp; De Vos &amp; Ingicco" authorityYear="2022" box="[625,755,1764,1785]" class="Mammalia" family="Rhinocerotidae" genus="Nesorhinus" kingdom="Animalia" order="Perissodactyla" pageId="7" pageNumber="423" phylum="Chordata" rank="genus">
<emphasis box="[625,755,1764,1785]" italics="true" pageId="7" pageNumber="423">Nesorhinus</emphasis>
</taxonomicName>
,
<taxonomicName baseAuthorityName="von Koenigswald" baseAuthorityYear="1956" box="[145,341,1794,1816]" class="Mammalia" family="Rhinocerotidae" genus="Nesorhinus" kingdom="Animalia" order="Perissodactyla" pageId="7" pageNumber="423" phylum="Chordata" rank="species" species="philippinensis">
<emphasis box="[145,341,1794,1816]" italics="true" pageId="7" pageNumber="423">N. philippinensis</emphasis>
</taxonomicName>
was also much more slender-limbed than
<taxonomicName authorityName="Antoine &amp; Reyes &amp; Amano &amp; Bautista &amp; Chang &amp; Claude &amp; De Vos &amp; Ingicco" authorityYear="2022" baseAuthorityName="Otsuka &amp; Lin" baseAuthorityYear="1984" box="[208,363,1825,1847]" class="Mammalia" family="Rhinocerotidae" genus="Nesorhinus" kingdom="Animalia" order="Perissodactyla" pageId="7" pageNumber="423" phylum="Chordata" rank="species" species="hayasakai">
<emphasis box="[208,363,1825,1847]" italics="true" pageId="7" pageNumber="423">N. hayasakai</emphasis>
</taxonomicName>
, which is further consistent with significantly lighter body mass estimates based on radius (998 vs.
<quantity box="[341,441,1886,1908]" metricMagnitude="3" metricUnit="kg" metricValue="1.3059999999999998" pageId="7" pageNumber="423" unit="kg" value="1306.0">1306 kg</quantity>
, respectively; Supporting Information, Table S5). This discrepancy may be related either to interindividual variability [e.g. sexual dimorphism although it seems to be exaggerated for a rhinocerotine (
<bibRefCitation author="Guerin" box="[1042,1194,289,311]" firstAuthor="Guerin" pageId="7" pageNumber="423" refId="ref9568" refString="Guerin C. 1980. Les rhinoceros (Mammalia, Perissodactyla) du Miocene terminal au Pleistocene superieur en Europe occidentale: comparaison avec les especes actuelles. Villeurbanne: Departement des sciences de la terre, Universite Claude-Bernard Lyon 1." type="journal volume" year="1980">Guérin, 1980</bibRefCitation>
)], or to a secondary adaptation to the unbalanced insular environment of the
<collectingCountry box="[856,992,350,372]" name="Philippines" pageId="7" pageNumber="423">Philippines</collectingCountry>
(Supporting Information, Table S8) with respect to mainland assemblages. Strikingly, the scapula of
<taxonomicName baseAuthorityName="von Koenigswald" baseAuthorityYear="1956" box="[940,1146,411,433]" class="Mammalia" family="Rhinocerotidae" genus="Nesorhinus" kingdom="Animalia" order="Perissodactyla" pageId="7" pageNumber="423" phylum="Chordata" rank="species" species="philippinensis">
<emphasis box="[940,1146,411,433]" italics="true" pageId="7" pageNumber="423">N. philippinensis</emphasis>
</taxonomicName>
is neither particularly spatulate nor elongated. Its gracility equals that of
<taxonomicName baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[809,998,473,494]" class="Mammalia" family="Rhinocerotidae" genus="Diceros" kingdom="Animalia" order="Perissodactyla" pageId="7" pageNumber="423" phylum="Chordata" rank="species" species="bicornis">
<emphasis box="[809,998,473,494]" italics="true" pageId="7" pageNumber="423">Diceros bicornis</emphasis>
</taxonomicName>
and it is intermediate to the living
<taxonomicName box="[809,1032,503,525]" pageId="7" pageNumber="423">
<emphasis box="[809,939,503,524]" italics="true" pageId="7" pageNumber="423">Rhinoceros</emphasis>
species
</taxonomicName>
, while
<taxonomicName baseAuthorityName="Fischer" baseAuthorityYear="1814" box="[1119,1425,503,525]" class="Mammalia" family="Rhinocerotidae" genus="Dicerorhinus" kingdom="Animalia" order="Perissodactyla" pageId="7" pageNumber="423" phylum="Chordata" rank="species" species="sumatrensis">
<emphasis box="[1119,1425,503,525]" italics="true" pageId="7" pageNumber="423">Dicerorhinus sumatrensis</emphasis>
</taxonomicName>
has by far the most robust scapula (Supporting Information, Table S8).
</paragraph>
</subSubSection>
</treatment>
</document>