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<mods:title id="FB005DBFC6FF35A52FA6B221039FD318">The Middle Pleistocene vertebrate fauna from Khok Sung (Nakhon Ratchasima, Thailand): biochronological and paleobiogeographical implications</mods:title>
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<mods:namePart id="E9B6A356E01E4C7D944CBB3560A3454E">Suraprasit, Kantapon</mods:namePart>
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<mods:namePart id="D89E5A03D4D06FB7556DCD3B3754BA00">Jaeger, Jean-Jacques</mods:namePart>
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<mods:namePart id="22E3B5E1A8E5FDCD0882EFF62D73D15B">Chaimanee, Yaowalak</mods:namePart>
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<mods:namePart id="91DF6CD70CAC178A1EDFF2FCB43F1714">Yamee, Chotima</mods:namePart>
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<mods:namePart id="16B072B59538B0BE8B086A6A8318CC8F">Tian, Pannipa</mods:namePart>
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<mods:namePart id="2D7BB3E665E214A3F38A56AA50F98EE0">Panha, Somsak</mods:namePart>
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<paragraph id="EB76B682E3EB6612F2F89D5B86A7A4E9" pageId="42" pageNumber="43">Taxon classification Animalia Artiodactyla Bovidae</paragraph>
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<subSubSection id="944244D0E22E07743489DE33FE8A68CC" pageId="42" pageNumber="43" type="nomenclature">
<paragraph id="9B109E78084CA6CBB23DB04E8F0095E7" pageId="42" pageNumber="43">
<taxonomicName id="5DDEAD875F92999B2AC6540FD000E5AF" ID-CoL="MLPV" authority="Hamilton-Smith, 1827" authorityName="Hamilton-Smith" authorityYear="1827" class="Mammalia" family="Bovidae" genus="Bos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bos gaurus" order="Artiodactyla" pageId="42" pageNumber="43" phylum="Chordata" rank="species" species="gaurus">Bos gaurus (Hamilton-Smith, 1827)</taxonomicName>
</paragraph>
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<subSubSection id="7F7CCFF879EDA9478F9C8186ABDECDA9" pageId="42" pageNumber="43" type="materials_examined">
<paragraph id="5FCC09771C640F8F77606C179A7A7D86" pageId="42" pageNumber="43">Referred material.</paragraph>
<paragraph id="BBCF6B491137F23EC5025321497B9157" pageId="42" pageNumber="43">A left horn core, DMR-KS-05-03-26-22; a right DP2, DMR-KS-05-03-20-4; two right P2-DMR-KS-05-03-19-27 and DMR-KS-05-04-03-3; a right DP3, DMR-KS-05-03-20-3; a right DP4, DMR-KS-05-03-17-3; a right M1, DMR-KS-05-03-00-20; a right M3, DMR-KS-05-03-17-1; a right mandible with m1-m3, DMR-KS-05-03-00-1; a left mandible with p2-m3, DMR-KS-05-04-3-1; a left i1, DMR-KS-05-03-00-27; two left m2-DMR-KS-05-03-19-26 and DMR-KS-05-03-16-1; two humeri-DMR-KS-05-05-1-1 (right) and DMR-KS-05-03-00-62 (left); a right metacarpus, DMR-KS-05-03-26-27; two left femora-DMR-KS-05-03-9-2 and DMR-KS-05-04-30-1 (proximal part).</paragraph>
</subSubSection>
<subSubSection id="178FE14294E40EDB0BF4A7995EB56084" lastPageId="44" lastPageNumber="45" pageId="42" pageNumber="43" type="materials_examined">
<paragraph id="4EF7122CD876734474B7A722197BA2AC" pageId="42" pageNumber="43">Material description.</paragraph>
<paragraph id="0D192F2C119DFB29059333F4A0C1C339" pageId="42" pageNumber="43">Horn core: a single horn core (DMR-KS-05-03-26-22) is small, curved upward (Fig. 28A, B) and slightly backward. The horn core base is oval in cross-section (Fig. 28A). A longitudinal ridge on the anterior surface of the horn core is present (Fig. 28B). This specimen belongs to a juvenile individual according to its very small size.</paragraph>
<caption id="E98EA959FE1862FFBFDCA56B5B0F84EC" pageId="42" pageNumber="43">
<paragraph id="D8EBC2925775355CDED026516D48DA56" pageId="42" pageNumber="43">
Figure 28. Remains of
<taxonomicName id="DD3F2E7AF42317D6C50675DBBD4C184E" class="Mammalia" family="Bovidae" genus="Bos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bos gaurus" order="Artiodactyla" pageId="42" pageNumber="43" phylum="Chordata" rank="species" species="gaurus">Bos gaurus</taxonomicName>
from Khok Sung:
<normalizedToken id="6D6F3FD444A7A42E2B3741B0A00E0B17" originalValue="AB">A-B</normalizedToken>
DMR-KS-05-03-26-22, a left horn core in posterior (A) and anterior (B) view C DMR-KS-05-03-20-1, a right DP2 D DMR-KS-05-03-19-27, a right P2 E DMR-KS-05-04-03-03, a right P2 F DMR-KS-05-03-20-3, a right DP3
<normalizedToken id="8BA9D5374E74E20A5E215D80FB795E4B" originalValue="GHDMR-">G-HDMR-</normalizedToken>
05-03-17-3, a right DP4 in occlusal (G) and lingual (H) views IDMR-05-03-17-1, a right M3 JDMR-05-03-19-26, a left m2
<normalizedToken id="3A3ABCA7467E73C014B3B5C69AE2837C" originalValue="KL">K-L</normalizedToken>
DMR-KS-05-04-3-1, a left mandible in and occlusal (K) and lateral (L) views M DMR-KS-05-03-00-1, a fragmentary mandible in occlusal view. All isolated teeth are shown in occlusal view.
</paragraph>
</caption>
<paragraph id="5846C01BEF4FDEE3F1A73B015F7AC5D6" lastPageId="44" lastPageNumber="45" pageId="42" pageNumber="43">
Upper dentition: DP2 (DMR-KS-05-03-20-4) is small and elongated, characterized by three main cones (anterior cone, paracone, and metacone) and a well-developed metastyle (Fig. 28C) (for measurements, see Tab. 15). The anterior and poste
<pageBreakToken id="BDAF513FAF1C9C2ACC62B35D6AB15A89" pageId="43" pageNumber="44" start="start">rior</pageBreakToken>
fossettes fuse together. Two P2 (DMR-KS-05-03-19-27; Fig. 28D and DMR-KS-05-04-03-3: Fig. 28E) have a well developed paracone rib close to the parastyle and a nearly flat lingual wall. The fossettes are separated into two islands (larger for the anterior one) due to the heavy wear stage (Fig. 28D). The P2 shows a nearly straight posterior wall and is wider than the DP2 (Fig. 28E). On the molarized DP3, the posterior lobe is broader than the anterior lobe (Fig. 28F). A small medial fossette is present. The entostyle is short and projects posteriorly. The molarized DP4 (DMR-KS-05-03-17-3)
<pageBreakToken id="DA926961041CCB68C2471E5B2D197625" pageId="44" pageNumber="45" start="start">is</pageBreakToken>
slightly worn, characterized by a rectangular outline, well-developed buccal styles, an unfused entostyle, and two separated medial fossette (Fig. 28
<normalizedToken id="4A038C31C4CAA6ED249B177C6E107DD4" originalValue="GH">G-H</normalizedToken>
). The entostyle is bifurcated and situated between the protocone and hypocone (Fig. 28G). Two parallel longitudinal grooves are present along the lingual surface of the enstostyle, likely resulting in a trifurcated pattern in relation to the middle wear stage (Fig. 28H). The heavily worn M1 (DMR-KS-05-03-00-20) displays a subsquare outline and an unbifurcated entostyle positioned between the protocone and hypocone (Fig. 28I). The medial fossette is absent due to the heavy wear stage. The M3 (DMR-KS-05-03-17-1) exhibits well-developed buccal styles and large medial fossettes splitting into 2 islands with wear (Fig. 28J). The entostyle on the M3 is short, not bifurcated, and close to the hypocone.
</paragraph>
<paragraph id="27503D5718F25929328EF7C0E5AAC1C1" pageId="44" pageNumber="45">
Mandibles and lower dentition: DMR-KS-05-04-3-1 is complete, posterior to the p2, with the exception of a small part of the angular region (Fig. 28K, L) (for measurements, see Appendix 13). Another mandible (DMR-KS-05-03-00-1) preserves only a portion of the ramus with the complete molar row (Fig. 28M and Appendix 13). The isolated i1 (DMR-KS-05-03-00-27) is heavily worn, spatulate, and robust. Lower premolars have well-developed main cuspids and cristids (Fig. 28K, M). On the p2, the protocone is the highest cuspid and the posterior fossette is present. The p3 is elongated as long as the p4. The premetacristid is poorly developed. The postprotocristid on the p3 is larger than that on the p4. On the p4, the postprotocristid is narrow and anteroposteriorly constricted. The metaconid is most developed, compared to
<taxonomicName id="43A2847E95D2FB85FE06C65515CE2484" class="Mammalia" family="Bovidae" genus="Bos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bos sauveli" order="Artiodactyla" pageId="44" pageNumber="45" phylum="Chordata" rank="species" species="sauveli">Bos sauveli</taxonomicName>
and
<taxonomicName id="AE102951416E1CA198F1ABC4DF73E6FB" class="Mammalia" family="Bovidae" genus="Bos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bos javanicus" order="Artiodactyla" pageId="44" pageNumber="45" phylum="Chordata" rank="species" species="javanicus">Bos javanicus</taxonomicName>
as well as
<taxonomicName id="75C57124BB9D596272EF2C3787ECBAD0" class="Mammalia" family="Bovidae" genus="Bubalus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bubalus arnee" order="Artiodactyla" pageId="44" pageNumber="45" phylum="Chordata" rank="species" species="arnee">Bubalus arnee</taxonomicName>
. For all lower molars, the ectostylid is slightly developed and not bifurcated (Fig. 28K,
<normalizedToken id="EA102B1EC822406CE7A07FB0A902DA76" originalValue="MN">M-N</normalizedToken>
) (for measurements, see Tab. 15). In lingual view, the metastylid is absent at the medium wear stage (Fig. 28K, M). In occlusal view, the entostylid is straight and short. The buccal outline of the protoconid and hypoconid is U-shaped in relation to the strong wear (Fig. 28M). The posterior talonid on the m3 is well-developed. The posthypoconulidcristid protrudes posteriorly.
</paragraph>
<paragraph id="52AB230D5B3A64E4D082E635A990555C" pageId="44" pageNumber="45">
Postcranial remains: postcranial elements include humeri (Fig. 29
<normalizedToken id="CBDB43144AAAFF07D852141DAD85784A" originalValue="AD">A-D</normalizedToken>
), a metacarpus (Fig. 29
<normalizedToken id="1BA2756657BF71A0F142B27BCCA01DBF" originalValue="EG">E-G</normalizedToken>
), and femora (Fig. 29
<normalizedToken id="79D1C997A030E5CC9BE12B787CD84181" originalValue="HJ">H-J</normalizedToken>
) (for measurements, see Appendix 1). The femur DMR-KS-05-04-30-1 lacks a distal portion. We assign these postcranial bones based on the proportional correlations with the recent specimens of
<taxonomicName id="E6A3B602086B2CC11604BA97E58AB6A5" class="Mammalia" family="Bovidae" genus="Bos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bos gaurus" order="Artiodactyla" pageId="44" pageNumber="45" phylum="Chordata" rank="species" species="gaurus">Bos gaurus</taxonomicName>
(Tab. 13 and Appendices 7 and 9-12).
</paragraph>
<caption id="9449281C878622B479AC2977B7BEBF90" pageId="44" pageNumber="45">
<paragraph id="6F56C02D18E661AA0312EA2431B9E3A8" pageId="44" pageNumber="45">
Figure 29. Postcranial remains of
<taxonomicName id="450F3349DF0E1B7A6CD336A95740DE89" class="Mammalia" family="Bovidae" genus="Bos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bos gaurus" order="Artiodactyla" pageId="44" pageNumber="45" phylum="Chordata" rank="species" species="gaurus">Bos gaurus</taxonomicName>
from Khok Sung:
<normalizedToken id="AFFE0445F72B6D83A426AAE2C56BD0B0" originalValue="AD">A-D</normalizedToken>
DMR-KS-05-05-1-1, a right humerus in proximal (A), posterior (B), anterior (C), and distal (D) views
<normalizedToken id="2587E05E450CB3850274EC6B561A1659" originalValue="EG">E-G</normalizedToken>
DMR-KS-05-03-26-27, a right metacarpus in proximal (E), anterior (F), and distal (G) views
<normalizedToken id="2EED070A39B3B86F0E7476BA56D954DE" originalValue="HJ">H-J</normalizedToken>
DMR-KS-05-03-9-2, a left femur in proximal (H), distal (I), and anterior (J) views.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="80F30190E31A13A9699003CC95C1AB03" lastPageId="45" lastPageNumber="46" pageId="44" pageNumber="45" type="remarks">
<paragraph id="9B6BECB65F1E6435228CB63249CB8B54" pageId="44" pageNumber="45">Taxonomic remarks and comparisons.</paragraph>
<paragraph id="8D66B38A773C45A1B0679A9F7E375C85" pageId="44" pageNumber="45">
According to
<bibRefCitation id="B9300AAE83890F4A5AF4F7C5B41E793A" author="Indriati, E" journalOrPublisher="PLoS ONE" pageId="83" pageNumber="84" title="The IUCN Red List of Threatened Species, version 2015.1" url="http://www.iucnredlist.org" year="2015">IUCN (2015)</bibRefCitation>
, the wild forms of gaurs are considered as
<taxonomicName id="D2BD7928D0D3650CDF8738336D90D9CC" class="Mammalia" family="Bovidae" genus="Bos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bos gaurus" order="Artiodactyla" pageId="44" pageNumber="45" phylum="Chordata" rank="species" species="gaurus">Bos gaurus</taxonomicName>
, while their domestic forms are recognized as
<taxonomicName id="81AA3C66D98FDD0124882A2AB2D95643" class="Mammalia" family="Bovidae" genus="Bos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bos frontalis" order="Artiodactyla" pageId="44" pageNumber="45" phylum="Chordata" rank="species" species="frontalis">Bos frontalis</taxonomicName>
(
<bibRefCitation id="54CD23EF690F975E086C22745C2D2AA7" author="Gentry, A" journalOrPublisher="Journal of Archaeological Science" pageId="81" pageNumber="82" pagination="645 - 651" title="The naming of wild animal species and their domestic derivatives." url="10.1016/j.jas.2003.10.006" volume="31" year="2004">Gentry et al. 2004</bibRefCitation>
). We consider here the Pleistocene fossil gaurs as belonging to wild forms in terms of taxonomic nomenclature.
</paragraph>
<paragraph id="5C3A32741A09B80DCFA7EB4900B6D57E" pageId="44" pageNumber="45">
We assign the juvenile horn core (DMR-KS-05-03-26-22) to
<taxonomicName id="0E458FB7D4535082DE7D71E6A916C511" class="Mammalia" family="Bovidae" genus="Bos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bos gaurus" order="Artiodactyla" pageId="44" pageNumber="45" phylum="Chordata" rank="species" species="gaurus">Bos gaurus</taxonomicName>
because the horn cores of gaurs are different from all other
<taxonomicName id="FA40B2B0D65CD56647D858331478071A" class="Mammalia" family="Bovidae" genus="Bos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bos" order="Artiodactyla" pageId="44" pageNumber="45" phylum="Chordata" rank="genus">Bos</taxonomicName>
species. They grow outward and curve upward, similar to those of
<taxonomicName id="44F3A251F7B629F7046F44DB461D4499" class="Mammalia" family="Bovidae" genus="Bubalus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bubalus arnee" order="Artiodactyla" pageId="44" pageNumber="45" phylum="Chordata" rank="species" species="arnee">Bubalus arnee</taxonomicName>
, but their apical portion curves inward and slightly forward (
<bibRefCitation id="DB0EF06DB86A253F27739636E483B3FF" author="Lekagul, B" journalOrPublisher="Association for the Conservation of Wildlife, Bangkok" pageId="83" pageNumber="84" title="Mammals of Thailand." year="1988">Lekagul and McNeely 1988</bibRefCitation>
).
</paragraph>
<paragraph id="6849D233A1684AC2405630397314B41E" lastPageId="45" lastPageNumber="46" pageId="44" pageNumber="45">
Mandibles and isolated teeth of
<taxonomicName id="FB733F8B3CD8920BF3072D490B88B629" class="Mammalia" family="Bovidae" genus="Bos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bos gaurus" order="Artiodactyla" pageId="44" pageNumber="45" phylum="Chordata" rank="species" species="gaurus">Bos gaurus</taxonomicName>
are also observed. The cheek teeth of
<taxonomicName id="845D3A809283B03CC97DAE05D9EC81B9" class="Mammalia" family="Bovidae" genus="Bos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bos gaurus" order="Artiodactyla" pageId="44" pageNumber="45" phylum="Chordata" rank="species" species="gaurus">Bos gaurus</taxonomicName>
are distinguished from
<taxonomicName id="250CA8D31E2FA0A81B3F6F8E1AA443D3" class="Mammalia" family="Bovidae" genus="Bos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bos sauveli" order="Artiodactyla" pageId="44" pageNumber="45" phylum="Chordata" rank="species" species="sauveli">Bos sauveli</taxonomicName>
and
<taxonomicName id="60947BC052038B3F9A269342A1AD2558" class="Mammalia" family="Bovidae" genus="Bos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bos javanicus" order="Artiodactyla" pageId="44" pageNumber="45" phylum="Chordata" rank="species" species="javanicus">Bos javanicus</taxonomicName>
by having two separate fossettes on the P2, more developed metaconids and more anteroposteriorly constricted
<pageBreakToken id="3D753F22437D008CADB7FAC632C31843" pageId="45" pageNumber="46" start="start">postprotocristids</pageBreakToken>
on the p3 and p4, and more robust cheek teeth (Figs 26 and 27, and Tab. 15). The entostyles are usually bifurcated or sometimes trifurcated on the slightly to moderately worn upper molars (our observations on the comparative material of recent
<taxonomicName id="9A1FE578240FDE310E5DFCD9C1D0DB7F" class="Mammalia" family="Bovidae" genus="Bos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bos gaurus" order="Artiodactyla" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="gaurus">Bos gaurus</taxonomicName>
: e.g., ZSM-1972-5 and ZSM-1961-313), similar to those of
<taxonomicName id="3FAE05A192AB90FFCE5A3FE548D17824" class="Mammalia" family="Bovidae" genus="Bos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bos javanicus" order="Artiodactyla" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="javanicus">Bos javanicus</taxonomicName>
. But the entostyle is not bifurcated, when the molar is extremely worn, as seen on the specimen DMR-KS-05-03-00-20 (Fig. 28I). This character is therefore morphologically variable through wear. On the m3, the entostylid and posterior talonid in
<taxonomicName id="F62671DAB466ED38482F977837F239A3" class="Mammalia" family="Bovidae" genus="Bos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bos gaurus" order="Artiodactyla" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="gaurus">Bos gaurus</taxonomicName>
is almost more developed than that in
<taxonomicName id="4E9BEB939F861E151553544AB719749F" class="Mammalia" family="Bovidae" genus="Bos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bos javanicus" order="Artiodactyla" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="javanicus">Bos javanicus</taxonomicName>
. The angle between the posthypocristid and prehypoconulidcristid is slightly more divergent in
<taxonomicName id="546FD693880265C2A14A056B13EB3E9C" class="Mammalia" family="Bovidae" genus="Bos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bos sauveli" order="Artiodactyla" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="sauveli">Bos sauveli</taxonomicName>
than in
<taxonomicName id="213B21508FC72B7144B3F7EBAD087A3B" class="Mammalia" family="Bovidae" genus="Bos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bos gaurus" order="Artiodactyla" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="gaurus">Bos gaurus</taxonomicName>
. The size of Khok Sung
<taxonomicName id="D1B3795B932D82AE6BC6F112AC00B340" class="Mammalia" family="Bovidae" genus="Bos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bos gaurus" order="Artiodactyla" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="gaurus">Bos gaurus</taxonomicName>
falls within the range of the recent population (Figs 26 and 27, and Tab. 14). We elucidate here the co-occurrence of two
<taxonomicName id="0A7A12E510A5E315D7474A318D026176" class="Mammalia" family="Bovidae" genus="Bos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bos" order="Artiodactyla" pageId="45" pageNumber="46" phylum="Chordata" rank="genus">Bos</taxonomicName>
species,
<taxonomicName id="6342B0D4AD977B87296F1E124525E8D7" class="Mammalia" family="Bovidae" genus="Bos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bos sauveli" order="Artiodactyla" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="sauveli">Bos sauveli</taxonomicName>
and
<taxonomicName id="160A26C610CBA081B70D12C60154668C" class="Mammalia" family="Bovidae" genus="Bos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bos gaurus" order="Artiodactyla" pageId="45" pageNumber="46" phylum="Chordata" rank="species" species="gaurus">Bos gaurus</taxonomicName>
(larger), in Khok Sung.
</paragraph>
</subSubSection>
</treatment>
</document>