1572 lines
232 KiB
XML
1572 lines
232 KiB
XML
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<mods:title id="B60DF947B6675B2557BAF96830702FF1">New material of Parabrachyodus hyopotamoides from Samane Nala, Bugti Hills (Pakistan) and the origin of Merycopotamini (Mammalia: Hippopotamoidea)</mods:title>
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<mods:namePart id="5BCD7047B543ABD92C7DB956A262CB9B">Gernelle, Killian</mods:namePart>
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<mods:namePart id="4F3D291863DBB16C2B7E719F68485B1B">Boisserie, Jean-Renaud</mods:namePart>
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<mods:namePart id="79FF106F79B580052A3B1CFFBABD4A14">Marivaux, Laurent</mods:namePart>
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<mods:namePart id="6FAFA362D78C3D3C6218A6EC23B195DF">MéTais, Grégoire</mods:namePart>
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<mods:namePart id="3C3B98735A299993F4D348F31080D961">Antoine, Pierre-Olivier</mods:namePart>
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<mods:title id="B202E5FF642A77878B2EF9D81C6BBD2B">Zoological Journal of the Linnean Society</mods:title>
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<taxonomicName id="71F34D69FFBB9D124F7BF956FA36B6B0" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[1029,1438,1721,1744]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="20" pageNumber="309" phylum="Chordata" rank="species" species="hyopotamoides">
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<emphasis id="8487EAF8FFBB9D124F7BF956FA36B6B0" box="[1029,1438,1721,1744]" italics="true" pageId="20" pageNumber="298">
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<smallCapsWord id="B0AAA036FFBB9D124F7BF956FB7FB6B0" baselines="1738,1739" box="[1029,1239,1721,1744]" lowerCaseFontSize="8" mainFontSize="10" normCase="title" normString="Parabrachyodus" pageId="20" pageNumber="298">
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<emphasis id="8487EAF8FFBB9D124F7BF956FB7FB6B0" box="[1029,1239,1721,1744]" italics="true" pageId="20" pageNumber="298">PARABRACHYODUS</emphasis>
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</smallCapsWord>
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<smallCapsWord id="B0AAA036FFBB9D124FA1F953FA36B6B0" baselines="1739" box="[1247,1438,1724,1744]" lowerCaseFontSize="8" mainFontSize="10" normCase="lower" normString="hyopotamoides" pageId="20" pageNumber="298">HYOPOTAMOIDES</smallCapsWord>
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</emphasis>
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</taxonomicName>
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<subSubSection id="FEE96561FFBB9D1C4845F90EFD39B711" lastPageId="26" lastPageNumber="304" pageId="20" pageNumber="298" type="description">
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<paragraph id="B64C36EAFFBB9D134845F90EFF44B583" blockId="20.[827,1444,1761,1905]" lastBlockId="21.[145,764,197,1905]" lastPageId="21" lastPageNumber="299" pageId="20" pageNumber="298">
|
||
Based on unstable dental characters, early authors like Lydekker, Pilgrim and Forster-Cooper created up to ten species of
|
||
<taxonomicName id="71F34D69FFBB9D124F1BF8F1FB59B753" box="[1125,1265,1822,1843]" class="Mammalia" family="Anthracotheriidae" genus="Brachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="20" pageNumber="298" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFBB9D124F1BF8F1FB59B753" box="[1125,1265,1822,1843]" italics="true" pageId="20" pageNumber="298">Brachyodus</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<taxonomicName id="71F34D69FFBB9D124E7CF8F0FA34B754" box="[1282,1436,1823,1844]" class="Mammalia" family="Anthracotheriidae" genus="Brachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="20" pageNumber="298" phylum="Chordata" rank="species" species="africanus">
|
||
<emphasis id="8487EAF8FFBB9D124E7CF8F0FA34B754" box="[1282,1436,1823,1844]" italics="true" pageId="20" pageNumber="298">Br. africanus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="71F34D69FFBB9D124845F8D1FC5FB732" box="[827,1015,1853,1875]" class="Mammalia" family="Anthracotheriidae" genus="Brachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="20" pageNumber="298" phylum="Chordata" rank="species" species="gandoiensis">
|
||
<emphasis id="8487EAF8FFBB9D124845F8D1FC5FB732" box="[827,1015,1853,1875]" italics="true" pageId="20" pageNumber="298">Br. gandoiensis</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="71F34D69FFBB9D124F76F8D1FB02B733" box="[1032,1194,1854,1875]" class="Mammalia" family="Anthracotheriidae" genus="Brachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="20" pageNumber="298" phylum="Chordata" rank="species" species="giganteus">
|
||
<emphasis id="8487EAF8FFBB9D124F76F8D1FB02B733" box="[1032,1194,1854,1875]" italics="true" pageId="20" pageNumber="298">Br. giganteus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="71F34D69FFBB9D124FC5F8D1FA34B732" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[1211,1436,1853,1875]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="20" pageNumber="298" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFBB9D124FC5F8D1FA34B732" box="[1211,1436,1853,1875]" italics="true" pageId="20" pageNumber="298">Br. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="71F34D69FFBB9D124845F8B3FC6FB711" box="[827,967,1884,1905]" class="Mammalia" family="Anthracotheriidae" genus="Brachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="20" pageNumber="298" phylum="Chordata" rank="species" species="indicus">
|
||
<emphasis id="8487EAF8FFBB9D124845F8B3FC6FB711" box="[827,967,1884,1905]" italics="true" pageId="20" pageNumber="298">Br. indicus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="71F34D69FFBB9D1248A4F8B3FB61B711" authorityName="Prasad" authorityYear="1964" box="[986,1225,1884,1905]" class="Mammalia" family="Anthracotheriidae" genus="Brachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="20" pageNumber="298" phylum="Chordata" rank="species" species="manchharensis">
|
||
<emphasis id="8487EAF8FFBB9D1248A4F8B3FB61B711" box="[986,1225,1884,1905]" italics="true" pageId="20" pageNumber="298">Br. manchharensis</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="71F34D69FFBB9D124FA3F8B3FAC4B711" box="[1245,1388,1884,1905]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="20" pageNumber="298" phylum="Chordata" rank="species" species="obtusus">
|
||
<emphasis id="8487EAF8FFBB9D124FA3F8B3FAC4B711" box="[1245,1388,1884,1905]" italics="true" pageId="20" pageNumber="298">Br. obtusus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="71F34D69FFBB9D134EFEF8B3FEAAB0BA" authorityName="Forster-Cooper" authorityYear="1924" class="Mammalia" family="Anthracotheriidae" genus="Telmatodon" higherTaxonomySource="GBIF" kingdom="Animalia" lastPageId="21" lastPageNumber="299" order="Artiodactyla" pageId="20" pageNumber="298" phylum="Chordata" rank="species" species="orientalis">
|
||
<emphasis id="8487EAF8FFBB9D134EFEF8B3FEAAB0BA" italics="true" lastPageId="21" lastPageNumber="299" pageId="20" pageNumber="298">Br. orientalis</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="71F34D69FFBA9D134A6FFF29FE34B0BA" box="[273,412,197,219]" class="Mammalia" family="Anthracotheriidae" genus="Brachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="pilgrimi">
|
||
<emphasis id="8487EAF8FFBA9D134A6FFF29FE34B0BA" box="[273,412,197,219]" italics="true" pageId="21" pageNumber="299">Br. pilgrimi</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="71F34D69FFBA9D134AA7FF29FDDEB0BA" box="[473,630,197,219]" class="Mammalia" family="Anthracotheriidae" genus="Brachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="platydens">
|
||
<emphasis id="8487EAF8FFBA9D134AA7FF29FDDEB0BA" box="[473,630,197,219]" italics="true" pageId="21" pageNumber="299">Br. platydens</emphasis>
|
||
</taxonomicName>
|
||
). Our analyses of character variance (namely the multiplication of original crests and styles around the protocone of upper jugal teeth) led us to synonymize these taxa with
|
||
<taxonomicName id="71F34D69FFBA9D134BAEFEAFFE10B135" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[208,440,320,341]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFBA9D134BAEFEAFFE10B135" box="[208,440,320,341]" italics="true" pageId="21" pageNumber="299">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
. For example,
|
||
<taxonomicName id="71F34D69FFBA9D134912FEAFFEADB113" class="Mammalia" family="Anthracotheriidae" genus="Brachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="platydens">
|
||
<emphasis id="8487EAF8FFBA9D134912FEAFFEADB113" italics="true" pageId="21" pageNumber="299">Brachyodus platydens</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation id="2EC82A6FFFBA9D134A68FEB1FECCB114" box="[278,356,350,372]" captionStart="Figure 5" captionStartId="10.[163,241,1448,1470]" captionTargetBox="[243,1362,198,1415]" captionTargetId="figure-25@10.[240,1366,195,1418]" captionTargetPageId="10" captionText="Figure 5. Upper teeth of Parabrachyodus hyopotamoides. A, left upper incisor (I2/ or I3/) in lingual (left) and labial (right) view (UM-SAM4-024). B, P1/-P4/ from a left maxilla (M12034) in occlusal view. C, right P1/ (UM-SAM4-025) in occlusal view (mirror). D, left P3/ (M10598) in occlusal view. E, left maxilla with P4/-M1/ in occlusal view, with a drawing of P4/. F, left dP3/ (M12820) in occlusal view. Abbreviations derived from the nomenclature introduced in Figure 2: protoc., protocrista. G, dental variation of Parabrachyodus hyopotamoides figured on composite upper molars (N = 26 for M3/; N = 11 for M2/). Variable characters are coloured. Character frequency: black: 100%; red: (75–99%); yellow: (50–74%); green: (25–49%); blue: (0–24%). The presence/absence of the coloured character is evaluated. 1, mesostyle divided into two slender apices; 2, parastyle of the same size or smaller than mesostyle; 3, wing-like labial cingulum; 4, postectoprotocrista reaching the lingual edge of the tooth; 5, double curvature of the postmetacristule. The arrow indicates the mesiolingual direction. Teeth are not to scale. H, left maxilla with M1/-M3/ (M12718) in occlusal view. I, right maxilla with P4/-M3/ (M12824) in occlusal view (mirror). Comparisons with drawings of (J) left M3/ of Brachyodus onoideus in occlusal view, modified from Ducrocq & Lihoreau (2006); K, left M3/ of Telmatodon orientalis (M12040, holotype) in occlusal view; L, right M3/ of Gonotelma shahbazi (UM-SAM4-010) in occlusal view (mirror); M, left M3/ of SiƲameryx palaeindicus (UM-SAM4-005) in occlusal view. N, M3/ of Par. hyopotamoides (UM-SAM5-001) in distal view, showing the low value of the angle between the lingual slope of the metaconule and the socket. Scale bars all equal 10 mm." figureDoi="http://doi.org/10.5281/zenodo.7927073" httpUri="https://zenodo.org/record/7927073/files/figure.png" pageId="21" pageNumber="299">Fig. 5I</figureCitation>
|
||
;
|
||
<bibRefCitation id="D2624B1BFFBA9D134A0AFEB0FDC6B114" author="Forster-Cooper C" box="[372,622,351,372]" pageId="21" pageNumber="299" pagination="1 - 59" refId="ref24403" refString="Forster-Cooper C. 1924. The Anthracotheriidae of the Dera Bugti deposits in Baluchistan. Memoirs of the Geological SurVey of India. Palaeontologica Indica 4: 1 - 59." type="journal article" year="1924">Forster-Cooper, 1924</bibRefCitation>
|
||
), described by the flattest upper molars of the historical collections, cannot be maintained because its definition is based on a degree of wear. The square shape of the upper molars of
|
||
<taxonomicName id="71F34D69FFBA9D134A35FE35FE53B18E" box="[331,507,473,495]" class="Mammalia" family="Anthracotheriidae" genus="Brachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="gandoiensis">
|
||
<emphasis id="8487EAF8FFBA9D134A35FE35FE53B18E" box="[331,507,473,495]" italics="true" pageId="21" pageNumber="299">Br. gandoiensis</emphasis>
|
||
</taxonomicName>
|
||
is the only distinctive feature on which this species was founded (
|
||
<bibRefCitation id="D2624B1BFFBA9D1349E4FE17FE22B24C" author="Forster-Cooper C" pageId="21" pageNumber="299" pagination="1 - 59" refId="ref24403" refString="Forster-Cooper C. 1924. The Anthracotheriidae of the Dera Bugti deposits in Baluchistan. Memoirs of the Geological SurVey of India. Palaeontologica Indica 4: 1 - 59." type="journal article" year="1924">Forster-Cooper, 1924: 27–28</bibRefCitation>
|
||
), but we have shown that the shape of the upper molars in our sample highly depends on characters submitted to intraspecific variability. Moreover, the mesostyle is not loop-shaped but pinched in the
|
||
<typeStatus id="69488848FFBA9D134A07FD7DFE05B2C7" box="[377,429,658,679]" pageId="21" pageNumber="299">type</typeStatus>
|
||
species of
|
||
<taxonomicName id="71F34D69FFBA9D13494CFD7EFD17B2C6" box="[562,703,657,678]" class="Mammalia" family="Anthracotheriidae" genus="Brachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFBA9D13494CFD7EFD17B2C6" box="[562,703,657,678]" italics="true" pageId="21" pageNumber="299">Brachyodus</emphasis>
|
||
</taxonomicName>
|
||
(e.g.
|
||
<figureCitation id="2EC82A6FFFBA9D134BEFFD5FFF77B2A6" box="[145,223,688,710]" captionStart="Figure 5" captionStartId="10.[163,241,1448,1470]" captionTargetBox="[243,1362,198,1415]" captionTargetId="figure-25@10.[240,1366,195,1418]" captionTargetPageId="10" captionText="Figure 5. Upper teeth of Parabrachyodus hyopotamoides. A, left upper incisor (I2/ or I3/) in lingual (left) and labial (right) view (UM-SAM4-024). B, P1/-P4/ from a left maxilla (M12034) in occlusal view. C, right P1/ (UM-SAM4-025) in occlusal view (mirror). D, left P3/ (M10598) in occlusal view. E, left maxilla with P4/-M1/ in occlusal view, with a drawing of P4/. F, left dP3/ (M12820) in occlusal view. Abbreviations derived from the nomenclature introduced in Figure 2: protoc., protocrista. G, dental variation of Parabrachyodus hyopotamoides figured on composite upper molars (N = 26 for M3/; N = 11 for M2/). Variable characters are coloured. Character frequency: black: 100%; red: (75–99%); yellow: (50–74%); green: (25–49%); blue: (0–24%). The presence/absence of the coloured character is evaluated. 1, mesostyle divided into two slender apices; 2, parastyle of the same size or smaller than mesostyle; 3, wing-like labial cingulum; 4, postectoprotocrista reaching the lingual edge of the tooth; 5, double curvature of the postmetacristule. The arrow indicates the mesiolingual direction. Teeth are not to scale. H, left maxilla with M1/-M3/ (M12718) in occlusal view. I, right maxilla with P4/-M3/ (M12824) in occlusal view (mirror). Comparisons with drawings of (J) left M3/ of Brachyodus onoideus in occlusal view, modified from Ducrocq & Lihoreau (2006); K, left M3/ of Telmatodon orientalis (M12040, holotype) in occlusal view; L, right M3/ of Gonotelma shahbazi (UM-SAM4-010) in occlusal view (mirror); M, left M3/ of SiƲameryx palaeindicus (UM-SAM4-005) in occlusal view. N, M3/ of Par. hyopotamoides (UM-SAM5-001) in distal view, showing the low value of the angle between the lingual slope of the metaconule and the socket. Scale bars all equal 10 mm." figureDoi="http://doi.org/10.5281/zenodo.7927073" httpUri="https://zenodo.org/record/7927073/files/figure.png" pageId="21" pageNumber="299">Fig. 5J</figureCitation>
|
||
). The absence of biometric discriminations between upper teeth invalidates
|
||
<taxonomicName id="71F34D69FFBA9D134A86FD21FD51B283" box="[504,761,718,739]" class="Mammalia" family="Anthracotheriidae" genus="Brachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="orientalis">
|
||
<emphasis id="8487EAF8FFBA9D134A86FD21FD51B283" box="[504,761,718,739]" italics="true" pageId="21" pageNumber="299">Brachyodus orientalis</emphasis>
|
||
</taxonomicName>
|
||
defined essentially from upper molars smaller than other specimens (
|
||
<figureCitation id="2EC82A6FFFBA9D134A1BFCE3FE15B342" box="[357,445,780,802]" captionStart="Figure 5" captionStartId="10.[163,241,1448,1470]" captionTargetBox="[243,1362,198,1415]" captionTargetId="figure-25@10.[240,1366,195,1418]" captionTargetPageId="10" captionText="Figure 5. Upper teeth of Parabrachyodus hyopotamoides. A, left upper incisor (I2/ or I3/) in lingual (left) and labial (right) view (UM-SAM4-024). B, P1/-P4/ from a left maxilla (M12034) in occlusal view. C, right P1/ (UM-SAM4-025) in occlusal view (mirror). D, left P3/ (M10598) in occlusal view. E, left maxilla with P4/-M1/ in occlusal view, with a drawing of P4/. F, left dP3/ (M12820) in occlusal view. Abbreviations derived from the nomenclature introduced in Figure 2: protoc., protocrista. G, dental variation of Parabrachyodus hyopotamoides figured on composite upper molars (N = 26 for M3/; N = 11 for M2/). Variable characters are coloured. Character frequency: black: 100%; red: (75–99%); yellow: (50–74%); green: (25–49%); blue: (0–24%). The presence/absence of the coloured character is evaluated. 1, mesostyle divided into two slender apices; 2, parastyle of the same size or smaller than mesostyle; 3, wing-like labial cingulum; 4, postectoprotocrista reaching the lingual edge of the tooth; 5, double curvature of the postmetacristule. The arrow indicates the mesiolingual direction. Teeth are not to scale. H, left maxilla with M1/-M3/ (M12718) in occlusal view. I, right maxilla with P4/-M3/ (M12824) in occlusal view (mirror). Comparisons with drawings of (J) left M3/ of Brachyodus onoideus in occlusal view, modified from Ducrocq & Lihoreau (2006); K, left M3/ of Telmatodon orientalis (M12040, holotype) in occlusal view; L, right M3/ of Gonotelma shahbazi (UM-SAM4-010) in occlusal view (mirror); M, left M3/ of SiƲameryx palaeindicus (UM-SAM4-005) in occlusal view. N, M3/ of Par. hyopotamoides (UM-SAM5-001) in distal view, showing the low value of the angle between the lingual slope of the metaconule and the socket. Scale bars all equal 10 mm." figureDoi="http://doi.org/10.5281/zenodo.7927073" httpUri="https://zenodo.org/record/7927073/files/figure.png" pageId="21" pageNumber="299">Fig. 5H</figureCitation>
|
||
;
|
||
<bibRefCitation id="D2624B1BFFBA9D134AB2FCE3FD59B341" author="Forster-Cooper C" box="[460,753,780,801]" pageId="21" pageNumber="299" pagination="1 - 59" refId="ref24403" refString="Forster-Cooper C. 1924. The Anthracotheriidae of the Dera Bugti deposits in Baluchistan. Memoirs of the Geological SurVey of India. Palaeontologica Indica 4: 1 - 59." type="journal article" year="1924">Forster-Cooper, 1924: 32</bibRefCitation>
|
||
) but with morphology equivalent to all the synonyms of
|
||
<taxonomicName id="71F34D69FFBA9D134BCEFCA5FE34B33E" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[176,412,841,863]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFBA9D134BCEFCA5FE34B33E" box="[176,412,841,863]" italics="true" pageId="21" pageNumber="299">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
. On the basis of the morphology of the paraconule, upper molars from the Bugti Hills attributed to
|
||
<taxonomicName id="71F34D69FFBA9D134A07FC69FD29B3FB" box="[377,641,902,923]" class="Mammalia" family="Anthracotheriidae" genus="Brachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="africanus">
|
||
<emphasis id="8487EAF8FFBA9D134A07FC69FD29B3FB" box="[377,641,902,923]" italics="true" pageId="21" pageNumber="299">Brachyodus africanus</emphasis>
|
||
</taxonomicName>
|
||
belong to
|
||
<taxonomicName id="71F34D69FFBA9D134BEFFC4AFEECB3DA" baseAuthorityName="Ducrocq & Lihoreau" baseAuthorityYear="2006" box="[145,324,933,954]" class="Mammalia" family="Anthracotheriidae" genus="Sivameryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="palaeindicus">
|
||
<emphasis id="8487EAF8FFBA9D134BEFFC4AFEECB3DA" box="[145,324,933,954]" italics="true" pageId="21" pageNumber="299">S. palaeindicus</emphasis>
|
||
</taxonomicName>
|
||
(e.g. GSI B463;
|
||
<bibRefCitation id="D2624B1BFFBA9D13497AFC4AFD0BB3DA" author="Pilgrim GE" box="[516,675,933,955]" pageId="21" pageNumber="299" pagination="1 - 83" refId="ref26597" refString="Pilgrim GE. 1912. The vertebrate fauna of the Gaj series in the Bugti Hills and the Punjab. Memoirs of the Geological SurVey of India. Paleontologia Indica 4: 1 - 83." type="journal article" year="1912">Pilgrim, 1912</bibRefCitation>
|
||
: pl. 22, fig. 1) and to the larger
|
||
<taxonomicName id="71F34D69FFBA9D134AD7FC2BFD24B3B9" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[425,652,964,985]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFBA9D134AD7FC2BFD24B3B9" box="[425,652,964,985]" italics="true" pageId="21" pageNumber="299">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
(e.g. GSI B462;
|
||
<bibRefCitation id="D2624B1BFFBA9D134BA5FC0DFED1B397" author="Pilgrim GE" box="[219,377,994,1016]" pageId="21" pageNumber="299" pagination="1 - 83" refId="ref26597" refString="Pilgrim GE. 1912. The vertebrate fauna of the Gaj series in the Bugti Hills and the Punjab. Memoirs of the Geological SurVey of India. Paleontologia Indica 4: 1 - 83." type="journal article" year="1912">Pilgrim, 1912</bibRefCitation>
|
||
: pl. 22, fig. 2).
|
||
<bibRefCitation id="D2624B1BFFBA9D13495BFC0DFD50B398" author="Pilgrim GE" box="[549,760,994,1016]" pageId="21" pageNumber="299" pagination="1 - 83" refId="ref26597" refString="Pilgrim GE. 1912. The vertebrate fauna of the Gaj series in the Bugti Hills and the Punjab. Memoirs of the Geological SurVey of India. Paleontologia Indica 4: 1 - 83." type="journal article" year="1912">Pilgrim (1912: 50)</bibRefCitation>
|
||
considered the M/3 displaying postentostylids as pertaining to
|
||
<taxonomicName id="71F34D69FFBA9D134A4CFBCFFD9CB455" box="[306,564,1056,1077]" class="Mammalia" family="Anthracotheriidae" genus="Brachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="giganteus">
|
||
<emphasis id="8487EAF8FFBA9D134A4CFBCFFD9CB455" box="[306,564,1056,1077]" italics="true" pageId="21" pageNumber="299">Brachyodus giganteus</emphasis>
|
||
</taxonomicName>
|
||
, the M/3 lacking the postentostylid belonging to
|
||
<taxonomicName id="71F34D69FFBA9D13496AFBD0FD5AB433" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[532,754,1086,1108]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFBA9D13496AFBD0FD5AB433" box="[532,754,1086,1108]" italics="true" pageId="21" pageNumber="299">Br. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
. However, as we have shown, this character is commonly subjected to intraspecific variability in close relatives of
|
||
<taxonomicName id="71F34D69FFBA9D134A64FB74FE52B4CF" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[282,506,1178,1200]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFBA9D134A64FB74FE52B4CF" box="[282,506,1178,1200]" italics="true" pageId="21" pageNumber="299">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
. The single-cuspidate P4/ were grouped in
|
||
<taxonomicName id="71F34D69FFBA9D134AF4FB56FD07B4AE" box="[394,687,1209,1230]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="obtusus">
|
||
<emphasis id="8487EAF8FFBA9D134AF4FB56FD07B4AE" box="[394,687,1209,1230]" italics="true" pageId="21" pageNumber="299">Parabrachyodus obtusus</emphasis>
|
||
</taxonomicName>
|
||
(=
|
||
<taxonomicName id="71F34D69FFBA9D1349A6FB55FF45B48D" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="obtusus">
|
||
<emphasis id="8487EAF8FFBA9D1349A6FB55FF45B48D" italics="true" pageId="21" pageNumber="299">Br. obtusus</emphasis>
|
||
</taxonomicName>
|
||
) by Forster-Cooper ((1915, 1924: 33) based on this criterion, interpreted as a dental anomaly by
|
||
<bibRefCitation id="D2624B1BFFBA9D134BCAFAFAFEEAB54B" author="Viret J" box="[180,322,1301,1323]" pageId="21" pageNumber="299" pagination="887 - 1084" refId="ref27315" refString="Viret J. 1961. Artiodactyla. In: Piveteau J, ed. Traite de paleontologie VI. Paris: Masson, 887 - 1084." type="book chapter" year="1961">Viret (1961)</bibRefCitation>
|
||
among those that can appear in isolated series of anthracotheres (
|
||
<bibRefCitation id="D2624B1BFFBA9D13497FFADBFD42B529" author="Ducrocq S & Chaimanee Y & Suteethorn V & Jaeger JJ" box="[513,746,1331,1353]" pageId="21" pageNumber="299" pagination="355 - 360" refId="ref24223" refString="Ducrocq S, Chaimanee Y, Suteethorn V, Jaeger JJ. 1995. Dental anomalies in Upper Eocene Anthracotheriidae: a possible case of inbreeding. Lethaia 28: 355 - 360." type="journal article" year="1995">
|
||
Ducrocq
|
||
<emphasis id="8487EAF8FFBA9D134917FADBFD34B529" box="[617,668,1332,1353]" italics="true" pageId="21" pageNumber="299">et al</emphasis>
|
||
., 1995
|
||
</bibRefCitation>
|
||
). Considering the frequency of this character state in the studied series of anthracotheres, it is included in the common variability of
|
||
<taxonomicName id="71F34D69FFBA9D134AB1FA7FFD1EB5C5" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[463,694,1424,1445]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFBA9D134AB1FA7FFD1EB5C5" box="[463,694,1424,1445]" italics="true" pageId="21" pageNumber="299">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
, as it is for the P4/ of some hippopotamids (
|
||
<bibRefCitation id="D2624B1BFFBA9D13493DFA41FD5AB5A4" author="Boisserie JR" box="[579,754,1454,1476]" pageId="21" pageNumber="299" pagination="1 - 26" refId="ref23882" refString="Boisserie JR. 2005. The phylogeny and taxonomy of Hippopotamidae (Mammalia: Artiodactyla): a review based on morphology and cladistic analysis. Zoological Journal of the Linnean Society 143: 1 - 26." type="journal article" year="2005">Boisserie, 2005</bibRefCitation>
|
||
: fig. 5D).
|
||
</paragraph>
|
||
<paragraph id="B64C36EAFFBA9D134BD7FA03FAE7B22B" blockId="21.[145,764,197,1905]" lastBlockId="21.[809,1427,197,1414]" pageId="21" pageNumber="299">
|
||
Even after attempts to resolve the systematics of
|
||
<taxonomicName id="71F34D69FFBA9D134BD1F9E4FE3EB67F" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[175,406,1546,1568]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFBA9D134BD1F9E4FE3EB67F" box="[175,406,1546,1568]" italics="true" pageId="21" pageNumber="299">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation id="D2624B1BFFBA9D134AD6F9E5FD85B640" author="Viret J" box="[424,557,1546,1568]" pageId="21" pageNumber="299" pagination="887 - 1084" refId="ref27315" refString="Viret J. 1961. Artiodactyla. In: Piveteau J, ed. Traite de paleontologie VI. Paris: Masson, 887 - 1084." type="book chapter" year="1961">Viret, 1961</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="D2624B1BFFBA9D134942F9E5FD41B67F" author="Pickford M" box="[572,745,1546,1568]" pageId="21" pageNumber="299" pagination="289 - 350" refId="ref26447" refString="Pickford M. 1987. Revision des suiformes (Artiodactyla, Mammalia) de Bugti (Pakistan). Annales de Paleontologie 73: 289 - 350." type="journal article" year="1987">Pickford, 1987</bibRefCitation>
|
||
), the diagnostic characters of this species remained partially misunderstood, with the strength of the convex labial ribs on paracone and metacone or the loop-like styles on upper molars being considered here as plesiomorphic characters shared with all bothriodontines from the Bugti Hills. Our biometric analysis shows that a smaller mesial than distal width of the lower molars was erroneously considered by
|
||
<bibRefCitation id="D2624B1BFFBA9D134A66F8F1FE6AB753" author="Dineur H" box="[280,450,1822,1844]" pageId="21" pageNumber="299" refId="ref24047" refString="Dineur H. 1981. Le genre Brachyodus, Anthracotheriidae (Artiodactyla, Mammalia) du Miocene inferieur d'Europe et d'Afrique. Unpublished Ph. D. Thesis, Universite Pierre et Marie Curie - Paris VI." type="book" year="1981">Dineur (1981)</bibRefCitation>
|
||
and
|
||
<bibRefCitation id="D2624B1BFFBA9D13497DF8F1FD16B753" author="Pickford M" box="[515,702,1822,1844]" pageId="21" pageNumber="299" pagination="289 - 350" refId="ref26447" refString="Pickford M. 1987. Revision des suiformes (Artiodactyla, Mammalia) de Bugti (Pakistan). Annales de Paleontologie 73: 289 - 350." type="journal article" year="1987">Pickford (1987)</bibRefCitation>
|
||
as a distinctive character between
|
||
<taxonomicName id="71F34D69FFBA9D13497FF8D2FD6BB732" authorityName="Forster-Cooper" authorityYear="1915" box="[513,707,1853,1874]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFBA9D13497FF8D2FD6BB732" box="[513,707,1853,1874]" italics="true" pageId="21" pageNumber="299">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="71F34D69FFBA9D134BEFF8B4FEB4B710" box="[145,284,1883,1904]" class="Mammalia" family="Anthracotheriidae" genus="Brachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFBA9D134BEFF8B4FEB4B710" box="[145,284,1883,1904]" italics="true" pageId="21" pageNumber="299">Brachyodus</emphasis>
|
||
</taxonomicName>
|
||
. Conversely, the M/3 of
|
||
<taxonomicName id="71F34D69FFBA9D134944F8B4FD51B710" authorityName="Forster-Cooper" authorityYear="1915" box="[570,761,1883,1904]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFBA9D134944F8B4FD51B710" box="[570,761,1883,1904]" italics="true" pageId="21" pageNumber="299">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
is not defined by a trigonid larger than the talonid, as suggested by
|
||
<bibRefCitation id="D2624B1BFFBA9D13488FFF0BFB52B099" author="Forster-Cooper C" box="[1009,1274,228,249]" pageId="21" pageNumber="299" pagination="514 - 522" refId="ref24343" refString="Forster-Cooper C. 1913. New anthracotheres and allied forms from Baluchistan - preliminary notice. Annals and Magazine of Natural History 12: 514 - 522." type="journal article" year="1913">Forster-Cooper (1913)</bibRefCitation>
|
||
, despite the proportions of the first M/3 allocated to the genus (
|
||
<figureCitation id="2EC82A6FFFBA9D13484DFECEFC3AB156" box="[819,914,289,311]" captionStart="Figure 1" captionStartId="1.[145,223,1829,1851]" captionTargetBox="[144,1425,1495,1796]" captionTargetId="figure-713@1.[141,1428,1492,1799]" captionTargetPageId="1" captionText="Figure 1. Drawings from Forster-Cooper (1915) of A, the first upper dental row (P4/-M3/, M12031) and B, lower molar (M/3, M12030) allocated to the anthracotheriid Parabrachyodus, from the Bugti Hills, Pakistan. Note the high degree of wear and the focus of the author on P4/’s morphology. Scale bars equal 10 mm." figureDoi="http://doi.org/10.5281/zenodo.7927063" httpUri="https://zenodo.org/record/7927063/files/figure.png" pageId="21" pageNumber="299">Fig. 1B</figureCitation>
|
||
). Furthermore, drawings of early authors show that the presence of the ectoprotocrista linked to a protostyle (a diagnostic character of
|
||
<taxonomicName id="71F34D69FFBA9D134857FE92FBBEB1F2" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[809,1046,381,402]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFBA9D134857FE92FBBEB1F2" box="[809,1046,381,402]" italics="true" pageId="21" pageNumber="299">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
) was not considered as a relevant feature (
|
||
<figureCitation id="2EC82A6FFFBA9D1348B2FE74FB89B1D1" box="[972,1057,411,433]" captionStart="Figure 1" captionStartId="1.[145,223,1829,1851]" captionTargetBox="[144,1425,1495,1796]" captionTargetId="figure-713@1.[141,1428,1492,1799]" captionTargetPageId="1" captionText="Figure 1. Drawings from Forster-Cooper (1915) of A, the first upper dental row (P4/-M3/, M12031) and B, lower molar (M/3, M12030) allocated to the anthracotheriid Parabrachyodus, from the Bugti Hills, Pakistan. Note the high degree of wear and the focus of the author on P4/’s morphology. Scale bars equal 10 mm." figureDoi="http://doi.org/10.5281/zenodo.7927063" httpUri="https://zenodo.org/record/7927063/files/figure.png" pageId="21" pageNumber="299">Fig. 1A</figureCitation>
|
||
). It is noteworthy that
|
||
<bibRefCitation id="D2624B1BFFBA9D134E46FE74FC08B1B0" author="Pilgrim GE" pageId="21" pageNumber="299" pagination="1 - 83" refId="ref26597" refString="Pilgrim GE. 1912. The vertebrate fauna of the Gaj series in the Bugti Hills and the Punjab. Memoirs of the Geological SurVey of India. Paleontologia Indica 4: 1 - 83." type="journal article" year="1912">Pilgrim (1912: 55)</bibRefCitation>
|
||
was the first to mention some character states of
|
||
<taxonomicName id="71F34D69FFBA9D1348EAFE36FBFAB18E" authorityName="Forster-Cooper" authorityYear="1915" box="[916,1106,473,494]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFBA9D1348EAFE36FBFAB18E" box="[916,1106,473,494]" italics="true" pageId="21" pageNumber="299">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
(=
|
||
<taxonomicName id="71F34D69FFBA9D134F04FE36FAACB18E" box="[1146,1284,473,494]" class="Mammalia" family="Anthracotheriidae" genus="Brachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFBA9D134F04FE36FAACB18E" box="[1146,1284,473,494]" italics="true" pageId="21" pageNumber="299">Brachyodus</emphasis>
|
||
</taxonomicName>
|
||
‘
|
||
<taxonomicName id="71F34D69FFBA9D134E6FFE36FA2AB18E" box="[1297,1410,473,494]" class="Mammalia" family="Anthracotheriidae" genus="Brachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="giganteus">
|
||
<emphasis id="8487EAF8FFBA9D134E6FFE36FA2AB18E" box="[1297,1410,473,494]" italics="true" pageId="21" pageNumber="299">giganteus</emphasis>
|
||
</taxonomicName>
|
||
’) such as the ‘rudimentary additional cusp’ on P3/ underscoring the anthracotheriine affinity of this tooth, without interpreting them as diagnostic.
|
||
</paragraph>
|
||
<paragraph id="B64C36EAFFBA9D13483FFDBCFC1CB5E6" blockId="21.[809,1427,197,1414]" pageId="21" pageNumber="299">
|
||
The sample of
|
||
<taxonomicName id="71F34D69FFBA9D134894FDBCFB0CB208" authorityName="Forster-Cooper" authorityYear="1915" box="[1002,1188,595,616]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFBA9D134894FDBCFB0CB208" box="[1002,1188,595,616]" italics="true" pageId="21" pageNumber="299">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
is comprehensive in the Bugti Hills, with at least
|
||
<specimenCount id="A0F5FD63FFBA9D134FF8FD9DFA99B2E8" box="[1158,1329,626,648]" pageId="21" pageNumber="299" type="generic">103 specimens</specimenCount>
|
||
(mostly molars) when including the remains newly described here. All these remains belong to the single species of the genus,
|
||
<taxonomicName id="71F34D69FFBA9D1348E1FD20FBD0B283" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[927,1144,718,740]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFBA9D1348E1FD20FBD0B283" box="[927,1144,718,740]" italics="true" pageId="21" pageNumber="299">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
<bibRefCitation id="D2624B1BFFBA9D134FFCFD21FA86B283" author="Pickford M" box="[1154,1326,718,740]" pageId="21" pageNumber="299" pagination="289 - 350" refId="ref26447" refString="Pickford M. 1987. Revision des suiformes (Artiodactyla, Mammalia) de Bugti (Pakistan). Annales de Paleontologie 73: 289 - 350." type="journal article" year="1987">Pickford (1987)</bibRefCitation>
|
||
stressed the difficulties of discriminating the lower molars of
|
||
<taxonomicName id="71F34D69FFBA9D134857FCE3FBBDB340" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[809,1045,779,801]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFBA9D134857FCE3FBBDB340" box="[809,1045,779,801]" italics="true" pageId="21" pageNumber="299">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
from those of
|
||
<taxonomicName id="71F34D69FFBA9D134FB5FCE4FAF1B340" baseAuthorityName="Forster-Cooper" baseAuthorityYear="1924" box="[1227,1369,779,800]" class="Mammalia" family="Anthracotheriidae" genus="Telmatodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFBA9D134FB5FCE4FAF1B340" box="[1227,1369,779,800]" italics="true" pageId="21" pageNumber="299">Telmatodon</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="71F34D69FFBA9D134857FCC4FC1AB320" box="[809,946,811,832]" class="Mammalia" family="Anthracotheriidae" genus="Hemimeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFBA9D134857FCC4FC1AB320" box="[809,946,811,832]" italics="true" pageId="21" pageNumber="299">Hemimeryx</emphasis>
|
||
</taxonomicName>
|
||
, presumably because the lower molars display a greater number of polymorphic traits and are less represented in number of specimens than the upper molars. Also, the convergent morphology of the hypoconulids of
|
||
<taxonomicName id="71F34D69FFBA9D1348A3FC4AFB0AB3DA" box="[989,1186,933,954]" class="Mammalia" family="Anthracotheriidae" genus="Anthracotherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFBA9D1348A3FC4AFB0AB3DA" box="[989,1186,933,954]" italics="true" pageId="21" pageNumber="299">Anthracotherium</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="71F34D69FFBA9D134FA9FC4AFA39B3DA" authorityName="Forster-Cooper" authorityYear="1915" box="[1239,1425,933,954]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFBA9D134FA9FC4AFA39B3DA" box="[1239,1425,933,954]" italics="true" pageId="21" pageNumber="299">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
probably brought confusion in distinguishing these genera (e.g.
|
||
<bibRefCitation id="D2624B1BFFBA9D1348C8FC0DFAF9B397" author="Welcomme JL & Ginsburg L" box="[950,1361,994,1016]" pageId="21" pageNumber="299" pagination="999 - 1004" refId="ref27403" refString="Welcomme JL, Ginsburg L. 1997. Mise en evidence de l'Oligocene sur le territoire des Bugti (Balouchistan, Pakistan). Comptes Rendus de l'Academie des Sciences, Paris 325: 999 - 1004." type="journal article" year="1997">Welcomme & Ginsburg, 1997: 1002</bibRefCitation>
|
||
), further inspiring the assignment of
|
||
<taxonomicName id="71F34D69FFBA9D134FCCFBEEFAD9B476" authorityName="Forster-Cooper" authorityYear="1915" box="[1202,1393,1025,1046]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFBA9D134FCCFBEEFAD9B476" box="[1202,1393,1025,1046]" italics="true" pageId="21" pageNumber="299">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
in the
|
||
<taxonomicName id="71F34D69FFBA9D13482AFBF0FB80B455" box="[852,1064,1055,1077]" class="Mammalia" family="Anthracotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="subFamily" subFamily="Anthracotheriinae">Anthracotheriinae</taxonomicName>
|
||
instead of the Bothriodontinae (
|
||
<bibRefCitation id="D2624B1BFFBA9D13484CFBD1FC77B433" author="Pickford M" box="[818,991,1086,1108]" pageId="21" pageNumber="299" pagination="289 - 350" refId="ref26447" refString="Pickford M. 1987. Revision des suiformes (Artiodactyla, Mammalia) de Bugti (Pakistan). Annales de Paleontologie 73: 289 - 350." type="journal article" year="1987">Pickford, 1987</bibRefCitation>
|
||
: fig. 4), as such following
|
||
<bibRefCitation id="D2624B1BFFBA9D134E5EFBD1FCDBB412" author="Lydekker R" pageId="21" pageNumber="299" pagination="143 - 177" refId="ref25755" refString="Lydekker R. 1883. Siwalik selenodont suina. Memoirs of the Geological surVey of India. Palaeontologica Indica 10: 143 - 177." type="journal article" year="1883">Lydekker (1883)</bibRefCitation>
|
||
who considered this anthracothere as a representative of
|
||
<taxonomicName id="71F34D69FFBA9D1348ADFB94FB34B4F0" box="[979,1180,1147,1168]" class="Mammalia" family="Anthracotheriidae" genus="Anthracotherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFBA9D1348ADFB94FB34B4F0" box="[979,1180,1147,1168]" italics="true" pageId="21" pageNumber="299">Anthracotherium</emphasis>
|
||
</taxonomicName>
|
||
. However, the upper molars are more distinctive.
|
||
<taxonomicName id="71F34D69FFBA9D134FE7FB75FAD9B4D0" box="[1177,1393,1178,1200]" class="Mammalia" family="Anthracotheriidae" genus="Gonotelma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="major">
|
||
<emphasis id="8487EAF8FFBA9D134FE7FB75FAD9B4D0" box="[1177,1393,1178,1200]" italics="true" pageId="21" pageNumber="299">Gonotelma major</emphasis>
|
||
</taxonomicName>
|
||
is defined by a worn upper molar with a paraconule (
|
||
<bibRefCitation id="D2624B1BFFBA9D13484FFB38FB8CB48C" author="Forster-Cooper C" box="[817,1060,1239,1260]" pageId="21" pageNumber="299" pagination="1 - 59" refId="ref24403" refString="Forster-Cooper C. 1924. The Anthracotheriidae of the Dera Bugti deposits in Baluchistan. Memoirs of the Geological SurVey of India. Palaeontologica Indica 4: 1 - 59." type="journal article" year="1924">Forster-Cooper, 1924</bibRefCitation>
|
||
: pl. 5, fig. 1) and it is here considered as a junior synonym of
|
||
<taxonomicName id="71F34D69FFBA9D134FF9FB18FACBB56B" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[1159,1379,1270,1292]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFBA9D134FF9FB18FACBB56B" box="[1159,1379,1270,1292]" italics="true" pageId="21" pageNumber="299">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
. We concur with
|
||
<bibRefCitation id="D2624B1BFFBA9D1348C8FAFAFBCDB54A" author="Pickford M" box="[950,1125,1301,1323]" pageId="21" pageNumber="299" pagination="289 - 350" refId="ref26447" refString="Pickford M. 1987. Revision des suiformes (Artiodactyla, Mammalia) de Bugti (Pakistan). Annales de Paleontologie 73: 289 - 350." type="journal article" year="1987">Pickford (1987)</bibRefCitation>
|
||
in considering
|
||
<taxonomicName id="71F34D69FFBA9D134E6BFAFAFA39B54A" authorityName="Pilgrim" authorityYear="1908" box="[1301,1425,1301,1322]" class="Mammalia" family="Anthracotheriidae" genus="Gonotelma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFBA9D134E6BFAFAFA39B54A" box="[1301,1425,1301,1322]" italics="true" pageId="21" pageNumber="299">Gonotelma</emphasis>
|
||
</taxonomicName>
|
||
a monospecific genus with small tetracuspidate molars, first interpreted as pentacuspidate (
|
||
<bibRefCitation id="D2624B1BFFBA9D134E4CFABDFCC8B5E6" author="Pilgrim GE" pageId="21" pageNumber="299" pagination="139 - 166" refId="ref26562" refString="Pilgrim GE. 1908. The Tertiary and Post-Tertiary freshwater deposits of Baluchistan and Sind: with notices of new vertebrate. Records of the Geological SurVey of India 37: 139 - 166." type="journal article" year="1908">Pilgrim, 1908</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="D2624B1BFFBA9D134813FA9EFC0CB5E6" author="Pilgrim GE" box="[877,932,1393,1414]" pageId="21" pageNumber="299" pagination="1 - 83" refId="ref26597" refString="Pilgrim GE. 1912. The vertebrate fauna of the Gaj series in the Bugti Hills and the Punjab. Memoirs of the Geological SurVey of India. Paleontologia Indica 4: 1 - 83." type="journal article" year="1912">1912</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="B64C36EAFFBA9D134815FA2CFB6EB59A" blockId="21.[875,1358,1475,1530]" pageId="21" pageNumber="299">
|
||
<smallCapsWord id="B0AAA036FFBA9D134815FA2CFC33B5BB" baselines="1494,1494" box="[875,923,1475,1500]" lowerCaseFontSize="8" mainFontSize="10" normCase="title" normString="Age" pageId="21" pageNumber="299">AGE</smallCapsWord>
|
||
<smallCapsWord id="B0AAA036FFBA9D1348DDFA28FC7AB5BB" baselines="1494" box="[931,978,1479,1499]" lowerCaseFontSize="8" mainFontSize="10" normCase="lower" normString="and" pageId="21" pageNumber="299">AND</smallCapsWord>
|
||
<smallCapsWord id="B0AAA036FFBA9D1348A7FA28FB2DB5BB" baselines="1494" box="[985,1157,1479,1499]" lowerCaseFontSize="8" mainFontSize="10" normCase="lower" normString="geographical" pageId="21" pageNumber="299">GEOGRAPHICAL</smallCapsWord>
|
||
<smallCapsWord id="B0AAA036FFBA9D134FF2FA28FA82B5BB" baselines="1494" box="[1164,1322,1479,1499]" lowerCaseFontSize="8" mainFontSize="10" normCase="lower" normString="distribution" pageId="21" pageNumber="299">DISTRIBUTION</smallCapsWord>
|
||
<smallCapsWord id="B0AAA036FFBA9D134E4FFA28FAE6B5BB" baselines="1494" box="[1329,1358,1479,1499]" lowerCaseFontSize="8" mainFontSize="10" normCase="lower" normString="of" pageId="21" pageNumber="299">OF</smallCapsWord>
|
||
<taxonomicName id="71F34D69FFBA9D13488DFA0CFB6EB59A" authorityName="Forster-Cooper" authorityYear="1915" box="[1011,1222,1507,1530]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="genus">
|
||
<smallCapsWord id="B0AAA036FFBA9D13488DFA0CFB6EB59A" baselines="1524,1525" box="[1011,1222,1507,1530]" lowerCaseFontSize="8" mainFontSize="10" normCase="title" normString="Parabrachyodus" pageId="21" pageNumber="299">
|
||
<emphasis id="8487EAF8FFBA9D13488DFA0CFB6EB59A" box="[1011,1222,1507,1530]" italics="true" pageId="21" pageNumber="299">PARABRACHYODUS</emphasis>
|
||
</smallCapsWord>
|
||
</taxonomicName>
|
||
</paragraph>
|
||
<paragraph id="B64C36EAFFBA9D104857F9E4FDBDB54A" blockId="21.[809,1426,1546,1906]" lastBlockId="22.[163,780,197,1905]" lastPageId="22" lastPageNumber="300" pageId="21" pageNumber="299">
|
||
In addition to the series of
|
||
<taxonomicName id="71F34D69FFBA9D134FC6F9E4FC7FB65E" baseAuthorityName="Lydekker" baseAuthorityYear="1883" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFBA9D134FC6F9E4FC7FB65E" italics="true" pageId="21" pageNumber="299">Parabrachyodus hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
from SAM 4 (
|
||
<emphasis id="8487EAF8FFBA9D134F00F9C5FB38B65F" box="[1150,1168,1578,1599]" italics="true" pageId="21" pageNumber="299">c.</emphasis>
|
||
21 Mya), the isolated M3/ UM-SAM5-001 is the first reported occurrence of a fossil from SAM 5, which extends the age formally given to
|
||
<taxonomicName id="71F34D69FFBA9D1348A5F969FB16B6FA" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[987,1214,1669,1691]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFBA9D1348A5F969FB16B6FA" box="[987,1214,1669,1691]" italics="true" pageId="21" pageNumber="299">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
to approximately 19 Mya (
|
||
<bibRefCitation id="D2624B1BFFBA9D1348E8F94BFB29B6D9" author="Roddaz M & Said A & Guillot S & Antoine PO & Montel JM & Martin F & Darrozes J" box="[918,1153,1700,1722]" pageId="21" pageNumber="299" pagination="499 - 516" refId="ref26895" refString="Roddaz M, Said A, Guillot S, Antoine PO, Montel JM, Martin F, Darrozes J. 2011. Provenance of Cenozoic sedimentary rocks from the Sulaiman fold and thrust belt, Pakistan: implications for the palaeogeography of the Indus drainage system. Journal of the Geological Society 168: 499 - 516." type="journal article" year="2011">
|
||
Roddaz
|
||
<emphasis id="8487EAF8FFBA9D134886F94BFB9EB6D9" box="[1016,1078,1700,1721]" italics="true" pageId="21" pageNumber="299">et al.</emphasis>
|
||
, 2011
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="D2624B1BFFBA9D134FEEF94BFA29B6D9" author="Antoine PO & Metais G & Orliac MJ & Crochet JY & Flynn LJ & Marivaux L & Rajpar AR & Roohi G & Welcomme JL" box="[1168,1409,1700,1722]" pageId="21" pageNumber="299" pagination="400 - 422" refId="ref23524" refString="Antoine PO, Metais G, Orliac MJ, Crochet JY, Flynn LJ, Marivaux L, Rajpar AR, Roohi G, Welcomme JL. 2013. Mammalian Neogene biostratigraphy of the Sulaiman Province, Pakistan. In: Xiaoming W, Flynn LJ, Fortelius M, eds. Fossil mammals of Asia. New York: Columbia University Press, 400 - 422." type="book chapter" year="2013">
|
||
Antoine
|
||
<emphasis id="8487EAF8FFBA9D134F87F94BFA9FB6D9" box="[1273,1335,1700,1721]" italics="true" pageId="21" pageNumber="299">et al.</emphasis>
|
||
, 2013
|
||
</bibRefCitation>
|
||
). We thus confirm the presence of the species in the Vihowa Formation (e.g.
|
||
<bibRefCitation id="D2624B1BFFBA9D134F39F90EFA9AB696" author="Antoine PO & Metais G & Orliac MJ & Crochet JY & Flynn LJ & Marivaux L & Rajpar AR & Roohi G & Welcomme JL" box="[1095,1330,1761,1783]" pageId="21" pageNumber="299" pagination="400 - 422" refId="ref23524" refString="Antoine PO, Metais G, Orliac MJ, Crochet JY, Flynn LJ, Marivaux L, Rajpar AR, Roohi G, Welcomme JL. 2013. Mammalian Neogene biostratigraphy of the Sulaiman Province, Pakistan. In: Xiaoming W, Flynn LJ, Fortelius M, eds. Fossil mammals of Asia. New York: Columbia University Press, 400 - 422." type="book chapter" year="2013">
|
||
Antoine
|
||
<emphasis id="8487EAF8FFBA9D134FD0F90DFB42B696" box="[1198,1258,1761,1783]" italics="true" pageId="21" pageNumber="299">et al.</emphasis>
|
||
, 2013
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="D2624B1BFFBA9D134E3FF90EFC04B775" author="Nanda AC & Sehgal RK & Chauhan PR" pageId="21" pageNumber="299" pagination="54 - 68" refId="ref26200" refString="Nanda AC, Sehgal RK, Chauhan PR. 2017. Siwalik-age faunas from the Himalayan Foreland Basin of South Asia. Journal of Asian Earth Sciences 162: 54 - 68." type="journal article" year="2017">
|
||
Nanda
|
||
<emphasis id="8487EAF8FFBA9D134857F8EFFCCCB775" box="[809,868,1792,1813]" italics="true" pageId="21" pageNumber="299">et al.</emphasis>
|
||
, 2017
|
||
</bibRefCitation>
|
||
).
|
||
<taxonomicName id="71F34D69FFBA9D1348BAF8EFFA94B775" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[964,1340,1792,1813]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="21" pageNumber="299" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFBA9D1348BAF8EFFA94B775" box="[964,1340,1792,1813]" italics="true" pageId="21" pageNumber="299">Parabrachyodus hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
is also recorded with confidence in the Zinda Pir Dome, Sulaiman Province of
|
||
<collectingCountry id="CEE4767AFFBA9D134F50F8D2FB31B733" box="[1070,1177,1853,1875]" name="Pakistan" pageId="21" pageNumber="299">Pakistan</collectingCountry>
|
||
, about
|
||
<quantity id="710B9B0FFFBA9D134F8EF8D2FAE0B733" box="[1264,1352,1853,1875]" metricMagnitude="5" metricUnit="m" metricValue="2.0" pageId="21" pageNumber="299" unit="km" value="200.0">200 km</quantity>
|
||
north of the Bugti Hills, through three isolated specimens in localities Z114, Z154 and Safed Nala (
|
||
<bibRefCitation id="D2624B1BFFB99D1049F1FF2AFEA5B09A" author="Lindsay EH & Flynn LJ & Cheema IU & Barry JC & Downing K & Rajpar AR & Raza SM" pageId="22" pageNumber="300" pagination="1 - 19" refId="ref25636" refString="Lindsay EH, Flynn LJ, Cheema IU, Barry JC, Downing K, Rajpar AR, Raza SM. 2005. Will Downs and the Zinda Pir Dome. Palaeontologia Electronica 8: 1 - 19." type="journal article" year="2005">
|
||
Lindsay
|
||
<emphasis id="8487EAF8FFB99D104988FF29FF6AB099" italics="true" pageId="22" pageNumber="300">et al.</emphasis>
|
||
, 2005
|
||
</bibRefCitation>
|
||
). The ‘Interpretation B’ of
|
||
<bibRefCitation id="D2624B1BFFB99D10491FFF0BFF4EB178" author="Lindsay EH & Flynn LJ & Cheema IU & Barry JC & Downing K & Rajpar AR & Raza SM" pageId="22" pageNumber="300" pagination="1 - 19" refId="ref25636" refString="Lindsay EH, Flynn LJ, Cheema IU, Barry JC, Downing K, Rajpar AR, Raza SM. 2005. Will Downs and the Zinda Pir Dome. Palaeontologia Electronica 8: 1 - 19." type="journal article" year="2005">
|
||
Lindsay
|
||
<emphasis id="8487EAF8FFB99D1049B3FF0AFCA3B099" box="[717,779,228,250]" italics="true" pageId="22" pageNumber="300">et al.</emphasis>
|
||
(2005
|
||
</bibRefCitation>
|
||
: fig. 6B) for the correlation of the Zinda Pir localities to the Geomagnetic Polarity Time Scale (GPTS) being the most satisfactory (
|
||
<bibRefCitation id="D2624B1BFFB99D10491EFEAFFEBFB114" author="Antoine PO & Metais G & Orliac MJ & Crochet JY & Flynn LJ & Marivaux L & Rajpar AR & Roohi G & Welcomme JL" pageId="22" pageNumber="300" pagination="400 - 422" refId="ref23524" refString="Antoine PO, Metais G, Orliac MJ, Crochet JY, Flynn LJ, Marivaux L, Rajpar AR, Roohi G, Welcomme JL. 2013. Mammalian Neogene biostratigraphy of the Sulaiman Province, Pakistan. In: Xiaoming W, Flynn LJ, Fortelius M, eds. Fossil mammals of Asia. New York: Columbia University Press, 400 - 422." type="book chapter" year="2013">
|
||
Antoine
|
||
<emphasis id="8487EAF8FFB99D1049B6FEAFFCACB135" box="[712,772,320,341]" italics="true" pageId="22" pageNumber="300">et al.</emphasis>
|
||
, 2013: 416
|
||
</bibRefCitation>
|
||
), the localities Z114 and Z154 then lie between the magnetochrons C6AA and C6Bn. The corresponding approximate age is 21.95 Mya (
|
||
<bibRefCitation id="D2624B1BFFB99D1049E2FE73FEA0B1B0" author="Speijer RP & Palike H & Hollis CJ & Hooker JJ & Ogg JG" pageId="22" pageNumber="300" pagination="1087 - 1140" refId="ref27159" refString="Speijer RP, Palike H, Hollis CJ, Hooker JJ, Ogg JG. 2020. The paleogene period. In: Gradstein FM, Ogg JG, Schmitz M, Ogg GM, eds. Geologic time scale 2020. Amsterdam: Elsevier, 1087 - 1140." type="book chapter" year="2020">
|
||
Speijer
|
||
<emphasis id="8487EAF8FFB99D104989FE73FF69B1AF" italics="true" pageId="22" pageNumber="300">et al.</emphasis>
|
||
, 2020
|
||
</bibRefCitation>
|
||
; J. Barry, pers. comm., 2022), which makes it the oldest known occurrence of
|
||
<taxonomicName id="71F34D69FFB99D104956FE35FCADB18E" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[552,773,473,495]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFB99D104956FE35FCADB18E" box="[552,773,473,495]" italics="true" pageId="22" pageNumber="300">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
. The species is also documented in the Khari Nadi Formation (Kutch Basin,
|
||
<collectingRegion id="7437F808FFB99D104AB6FDF9FD8CB24C" box="[456,548,534,556]" country="India" name="Gujarat" pageId="22" pageNumber="300">Gujarat</collectingRegion>
|
||
,
|
||
<collectingCountry id="CEE4767AFFB99D10494EFDF9FDC7B24C" box="[560,623,534,556]" name="India" pageId="22" pageNumber="300">India</collectingCountry>
|
||
), via a palate with molars that have, among other traits, the characteristic morphology of the protocone and a large distostyle (
|
||
<bibRefCitation id="D2624B1BFFB99D104A5FFD9DFDBBB2E8" author="Bhandari A & Mohabey DM & Bajpai S & Tiwari BN & Pickford M" box="[289,531,626,648]" pageId="22" pageNumber="300" pagination="69 - 97" refId="ref23717" refString="Bhandari A, Mohabey DM, Bajpai S, Tiwari BN, Pickford M. 2010. Early Miocene mammals from central Kutch (Gujarat), Western India: implications for geochronology, biogeography, eustacy and intercontinental dispersals. Neues Jahrbuch fur Geologie und Palaontologie - Abhandlungen 256: 69 - 97." type="journal article" year="2010">
|
||
Bhandari
|
||
<emphasis id="8487EAF8FFB99D104AE8FD9CFE66B2E7" box="[406,462,626,648]" italics="true" pageId="22" pageNumber="300">et al.</emphasis>
|
||
, 2010
|
||
</bibRefCitation>
|
||
: fig. 7A). It is also accompanied by
|
||
<taxonomicName id="71F34D69FFB99D104A31FD7EFDC3B2C6" baseAuthorityName="Ducrocq & Lihoreau" baseAuthorityYear="2006" box="[335,619,657,678]" class="Mammalia" family="Anthracotheriidae" genus="Sivameryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="species" species="palaeindicus">
|
||
<emphasis id="8487EAF8FFB99D104A31FD7EFDC3B2C6" box="[335,619,657,678]" italics="true" pageId="22" pageNumber="300">SiƲameryx palaeindicus</emphasis>
|
||
</taxonomicName>
|
||
in the Kutch fauna. This assemblage could be younger than Samane Nala 4 and 5 according to Bhandari
|
||
<emphasis id="8487EAF8FFB99D10493EFD20FDD0B283" box="[576,632,718,740]" italics="true" pageId="22" pageNumber="300">et al.</emphasis>
|
||
(16.5 +/– 0.5 Mya). However, they precisely considered a last local appearance (LLA) of 16.5 Mya for
|
||
<taxonomicName id="71F34D69FFB99D104950FCE3FCA3B341" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[558,779,780,801]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFB99D104950FCE3FCA3B341" box="[558,779,780,801]" italics="true" pageId="22" pageNumber="300">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
to propose the most likely age of the Kutch mammal fauna. It is unclear whether or not
|
||
<taxonomicName id="71F34D69FFB99D104935FCA6FCA3B33E" authorityName="Forster-Cooper" authorityYear="1915" box="[587,779,841,862]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB99D104935FCA6FCA3B33E" box="[587,779,841,862]" italics="true" pageId="22" pageNumber="300">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
also occurs in the Level 6 of the Chitarwata Formation (
|
||
<emphasis id="8487EAF8FFB99D104BD5FC68FF14B3FC" box="[171,188,903,924]" italics="true" pageId="22" pageNumber="300">c.</emphasis>
|
||
17.5–18.0 Mya,
|
||
<bibRefCitation id="D2624B1BFFB99D104A03FC69FDF2B3FB" author="Roddaz M & Said A & Guillot S & Antoine PO & Montel JM & Martin F & Darrozes J" box="[381,602,902,924]" pageId="22" pageNumber="300" pagination="499 - 516" refId="ref26895" refString="Roddaz M, Said A, Guillot S, Antoine PO, Montel JM, Martin F, Darrozes J. 2011. Provenance of Cenozoic sedimentary rocks from the Sulaiman fold and thrust belt, Pakistan: implications for the palaeogeography of the Indus drainage system. Journal of the Geological Society 168: 499 - 516." type="journal article" year="2011">
|
||
Roddaz
|
||
<emphasis id="8487EAF8FFB99D104AA4FC68FDBCB3FB" box="[474,532,902,924]" italics="true" pageId="22" pageNumber="300">et al.</emphasis>
|
||
, 2011
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="D2624B1BFFB99D104919FC69FF74B3DA" author="Antoine PO & Metais G & Orliac MJ & Crochet JY & Flynn LJ & Marivaux L & Rajpar AR & Roohi G & Welcomme JL" pageId="22" pageNumber="300" pagination="400 - 422" refId="ref23524" refString="Antoine PO, Metais G, Orliac MJ, Crochet JY, Flynn LJ, Marivaux L, Rajpar AR, Roohi G, Welcomme JL. 2013. Mammalian Neogene biostratigraphy of the Sulaiman Province, Pakistan. In: Xiaoming W, Flynn LJ, Fortelius M, eds. Fossil mammals of Asia. New York: Columbia University Press, 400 - 422." type="book chapter" year="2013">
|
||
Antoine
|
||
<emphasis id="8487EAF8FFB99D1049B5FC68FCADB3FB" box="[715,773,902,924]" italics="true" pageId="22" pageNumber="300">et al.</emphasis>
|
||
, 2013
|
||
</bibRefCitation>
|
||
: fig. 16.4) and
|
||
<taxonomicName id="71F34D69FFB99D104AEDFC49FDD2B3DA" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[403,634,933,955]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFB99D104AEDFC49FDD2B3DA" box="[403,634,933,955]" italics="true" pageId="22" pageNumber="300">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
is certainly not documented in younger levels in the Bugti Hills. Nevertheless, a gap is observed in terms of fossilyielding levels in the concerned overlying sequence, i.e. between the poorly documented Level 6sup (‘Assemblage B’) and the Middle Miocene ‘Assemblage C’ (Level W;
|
||
<bibRefCitation id="D2624B1BFFB99D104A47FBB2FD8CB412" author="Antoine PO & Metais G & Orliac MJ & Crochet JY & Flynn LJ & Marivaux L & Rajpar AR & Roohi G & Welcomme JL" box="[313,548,1117,1139]" pageId="22" pageNumber="300" pagination="400 - 422" refId="ref23524" refString="Antoine PO, Metais G, Orliac MJ, Crochet JY, Flynn LJ, Marivaux L, Rajpar AR, Roohi G, Welcomme JL. 2013. Mammalian Neogene biostratigraphy of the Sulaiman Province, Pakistan. In: Xiaoming W, Flynn LJ, Fortelius M, eds. Fossil mammals of Asia. New York: Columbia University Press, 400 - 422." type="book chapter" year="2013">
|
||
Antoine
|
||
<emphasis id="8487EAF8FFB99D104ADEFBB1FE74B412" box="[416,476,1117,1139]" italics="true" pageId="22" pageNumber="300">et al.</emphasis>
|
||
, 2013
|
||
</bibRefCitation>
|
||
). In any event, the biochronological age of the Kutch fauna is potentially questionable (
|
||
<emphasis id="8487EAF8FFB99D104A2AFB74FE0AB4D0" box="[340,418,1179,1200]" italics="true" pageId="22" pageNumber="300">contra</emphasis>
|
||
<bibRefCitation id="D2624B1BFFB99D104AD3FB75FD1FB4CF" author="Bhandari A & Mohabey DM & Bajpai S & Tiwari BN & Pickford M" box="[429,695,1178,1200]" pageId="22" pageNumber="300" pagination="69 - 97" refId="ref23717" refString="Bhandari A, Mohabey DM, Bajpai S, Tiwari BN, Pickford M. 2010. Early Miocene mammals from central Kutch (Gujarat), Western India: implications for geochronology, biogeography, eustacy and intercontinental dispersals. Neues Jahrbuch fur Geologie und Palaontologie - Abhandlungen 256: 69 - 97." type="journal article" year="2010">
|
||
Bhandari
|
||
<emphasis id="8487EAF8FFB99D104952FB74FDCCB4CF" box="[556,612,1178,1200]" italics="true" pageId="22" pageNumber="300">et al</emphasis>
|
||
., 2010
|
||
</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="D2624B1BFFB99D1049B7FB75FCABB4CF" author="Bhandari A & Bajpai S & Flynn LJ & Tiwari BN & Mandal N" box="[713,771,1178,1199]" pageId="22" pageNumber="300" pagination="3471 - 3479" refId="ref23772" refString="Bhandari A, Bajpai S, Flynn LJ, Tiwari BN, Mandal N. 2021. First Miocene rodents from Kutch, western India. Historical Biology 33: 3471 - 3479." type="journal article" year="2021">2021</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="D2624B1BFFB99D104BDDFB56FE67B4AE" author="Sehgal RK & Bhandari A" box="[163,463,1209,1231]" pageId="22" pageNumber="300" pagination="199 - 212" refId="ref27104" refString="Sehgal RK, Bhandari A. 2014. Miocene mammals from India: present status and future prospects. In: Tiwari, RP, ed. Indian Miocene: a geodynamic and chronological framework for palaeobiota, sedimentary environments and palaeoclimates. Palaeontological Society of India Special Publication 5: 199 - 212." type="journal article" year="2014">Sehgal & Bhandari, 2014</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="D2624B1BFFB99D104AA1FB56FD53B4AF" author="Patnaik R & Prasad V" box="[479,763,1209,1231]" pageId="22" pageNumber="300" pagination="605 - 615" refId="ref26414" refString="Patnaik R, Prasad V. 2016. Neogene climate, terrestrial mammals and flora of the Indian subcontinent. Proceedings of the Indian National Science Academy 82: 605 - 615." type="journal article" year="2016">Patnaik & Prasad, 2016</bibRefCitation>
|
||
). Conversely, it may restrain the use of anthracotheriid species from the Bugti Hills for biochronological purposes (
|
||
<bibRefCitation id="D2624B1BFFB99D104B8EFAFAFDA1B54A" author="Antoine PO & Metais G & Orliac MJ & Crochet JY & Flynn LJ & Marivaux L & Rajpar AR & Roohi G & Welcomme JL" box="[240,521,1301,1323]" pageId="22" pageNumber="300" pagination="400 - 422" refId="ref23524" refString="Antoine PO, Metais G, Orliac MJ, Crochet JY, Flynn LJ, Marivaux L, Rajpar AR, Roohi G, Welcomme JL. 2013. Mammalian Neogene biostratigraphy of the Sulaiman Province, Pakistan. In: Xiaoming W, Flynn LJ, Fortelius M, eds. Fossil mammals of Asia. New York: Columbia University Press, 400 - 422." type="book chapter" year="2013">
|
||
Antoine
|
||
<emphasis id="8487EAF8FFB99D104A2CFAF9FE23B54A" box="[338,395,1301,1323]" italics="true" pageId="22" pageNumber="300">et al.</emphasis>
|
||
, 2013: 413
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="B64C36EAFFB99D104BC5FADBFBB7B209" blockId="22.[163,780,197,1905]" lastBlockId="22.[827,1444,197,1445]" pageId="22" pageNumber="300">
|
||
<bibRefCitation id="D2624B1BFFB99D104BC5FADBFE61B529" author="Russell DE & Zhai RJ" box="[187,457,1332,1354]" pageId="22" pageNumber="300" pagination="1 - 490" refId="ref26996" refString="Russell DE, Zhai RJ. 1987. The Paleogene of Asia: mammals and stratigraphy. Memoires du Museum National d'Histoire Naturelle 52: 1 - 490." type="journal article" year="1987">Russell & Zhai (1987)</bibRefCitation>
|
||
reported one P4/ and one M/2 from the Benara fauna in
|
||
<collectingCountry id="CEE4767AFFB99D104960FABDFDD4B508" box="[542,636,1362,1384]" name="Georgia" pageId="22" pageNumber="300">Georgia</collectingCountry>
|
||
(
|
||
<bibRefCitation id="D2624B1BFFB99D1049F1FABDFF74B5E7" author="Gabounia L" pageId="22" pageNumber="300" pagination="857 - 869" refId="ref24434" refString="Gabounia L. 1966. Sur les mammiferes oligocenes du Caucase. Bulletin de la Societe Geologique de France 8: 857 - 869." type="journal article" year="1966">Gabounia, 1966</bibRefCitation>
|
||
: fig. 9d, e) referred to as
|
||
<taxonomicName id="71F34D69FFB99D104970FA9EFD66B5E6" authorityName="Forster-Cooper" authorityYear="1915" box="[526,718,1393,1414]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB99D104970FA9EFD66B5E6" box="[526,718,1393,1414]" italics="true" pageId="22" pageNumber="300">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
. The upper premolar lacks the two distal ridges and large distostyle characteristic of this genus, and it is more oblong transversely. The lower molar is too small to belong to this taxon; its selenodont morphology with pinched lingual cuspids and the connection between the preprotocristid and the premetacristid are consistent with
|
||
<taxonomicName id="71F34D69FFB99D104A46F9A7FE0EB63D" baseAuthorityName="Hellmund" baseAuthorityYear="1991" box="[312,422,1608,1629]" class="Mammalia" family="Anthracotheriidae" genus="Elomeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB99D104A46F9A7FE0EB63D" box="[312,422,1608,1629]" italics="true" pageId="22" pageNumber="300">Elomeryx</emphasis>
|
||
</taxonomicName>
|
||
instead. According to the provided drawing, it could also be a worn, ruminant-like lower tooth. A fragmentary pentacuspidate upper molar from the Dingdanggou fauna (
|
||
<collectingCountry id="CEE4767AFFB99D104933F94BFD3DB6DA" box="[589,661,1700,1722]" name="China" pageId="22" pageNumber="300">China</collectingCountry>
|
||
) has been interpreted as
|
||
<taxonomicName id="71F34D69FFB99D104A1EF92DFDFDB6B8" box="[352,597,1730,1752]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="species" species="undetermined">
|
||
<emphasis id="8487EAF8FFB99D104A1EF92DFD80B6B7" box="[352,552,1730,1751]" italics="true" pageId="22" pageNumber="300">Parabrachyodus</emphasis>
|
||
sp.
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation id="D2624B1BFFB99D104917F92CFF75B696" author="Wang BY & Qiu ZX" pageId="22" pageNumber="300" pagination="130 - 143" refId="ref27373" refString="Wang BY, Qiu ZX. 2004. Discovery of Early Oligocene mammalian fossils form Danghe Area, Gansu, China. Vertebrata PalAsiatica 42: 130 - 143." type="journal article" year="2004">Wang & Qiu, 2004</bibRefCitation>
|
||
), but it has more selenodont ridges than unambiguous representatives of this genus, and the postparacristule joins the base of the paracone rather than the transverse valley. In this context, it is more likely that this tooth belongs to
|
||
<taxonomicName id="71F34D69FFB99D104924F8B3FCA3B710" baseAuthorityName="Ducrocq & Lihoreau" baseAuthorityYear="2006" box="[602,779,1883,1905]" class="Mammalia" family="Anthracotheriidae" genus="Sivameryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="species" species="palaeindicus">
|
||
<emphasis id="8487EAF8FFB99D104924F8B3FCA3B710" box="[602,779,1883,1905]" italics="true" pageId="22" pageNumber="300">S. palaeindicus</emphasis>
|
||
</taxonomicName>
|
||
than to
|
||
<taxonomicName id="71F34D69FFB99D1048E7FF29FBD4B0BA" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[921,1148,197,219]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFB99D1048E7FF29FBD4B0BA" box="[921,1148,197,219]" italics="true" pageId="22" pageNumber="300">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
. Hence, no occurrence of
|
||
<taxonomicName id="71F34D69FFB99D104845FF0CFC5DB098" authorityName="Forster-Cooper" authorityYear="1915" box="[827,1013,227,248]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB99D104845FF0CFC5DB098" box="[827,1013,227,248]" italics="true" pageId="22" pageNumber="300">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
is documented in Oligocene–Miocene deposits of
|
||
<collectingCountry id="CEE4767AFFB99D1048C1FEEDFBA0B178" box="[959,1032,258,280]" name="China" pageId="22" pageNumber="300">China</collectingCountry>
|
||
, and there is no dispersal event either between the north and south sides of the Tibetan Plateau involving this bothriodontine during the Early Oligocene (
|
||
<emphasis id="8487EAF8FFB99D1048C4FEB0FBABB114" box="[954,1027,351,372]" italics="true" pageId="22" pageNumber="300">contra</emphasis>
|
||
<bibRefCitation id="D2624B1BFFB99D104F74FEB1FB0EB114" author="Li ZC & Li YX & Zhang YX & Li WH & Xie K" box="[1034,1190,350,372]" pageId="22" pageNumber="300" pagination="1258 - 1266" refId="ref25017" refString="Li ZC, Li YX, Zhang YX, Li WH, Xie K. 2016. Nanpoping fauna of the Lanzhou Basin and its environmental significance. Science China Earth Sciences 59: 1258 - 1266." type="journal article" year="2016">
|
||
Li
|
||
<emphasis id="8487EAF8FFB99D104F57FEB0FBCAB113" box="[1065,1122,350,372]" italics="true" pageId="22" pageNumber="300">et al.</emphasis>
|
||
, 2016
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="D2624B1BFFB99D104FCDFEB0FA9FB113" author="Wang SQ" box="[1203,1335,350,372]" pageId="22" pageNumber="300" pagination="839 - 852" refId="ref27342" refString="Wang SQ. 2020. The anthracotheres from northern Junggar Basin and their palaeoclimatic significance in relation to the Tibetan Plateau. PalaeobiodiVersity and PalaeoenVironments 101: 839 - 852." type="journal article" year="2020">Wang, 2020</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="D2624B1BFFB99D104E3BFEB1FCDCB1F2" author="Li C & Wang SQ & Yang Q" pageId="22" pageNumber="300" pagination="1 - 9" refId="ref25055" refString="Li C, Wang SQ, Yang Q. 2022. Discovery of a primitive Gomphotherium from the Early Miocene of northern China and its biochronology and palaeobiogeography significance. Historical Biology 1 - 9." type="book chapter" year="2022">
|
||
Li
|
||
<emphasis id="8487EAF8FFB99D104E1AFEB0FA35B113" box="[1380,1437,350,372]" italics="true" pageId="22" pageNumber="300">et al.</emphasis>
|
||
, 2022
|
||
</bibRefCitation>
|
||
). The occurrence of
|
||
<taxonomicName id="71F34D69FFB99D104F16FE92FAFBB1F2" box="[1128,1363,381,402]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="species" species="undetermined">
|
||
<emphasis id="8487EAF8FFB99D104F16FE92FA81B1F2" box="[1128,1321,381,402]" italics="true" pageId="22" pageNumber="300">Parabrachyodus</emphasis>
|
||
sp.
|
||
</taxonomicName>
|
||
in the
|
||
<collectingRegion id="7437F808FFB99D104845FE74FC11B1D1" box="[827,953,411,433]" country="Myanmar" name="Ayeyawady" pageId="22" pageNumber="300">Irrawaddy</collectingRegion>
|
||
area of
|
||
<collectingCountry id="CEE4767AFFB99D104F5EFE73FB3DB1D1" box="[1056,1173,412,433]" name="Myanmar" pageId="22" pageNumber="300">Myanmar</collectingCountry>
|
||
(
|
||
<collectingCountry id="CEE4767AFFB99D104FD8FE73FB52B1D1" box="[1190,1274,412,433]" name="Myanmar" pageId="22" pageNumber="300">Burma</collectingCountry>
|
||
) immediately east of the Indian subcontinent (
|
||
<bibRefCitation id="D2624B1BFFB99D104FD3FE55FA33B1AF" author="Bhandari A & Mohabey DM & Bajpai S & Tiwari BN & Pickford M" box="[1197,1435,442,464]" pageId="22" pageNumber="300" pagination="69 - 97" refId="ref23717" refString="Bhandari A, Mohabey DM, Bajpai S, Tiwari BN, Pickford M. 2010. Early Miocene mammals from central Kutch (Gujarat), Western India: implications for geochronology, biogeography, eustacy and intercontinental dispersals. Neues Jahrbuch fur Geologie und Palaontologie - Abhandlungen 256: 69 - 97." type="journal article" year="2010">
|
||
Bhandari
|
||
<emphasis id="8487EAF8FFB99D104E61FE54FAFFB1AF" box="[1311,1367,442,464]" italics="true" pageId="22" pageNumber="300">et al.</emphasis>
|
||
, 2010
|
||
</bibRefCitation>
|
||
) remains uncertain due to the lack of illustration for the referred specimen. The provided measurements of the concerned M3/ (46.5 × 54.9 mm) fall outside the range of variation defined for
|
||
<taxonomicName id="71F34D69FFB99D104FDEFDDAFA2BB22A" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[1184,1411,565,586]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFB99D104FDEFDDAFA2BB22A" box="[1184,1411,565,586]" italics="true" pageId="22" pageNumber="300">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
in this study (
|
||
<tableCitation id="FB710351FFB99D1048C2FDBCFBA7B209" box="[956,1039,595,617]" captionStart="Table 2" captionStartId="12.[164,228,196,217]" captionTargetPageId="12" captionText="Table 2. Measurements of upper jugal teeth of Parabrachyodus hyopotamoides (in mm) from the historical quarry sites ‘Dera Bugti’, ‘Chur Lando’ (= Lundo Chur), ‘Bugti’, ‘Kumbi’ (only those conserved in the NHM) and from Samane Nala. Abbreviations: N, number of specimens measured; Lmd, mesiodistal length; Lmd, mesiodistal lengths; Lll1-3, labiolingual lengths; SD, standard deviation; CV, unbiased coefficient of variation (%); W, Shapiro–Wilk test (ns, normal distribution cannot be rejected; s, normal distribution is rejected; P <0.05); t-test, student test between mesiodistal length (Lmd)" httpUri="http://table.plazi.org/id/E28C6662FFA39D0A4BDAFF2BFE2DB237" pageId="22" pageNumber="300" tableUuid="E28C6662FFA39D0A4BDAFF2BFE2DB237">Table 2</tableCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="B64C36EAFFB99D10482DFD9CFBFDB5C5" blockId="22.[827,1444,197,1445]" pageId="22" pageNumber="300">
|
||
In the current stage of our knowledge,
|
||
<taxonomicName id="71F34D69FFB99D104E0CFD9CFC40B2C6" baseAuthorityName="Lydekker" baseAuthorityYear="1883" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFB99D104E0CFD9CFC40B2C6" italics="true" pageId="22" pageNumber="300">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
seems to be restricted to the western part of the Indian subcontinent and it is not formally known in the Palaeogene, in contrast with what the compilation of Sulaiman Range faunas studied by
|
||
<bibRefCitation id="D2624B1BFFB99D104845FCE3FB9FB341" author="Raza SM & Meyer GE" box="[827,1079,780,801]" pageId="22" pageNumber="300" pagination="43 - 63" refId="ref26783" refString="Raza SM, Meyer GE. 1984. Early Miocene geology and paleontology of the Bugti Hills. Geological SurVey of Pakistan 11: 43 - 63." type="journal article" year="1984">Raza & Meyer (1984)</bibRefCitation>
|
||
and
|
||
<bibRefCitation id="D2624B1BFFB99D104F0AFCE4FA80B341" author="Pickford M" box="[1140,1320,779,801]" pageId="22" pageNumber="300" pagination="289 - 350" refId="ref26447" refString="Pickford M. 1987. Revision des suiformes (Artiodactyla, Mammalia) de Bugti (Pakistan). Annales de Paleontologie 73: 289 - 350." type="journal article" year="1987">Pickford (1987)</bibRefCitation>
|
||
may have suggested. Despite the ‘gigantic
|
||
<taxonomicName id="71F34D69FFB99D104FC8FCC4FAE0B320" box="[1206,1352,811,832]" class="Mammalia" family="Anthracotheriidae" genus="Hyopotamus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB99D104FC8FCC4FAE0B320" box="[1206,1352,811,832]" italics="true" pageId="22" pageNumber="300">Hyopotamus</emphasis>
|
||
</taxonomicName>
|
||
’ known ‘from Sind’ (
|
||
<bibRefCitation id="D2624B1BFFB99D1048B4FCA6FB1EB33E" author="Lydekker R" box="[970,1206,841,863]" pageId="22" pageNumber="300" pagination="107" refId="ref25726" refString="Lydekker R. 1882. Note on some Siwalik and Narbada fossils. 7. Gigantic Hyopotamus. Records of the Geological SurVey of India 15: 107." type="journal article" year="1882">Lydekker, 1882: 107</bibRefCitation>
|
||
),
|
||
<taxonomicName id="71F34D69FFB99D104FB5FCA6FA20B33E" authorityName="Forster-Cooper" authorityYear="1915" box="[1227,1416,841,862]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB99D104FB5FCA6FA20B33E" box="[1227,1416,841,862]" italics="true" pageId="22" pageNumber="300">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
is not mentioned in the Manchar Formation, with maybe the exception of ‘cf.
|
||
<taxonomicName id="71F34D69FFB99D104F5CFC69FB7BB3FB" box="[1058,1235,902,923]" class="Mammalia" family="Anthracotheriidae" genus="Brachyodus" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="species" species="undetermined">
|
||
<emphasis id="8487EAF8FFB99D104F5CFC69FB03B3FB" box="[1058,1195,902,923]" italics="true" pageId="22" pageNumber="300">Brachyodus</emphasis>
|
||
sp.
|
||
</taxonomicName>
|
||
’ listed by Raza
|
||
<emphasis id="8487EAF8FFB99D104EF1FC68FCF0B3DA" italics="true" pageId="22" pageNumber="300">et al.</emphasis>
|
||
(1984: table 2) in its lower member. Given that the
|
||
<collectingRegion id="7437F808FFB99D104845FC2CFCD9B3B9" box="[827,881,963,985]" country="Pakistan" name="Sindh" pageId="22" pageNumber="300">Sind</collectingRegion>
|
||
deposits lie south of the Bugti Hills and north of the Kutch Province (
|
||
<bibRefCitation id="D2624B1BFFB99D104F2EFC0DFAE3B397" author="Bhandari A & Mohabey DM & Bajpai S & Tiwari BN & Pickford M" box="[1104,1355,994,1016]" pageId="22" pageNumber="300" pagination="69 - 97" refId="ref23717" refString="Bhandari A, Mohabey DM, Bajpai S, Tiwari BN, Pickford M. 2010. Early Miocene mammals from central Kutch (Gujarat), Western India: implications for geochronology, biogeography, eustacy and intercontinental dispersals. Neues Jahrbuch fur Geologie und Palaontologie - Abhandlungen 256: 69 - 97." type="journal article" year="2010">
|
||
Bhandari
|
||
<emphasis id="8487EAF8FFB99D104FB6FC0CFAACB397" box="[1224,1284,994,1016]" italics="true" pageId="22" pageNumber="300">et al.</emphasis>
|
||
, 2010
|
||
</bibRefCitation>
|
||
: fig. 1), fossils of
|
||
<taxonomicName id="71F34D69FFB99D1048D4FBEEFB25B476" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[938,1165,1025,1046]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFB99D1048D4FBEEFB25B476" box="[938,1165,1025,1046]" italics="true" pageId="22" pageNumber="300">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
are also likely to occur in the Manchar Formation. Fossil collections without any stratigraphical context in the Bugti Hills where Oligocene sediments have been identified (M12030 and M12033) are suspected to somewhat pre-date the Oligocene–Miocene transition (
|
||
<bibRefCitation id="D2624B1BFFB99D104FD7FB75FA34B4CF" author="Forster-Cooper C" box="[1193,1436,1178,1199]" pageId="22" pageNumber="300" pagination="514 - 522" refId="ref24343" refString="Forster-Cooper C. 1913. New anthracotheres and allied forms from Baluchistan - preliminary notice. Annals and Magazine of Natural History 12: 514 - 522." type="journal article" year="1913">Forster-Cooper, 1913</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="D2624B1BFFB99D104845FB56FB88B4AE" author="Antoine PO & Metais G & Orliac MJ & Crochet JY & Flynn LJ & Marivaux L & Rajpar AR & Roohi G & Welcomme JL" box="[827,1056,1209,1231]" pageId="22" pageNumber="300" pagination="400 - 422" refId="ref23524" refString="Antoine PO, Metais G, Orliac MJ, Crochet JY, Flynn LJ, Marivaux L, Rajpar AR, Roohi G, Welcomme JL. 2013. Mammalian Neogene biostratigraphy of the Sulaiman Province, Pakistan. In: Xiaoming W, Flynn LJ, Fortelius M, eds. Fossil mammals of Asia. New York: Columbia University Press, 400 - 422." type="book chapter" year="2013">
|
||
Antoine
|
||
<emphasis id="8487EAF8FFB99D1048E1FB56FC72B4AE" box="[927,986,1209,1230]" italics="true" pageId="22" pageNumber="300">et al.</emphasis>
|
||
, 2013
|
||
</bibRefCitation>
|
||
: fig. 16.4) as those referred to as
|
||
<taxonomicName id="71F34D69FFB99D104845FB37FB9CB48C" box="[827,1076,1239,1261]" class="Mammalia" family="Anthracotheriidae" genus="Hemimeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="species" species="blanfordi">
|
||
<emphasis id="8487EAF8FFB99D104845FB37FB9CB48C" box="[827,1076,1239,1261]" italics="true" pageId="22" pageNumber="300">Hemimeryx blanfordi</emphasis>
|
||
</taxonomicName>
|
||
have been in the same work of Forster-Cooper (
|
||
<bibRefCitation id="D2624B1BFFB99D10488DFB19FB74B56C" author="Lihoreau F & Alloing-Seguier L & Antoine PO & Boisserie JR & Marivaux L & Metais G & Welcomme JL" box="[1011,1244,1270,1292]" pageId="22" pageNumber="300" pagination="947 - 957" refId="ref25524" refString="Lihoreau F, Alloing-Seguier L, Antoine PO, Boisserie JR, Marivaux L, Metais G, Welcomme JL. 2016. Enamel microstructure defines a major Paleogene hippopotamoid clade: the Merycopotamini (Cetartiodactyla, Hippopotamoidea). Historical Biology 29: 947 - 957." type="journal article" year="2016">
|
||
Lihoreau
|
||
<emphasis id="8487EAF8FFB99D104F1FFB18FB31B56B" box="[1121,1177,1270,1292]" italics="true" pageId="22" pageNumber="300">et al.</emphasis>
|
||
, 2016
|
||
</bibRefCitation>
|
||
). This occurrence of
|
||
<taxonomicName id="71F34D69FFB99D104826FAFAFBA1B54A" box="[856,1033,1301,1322]" class="Mammalia" family="Anthracotheriidae" genus="Hemimeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="species" species="blanfordi">
|
||
<emphasis id="8487EAF8FFB99D104826FAFAFBA1B54A" box="[856,1033,1301,1322]" italics="true" pageId="22" pageNumber="300">Hem. blanfordi</emphasis>
|
||
</taxonomicName>
|
||
in the Late Oligocene of the Bugti Hills, together with the basal phylogenetic position of the clade (
|
||
<taxonomicName id="71F34D69FFB99D1048CDFABCFB39B507" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[947,1169,1362,1384]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFB99D1048CDFABCFB39B507" box="[947,1169,1362,1384]" italics="true" pageId="22" pageNumber="300">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
+
|
||
<taxonomicName id="71F34D69FFB99D104FD3FABDFA9CB507" baseAuthorityName="Forster-Cooper" baseAuthorityYear="1924" box="[1197,1332,1362,1383]" class="Mammalia" family="Anthracotheriidae" genus="Telmatodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB99D104FD3FABDFA9CB507" box="[1197,1332,1362,1383]" italics="true" pageId="22" pageNumber="300">Telmatodon</emphasis>
|
||
</taxonomicName>
|
||
), support the potential appearance of
|
||
<taxonomicName id="71F34D69FFB99D104FFCFA9EFA96B5E6" authorityName="Forster-Cooper" authorityYear="1915" box="[1154,1342,1393,1414]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB99D104FFCFA9EFA96B5E6" box="[1154,1342,1393,1414]" italics="true" pageId="22" pageNumber="300">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
as early as in the Late Oligocene.
|
||
</paragraph>
|
||
<paragraph id="B64C36EAFFB99D1048FCFA0DFACCB678" blockId="22.[889,1380,1506,1562]" pageId="22" pageNumber="300">
|
||
<smallCapsWord id="B0AAA036FFB99D1048FCFA0DFB99B59A" baselines="1525,1525" box="[898,1073,1506,1531]" lowerCaseFontSize="8" mainFontSize="10" normCase="title" normString="Phylogenetic" pageId="22" pageNumber="300">PHYLOGENETIC</smallCapsWord>
|
||
<smallCapsWord id="B0AAA036FFB99D104F46FA09FB4FB59A" baselines="1525" box="[1080,1255,1510,1530]" lowerCaseFontSize="8" mainFontSize="10" normCase="lower" normString="relationships" pageId="22" pageNumber="300">RELATIONSHIPS</smallCapsWord>
|
||
<smallCapsWord id="B0AAA036FFB99D104F91FA09FAF3B59A" baselines="1525" box="[1263,1371,1510,1530]" lowerCaseFontSize="8" mainFontSize="10" normCase="lower" normString="between" pageId="22" pageNumber="300">BETWEEN</smallCapsWord>
|
||
<smallCapsWord id="B0AAA036FFB99D104807F9EBFBE6B678" baselines="1555" box="[889,1102,1540,1560]" lowerCaseFontSize="8" mainFontSize="10" normCase="lower" normString="bothriodontines" pageId="22" pageNumber="300">BOTHRIODONTINES</smallCapsWord>
|
||
<smallCapsWord id="B0AAA036FFB99D104F28F9EBFB3DB678" baselines="1555" box="[1110,1173,1540,1560]" lowerCaseFontSize="8" mainFontSize="10" normCase="lower" normString="from" pageId="22" pageNumber="300">FROM</smallCapsWord>
|
||
<smallCapsWord id="B0AAA036FFB99D104FE2F9EBFB61B678" baselines="1555" box="[1180,1225,1540,1560]" lowerCaseFontSize="8" mainFontSize="10" normCase="lower" normString="the" pageId="22" pageNumber="300">THE</smallCapsWord>
|
||
<smallCapsWord id="B0AAA036FFB99D104FAEF9EEFAB0B678" baselines="1556,1555" box="[1232,1304,1537,1562]" lowerCaseFontSize="8" mainFontSize="10" normCase="title" normString="Bugti" pageId="22" pageNumber="300">BUGTI</smallCapsWord>
|
||
<smallCapsWord id="B0AAA036FFB99D104E61F9EEFACCB678" baselines="1556,1555" box="[1311,1380,1537,1562]" lowerCaseFontSize="8" mainFontSize="10" normCase="title" normString="Hills" pageId="22" pageNumber="300">HILLS</smallCapsWord>
|
||
</paragraph>
|
||
<paragraph id="B64C36EAFFB99D114845F9C6FE75B48D" blockId="22.[827,1443,1577,1906]" lastBlockId="23.[145,762,197,1905]" lastPageId="23" lastPageNumber="301" pageId="22" pageNumber="300">
|
||
The three genera,
|
||
<taxonomicName id="71F34D69FFB99D104F65F9C6FB33B65E" authorityName="Pilgrim" authorityYear="1908" box="[1051,1179,1577,1598]" class="Mammalia" family="Anthracotheriidae" genus="Gonotelma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB99D104F65F9C6FB33B65E" box="[1051,1179,1577,1598]" italics="true" pageId="22" pageNumber="300">Gonotelma</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="71F34D69FFB99D104FD5F9C6FAC5B65E" authorityName="Forster-Cooper" authorityYear="1915" box="[1195,1389,1577,1598]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB99D104FD5F9C6FAC5B65E" box="[1195,1389,1577,1598]" italics="true" pageId="22" pageNumber="300">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="71F34D69FFB99D104845F9A7FC66B63D" baseAuthorityName="Forster-Cooper" baseAuthorityYear="1924" box="[827,974,1608,1629]" class="Mammalia" family="Anthracotheriidae" genus="Telmatodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB99D104845F9A7FC66B63D" box="[827,974,1608,1629]" italics="true" pageId="22" pageNumber="300">Telmatodon</emphasis>
|
||
</taxonomicName>
|
||
, recorded in the Bugti Hills can be seen as early
|
||
<taxonomicName id="71F34D69FFB99D10489EF989FC2EB6FB" authority="sensu Lihoreau et al. (2016)" authorityName="Lihoreau" authorityYear="2016" pageId="22" pageNumber="304" rank="tribe" tribe="Merycopotamini">
|
||
Merycopotamini
|
||
<emphasis id="8487EAF8FFB99D104FD7F988FB42B61C" box="[1193,1258,1639,1660]" italics="true" pageId="22" pageNumber="300">sensu</emphasis>
|
||
<bibRefCitation id="D2624B1BFFB99D104F8DF989FC2EB6FB" author="Lihoreau F & Alloing-Seguier L & Antoine PO & Boisserie JR & Marivaux L & Metais G & Welcomme JL" pageId="22" pageNumber="300" pagination="947 - 957" refId="ref25524" refString="Lihoreau F, Alloing-Seguier L, Antoine PO, Boisserie JR, Marivaux L, Metais G, Welcomme JL. 2016. Enamel microstructure defines a major Paleogene hippopotamoid clade: the Merycopotamini (Cetartiodactyla, Hippopotamoidea). Historical Biology 29: 947 - 957." type="journal article" year="2016">
|
||
Lihoreau
|
||
<emphasis id="8487EAF8FFB99D104E19F988FA0BB61B" box="[1383,1443,1638,1660]" italics="true" pageId="22" pageNumber="300">et al.</emphasis>
|
||
(2016)
|
||
</bibRefCitation>
|
||
</taxonomicName>
|
||
in the present study. We refute the hypothesis that
|
||
<taxonomicName id="71F34D69FFB99D1048CDF94BFB19B6D9" authorityName="Pilgrim" authorityYear="1908" box="[947,1201,1700,1721]" class="Mammalia" family="Anthracotheriidae" genus="Gonotelma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="species" species="shahbazi">
|
||
<emphasis id="8487EAF8FFB99D1048CDF94BFB19B6D9" box="[947,1201,1700,1721]" italics="true" pageId="22" pageNumber="300">Gonotelma shahbazi</emphasis>
|
||
</taxonomicName>
|
||
is more closely related to
|
||
<taxonomicName id="71F34D69FFB99D1048E0F92CFBB2B6B8" box="[926,1050,1731,1752]" class="Mammalia" family="Anthracotheriidae" genus="Sivameryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB99D1048E0F92CFBB2B6B8" box="[926,1050,1731,1752]" italics="true" pageId="22" pageNumber="300">SiƲameryx</emphasis>
|
||
</taxonomicName>
|
||
than to
|
||
<taxonomicName id="71F34D69FFB99D104FFFF92CFAE7B6B7" authorityName="Pickford" authorityYear="1991" box="[1153,1359,1730,1752]" class="Mammalia" family="Anthracotheriidae" genus="Afromeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="species" species="zelteni">
|
||
<emphasis id="8487EAF8FFB99D104FFFF92CFAE7B6B7" box="[1153,1359,1730,1752]" italics="true" pageId="22" pageNumber="300">Afromeryx zelteni</emphasis>
|
||
</taxonomicName>
|
||
due to the shared retention of the paraconule (
|
||
<bibRefCitation id="D2624B1BFFB99D104E64F90EFC72B775" author="Lihoreau F & Ducrocq S" pageId="22" pageNumber="300" pagination="89 - 105" refId="ref25128" refString="Lihoreau F, Ducrocq S. 2007. Family Anthracotheriidae. In: Protero DR, Foss SE, eds. The eVolution of artiodactyls. Baltimore: John Hopkins University Press, 89 - 105." type="book chapter" year="2007">Lihoreau & Ducrocq, 2007</bibRefCitation>
|
||
). First, the absence of a postparacristule on the M3/ of
|
||
<taxonomicName id="71F34D69FFB99D104893F8F0FBD7B753" authorityName="Pilgrim" authorityYear="1908" box="[1005,1151,1822,1844]" class="Mammalia" family="Anthracotheriidae" genus="Gonotelma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="species" species="shahbazi">
|
||
<emphasis id="8487EAF8FFB99D104893F8F0FBD7B753" box="[1005,1151,1822,1844]" italics="true" pageId="22" pageNumber="300">G. shahbazi</emphasis>
|
||
</taxonomicName>
|
||
shows that there is no paraconule (
|
||
<figureCitation id="2EC82A6FFFB99D1048A8F8D2FB84B733" box="[982,1068,1853,1875]" captionStart="Figure 5" captionStartId="10.[163,241,1448,1470]" captionTargetBox="[243,1362,198,1415]" captionTargetId="figure-25@10.[240,1366,195,1418]" captionTargetPageId="10" captionText="Figure 5. Upper teeth of Parabrachyodus hyopotamoides. A, left upper incisor (I2/ or I3/) in lingual (left) and labial (right) view (UM-SAM4-024). B, P1/-P4/ from a left maxilla (M12034) in occlusal view. C, right P1/ (UM-SAM4-025) in occlusal view (mirror). D, left P3/ (M10598) in occlusal view. E, left maxilla with P4/-M1/ in occlusal view, with a drawing of P4/. F, left dP3/ (M12820) in occlusal view. Abbreviations derived from the nomenclature introduced in Figure 2: protoc., protocrista. G, dental variation of Parabrachyodus hyopotamoides figured on composite upper molars (N = 26 for M3/; N = 11 for M2/). Variable characters are coloured. Character frequency: black: 100%; red: (75–99%); yellow: (50–74%); green: (25–49%); blue: (0–24%). The presence/absence of the coloured character is evaluated. 1, mesostyle divided into two slender apices; 2, parastyle of the same size or smaller than mesostyle; 3, wing-like labial cingulum; 4, postectoprotocrista reaching the lingual edge of the tooth; 5, double curvature of the postmetacristule. The arrow indicates the mesiolingual direction. Teeth are not to scale. H, left maxilla with M1/-M3/ (M12718) in occlusal view. I, right maxilla with P4/-M3/ (M12824) in occlusal view (mirror). Comparisons with drawings of (J) left M3/ of Brachyodus onoideus in occlusal view, modified from Ducrocq & Lihoreau (2006); K, left M3/ of Telmatodon orientalis (M12040, holotype) in occlusal view; L, right M3/ of Gonotelma shahbazi (UM-SAM4-010) in occlusal view (mirror); M, left M3/ of SiƲameryx palaeindicus (UM-SAM4-005) in occlusal view. N, M3/ of Par. hyopotamoides (UM-SAM5-001) in distal view, showing the low value of the angle between the lingual slope of the metaconule and the socket. Scale bars all equal 10 mm." figureDoi="http://doi.org/10.5281/zenodo.7927073" httpUri="https://zenodo.org/record/7927073/files/figure.png" pageId="22" pageNumber="300">Fig. 5L</figureCitation>
|
||
). In comparison, the
|
||
<typeStatus id="69488848FFB99D104E45F8D2FA0AB733" box="[1339,1442,1853,1875]" pageId="22" pageNumber="300">holotype</typeStatus>
|
||
of
|
||
<taxonomicName id="71F34D69FFB99D104827F8B3FAD8B711" authority="(Forster-Cooper, 1924)" baseAuthorityName="Forster-Cooper" baseAuthorityYear="1924" box="[857,1392,1884,1905]" class="Mammalia" family="Anthracotheriidae" genus="Telmatodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="22" pageNumber="300" phylum="Chordata" rank="species" species="orientalis">
|
||
<emphasis id="8487EAF8FFB99D104827F8B3FBF6B711" box="[857,1118,1884,1905]" italics="true" pageId="22" pageNumber="300">Telmatodon orientalis</emphasis>
|
||
(
|
||
<bibRefCitation id="D2624B1BFFB99D104F0EF8B3FAC0B711" author="Forster-Cooper C" box="[1136,1384,1884,1905]" pageId="22" pageNumber="300" pagination="1 - 59" refId="ref24403" refString="Forster-Cooper C. 1924. The Anthracotheriidae of the Dera Bugti deposits in Baluchistan. Memoirs of the Geological SurVey of India. Palaeontologica Indica 4: 1 - 59." type="journal article" year="1924">Forster-Cooper, 1924</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
has a reduced paraconule with a vestigial cristule (
|
||
<figureCitation id="2EC82A6FFFB89D1149B2FF2AFF1AB09A" captionStart="Figure 5" captionStartId="10.[163,241,1448,1470]" captionTargetBox="[243,1362,198,1415]" captionTargetId="figure-25@10.[240,1366,195,1418]" captionTargetPageId="10" captionText="Figure 5. Upper teeth of Parabrachyodus hyopotamoides. A, left upper incisor (I2/ or I3/) in lingual (left) and labial (right) view (UM-SAM4-024). B, P1/-P4/ from a left maxilla (M12034) in occlusal view. C, right P1/ (UM-SAM4-025) in occlusal view (mirror). D, left P3/ (M10598) in occlusal view. E, left maxilla with P4/-M1/ in occlusal view, with a drawing of P4/. F, left dP3/ (M12820) in occlusal view. Abbreviations derived from the nomenclature introduced in Figure 2: protoc., protocrista. G, dental variation of Parabrachyodus hyopotamoides figured on composite upper molars (N = 26 for M3/; N = 11 for M2/). Variable characters are coloured. Character frequency: black: 100%; red: (75–99%); yellow: (50–74%); green: (25–49%); blue: (0–24%). The presence/absence of the coloured character is evaluated. 1, mesostyle divided into two slender apices; 2, parastyle of the same size or smaller than mesostyle; 3, wing-like labial cingulum; 4, postectoprotocrista reaching the lingual edge of the tooth; 5, double curvature of the postmetacristule. The arrow indicates the mesiolingual direction. Teeth are not to scale. H, left maxilla with M1/-M3/ (M12718) in occlusal view. I, right maxilla with P4/-M3/ (M12824) in occlusal view (mirror). Comparisons with drawings of (J) left M3/ of Brachyodus onoideus in occlusal view, modified from Ducrocq & Lihoreau (2006); K, left M3/ of Telmatodon orientalis (M12040, holotype) in occlusal view; L, right M3/ of Gonotelma shahbazi (UM-SAM4-010) in occlusal view (mirror); M, left M3/ of SiƲameryx palaeindicus (UM-SAM4-005) in occlusal view. N, M3/ of Par. hyopotamoides (UM-SAM5-001) in distal view, showing the low value of the angle between the lingual slope of the metaconule and the socket. Scale bars all equal 10 mm." figureDoi="http://doi.org/10.5281/zenodo.7927073" httpUri="https://zenodo.org/record/7927073/files/figure.png" pageId="23" pageNumber="301">Fig. 5K</figureCitation>
|
||
). Second, the presence of the paraconule is here considered as a derived character that appeared independently in
|
||
<taxonomicName id="71F34D69FFB89D114A37FECEFE6CB156" box="[329,452,289,310]" class="Mammalia" family="Anthracotheriidae" genus="Sivameryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114A37FECEFE6CB156" box="[329,452,289,310]" italics="true" pageId="23" pageNumber="301">SiƲameryx</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="71F34D69FFB89D11497EFECEFD16B156" authorityName="Forster-Cooper" authorityYear="1915" box="[512,702,289,310]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D11497EFECEFD16B156" box="[512,702,289,310]" italics="true" pageId="23" pageNumber="301">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
. The position of
|
||
<taxonomicName id="71F34D69FFB89D114A70FEAFFE21B135" authorityName="Pilgrim" authorityYear="1908" box="[270,393,320,341]" class="Mammalia" family="Anthracotheriidae" genus="Gonotelma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114A70FEAFFE21B135" box="[270,393,320,341]" italics="true" pageId="23" pageNumber="301">Gonotelma</emphasis>
|
||
</taxonomicName>
|
||
as sister-group to
|
||
<taxonomicName id="71F34D69FFB89D114923FEAFFD7AB135" authorityName="Pickford" authorityYear="1991" box="[605,722,320,341]" class="Mammalia" family="Anthracotheriidae" genus="Afromeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114923FEAFFD7AB135" box="[605,722,320,341]" italics="true" pageId="23" pageNumber="301">Afromeryx</emphasis>
|
||
</taxonomicName>
|
||
instead supports the hypothesis of closer phylogenetic relationships with this small Libyan merycopotamine (
|
||
<bibRefCitation id="D2624B1BFFB89D114BE7FE73FEE9B1D1" author="Pickford M" box="[153,321,412,434]" pageId="23" pageNumber="301" pagination="289 - 350" refId="ref26447" refString="Pickford M. 1987. Revision des suiformes (Artiodactyla, Mammalia) de Bugti (Pakistan). Annales de Paleontologie 73: 289 - 350." type="journal article" year="1987">Pickford, 1987</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="D2624B1BFFB89D114A2EFE73FE20B1D1" author="Pickford M" box="[336,392,412,433]" pageId="23" pageNumber="301" pagination="1491 - 1525" refId="ref26475" refString="Pickford M. 1991. Revision of the Neogene Anthracotheriidae of Africa. The Geology of Libya 4: 1491 - 1525." type="journal article" year="1991">1991</bibRefCitation>
|
||
). Shared characters relating to the morphology of the lower molars of these genera explain they are close relatives, while only the similarity of their upper molars had been pointed out by
|
||
<bibRefCitation id="D2624B1BFFB89D114BCBFDF9FEC9B24C" author="Pickford M" box="[181,353,534,556]" pageId="23" pageNumber="301" pagination="289 - 350" refId="ref26447" refString="Pickford M. 1987. Revision des suiformes (Artiodactyla, Mammalia) de Bugti (Pakistan). Annales de Paleontologie 73: 289 - 350." type="journal article" year="1987">Pickford (1987</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="D2624B1BFFB89D114A0FFDF8FE01B24C" author="Pickford M" box="[369,425,535,556]" pageId="23" pageNumber="301" pagination="1491 - 1525" refId="ref26475" refString="Pickford M. 1991. Revision of the Neogene Anthracotheriidae of Africa. The Geology of Libya 4: 1491 - 1525." type="journal article" year="1991">1991</bibRefCitation>
|
||
). As
|
||
<taxonomicName id="71F34D69FFB89D114A98FDF9FDD8B24B" baseAuthorityName="Forster-Cooper" baseAuthorityYear="1924" box="[486,624,534,555]" class="Mammalia" family="Anthracotheriidae" genus="Telmatodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114A98FDF9FDD8B24B" box="[486,624,534,555]" italics="true" pageId="23" pageNumber="301">Telmatodon</emphasis>
|
||
</taxonomicName>
|
||
is closer to
|
||
<taxonomicName id="71F34D69FFB89D114BEFFDDAFEFAB22A" authorityName="Forster-Cooper" authorityYear="1915" box="[145,338,565,586]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114BEFFDDAFEFAB22A" box="[145,338,565,586]" italics="true" pageId="23" pageNumber="301">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
than to
|
||
<taxonomicName id="71F34D69FFB89D114AC4FDDAFD92B22A" authorityName="Pilgrim" authorityYear="1908" box="[442,570,565,586]" class="Mammalia" family="Anthracotheriidae" genus="Gonotelma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114AC4FDDAFD92B22A" box="[442,570,565,586]" italics="true" pageId="23" pageNumber="301">Gonotelma</emphasis>
|
||
</taxonomicName>
|
||
, the hypothesis of a synonymy between
|
||
<taxonomicName id="71F34D69FFB89D114ACFFDBBFD94B209" baseAuthorityName="Forster-Cooper" baseAuthorityYear="1924" box="[433,572,596,617]" class="Mammalia" family="Anthracotheriidae" genus="Telmatodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114ACFFDBBFD94B209" box="[433,572,596,617]" italics="true" pageId="23" pageNumber="301">Telmatodon</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="71F34D69FFB89D114904FDBBFD51B209" authorityName="Pilgrim" authorityYear="1908" box="[634,761,596,617]" class="Mammalia" family="Anthracotheriidae" genus="Gonotelma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114904FDBBFD51B209" box="[634,761,596,617]" italics="true" pageId="23" pageNumber="301">Gonotelma</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation id="D2624B1BFFB89D114BE7FD9DFEB5B2E8" author="Viret J" box="[153,285,626,648]" pageId="23" pageNumber="301" pagination="887 - 1084" refId="ref27315" refString="Viret J. 1961. Artiodactyla. In: Piveteau J, ed. Traite de paleontologie VI. Paris: Masson, 887 - 1084." type="book chapter" year="1961">Viret, 1961</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="D2624B1BFFB89D114A53FD9CFDB5B2E8" author="Kumar K & Kad S" box="[301,541,626,648]" pageId="23" pageNumber="301" pagination="29 - 53" refId="ref24955" refString="Kumar K, Kad S. 2003. Early Miocene vertebrates from the Murree Group, northwest Himalaya, India: affinities and age implications. Himalayan Geology 24: 29 - 53." type="journal article" year="2003">Kumar & Kad, 2003</bibRefCitation>
|
||
) is not verified by the present analysis, despite their general similarity. These findings involving the phylogenetic position of
|
||
<taxonomicName id="71F34D69FFB89D114BEFFD21FEB2B283" baseAuthorityName="Forster-Cooper" baseAuthorityYear="1924" box="[145,282,718,739]" class="Mammalia" family="Anthracotheriidae" genus="Telmatodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114BEFFD21FEB2B283" box="[145,282,718,739]" italics="true" pageId="23" pageNumber="301">Telmatodon</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="71F34D69FFB89D114A28FD21FE7CB283" authorityName="Pilgrim" authorityYear="1908" box="[342,468,718,739]" class="Mammalia" family="Anthracotheriidae" genus="Gonotelma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114A28FD21FE7CB283" box="[342,468,718,739]" italics="true" pageId="23" pageNumber="301">Gonotelma</emphasis>
|
||
</taxonomicName>
|
||
suggest that they do not constitute a clade with
|
||
<taxonomicName id="71F34D69FFB89D114AD5FD02FDC3B362" authorityName="Forster-Cooper" authorityYear="1915" box="[427,619,749,770]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114AD5FD02FDC3B362" box="[427,619,749,770]" italics="true" pageId="23" pageNumber="301">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
contrary to what
|
||
<bibRefCitation id="D2624B1BFFB89D114BADFCE3FED6B341" author="Pickford M" box="[211,382,780,802]" pageId="23" pageNumber="301" pagination="289 - 350" refId="ref26447" refString="Pickford M. 1987. Revision des suiformes (Artiodactyla, Mammalia) de Bugti (Pakistan). Annales de Paleontologie 73: 289 - 350." type="journal article" year="1987">Pickford (1987</bibRefCitation>
|
||
: fig. 4) had informally proposed. Instead,
|
||
<taxonomicName id="71F34D69FFB89D114B8BFCC5FE06B35F" authorityName="Forster-Cooper" authorityYear="1915" box="[245,430,810,831]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114B8BFCC5FE06B35F" box="[245,430,810,831]" italics="true" pageId="23" pageNumber="301">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="71F34D69FFB89D114A99FCC5FDC5B35F" baseAuthorityName="Forster-Cooper" baseAuthorityYear="1924" box="[487,621,810,831]" class="Mammalia" family="Anthracotheriidae" genus="Telmatodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114A99FCC5FDC5B35F" box="[487,621,810,831]" italics="true" pageId="23" pageNumber="301">Telmatodon</emphasis>
|
||
</taxonomicName>
|
||
can be seen as a lineage that appears to be restricted to the Indian subcontinent during the Early Miocene. These conclusions remain uncertain insofar as the scarce fossil material for
|
||
<taxonomicName id="71F34D69FFB89D114A7DFC4AFE25B3DA" baseAuthorityName="Forster-Cooper" baseAuthorityYear="1924" box="[259,397,933,954]" class="Mammalia" family="Anthracotheriidae" genus="Telmatodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114A7DFC4AFE25B3DA" box="[259,397,933,954]" italics="true" pageId="23" pageNumber="301">Telmatodon</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="71F34D69FFB89D114AB4FC4AFDE1B3DA" authorityName="Pilgrim" authorityYear="1908" box="[458,585,933,954]" class="Mammalia" family="Anthracotheriidae" genus="Gonotelma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114AB4FC4AFDE1B3DA" box="[458,585,933,954]" italics="true" pageId="23" pageNumber="301">Gonotelma</emphasis>
|
||
</taxonomicName>
|
||
(lacking lower premolars, rostral teeth, mandible and skull) has not allowed us to discriminate their diagnostic characters as precisely as for
|
||
<taxonomicName id="71F34D69FFB89D114A1CFBEDFD94B476" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[354,572,1025,1047]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFB89D114A1CFBEDFD94B476" box="[354,572,1025,1047]" italics="true" pageId="23" pageNumber="301">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
. In all cases, the phylogenetic proximity of these three associated early merycopotamines, together with their size differences (
|
||
<taxonomicName id="71F34D69FFB89D114BE7FBB2FEFAB412" authorityName="Forster-Cooper" authorityYear="1915" box="[153,338,1117,1138]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114BE7FBB2FEFAB412" box="[153,338,1117,1138]" italics="true" pageId="23" pageNumber="301">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="71F34D69FFB89D114AF2FBB2FDBAB412" baseAuthorityName="Forster-Cooper" baseAuthorityYear="1924" box="[396,530,1117,1138]" class="Mammalia" family="Anthracotheriidae" genus="Telmatodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114AF2FBB2FDBAB412" box="[396,530,1117,1138]" italics="true" pageId="23" pageNumber="301">Telmatodon</emphasis>
|
||
</taxonomicName>
|
||
being gigantic compared to
|
||
<taxonomicName id="71F34D69FFB89D114B8AFB93FEC7B4F1" authorityName="Pilgrim" authorityYear="1908" box="[244,367,1148,1169]" class="Mammalia" family="Anthracotheriidae" genus="Gonotelma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114B8AFB93FEC7B4F1" box="[244,367,1148,1169]" italics="true" pageId="23" pageNumber="301">Gonotelma</emphasis>
|
||
</taxonomicName>
|
||
), contradict the existence of a body size increase through time in this lineage, as argued by
|
||
<bibRefCitation id="D2624B1BFFB89D114BCBFB56FEC2B4AE" author="Pickford M" box="[181,362,1209,1231]" pageId="23" pageNumber="301" pagination="107 - 120" refId="ref26498" refString="Pickford M. 2009. Metric variation in Afromeryx and Libycosaurus (Anthracotheriidae: mammalia) and its utility for biochronology. ReVista Espanola de paleontolog ι a 24: 107 - 120." type="journal article" year="2009">Pickford (2009)</bibRefCitation>
|
||
for the African merycopotamines
|
||
<taxonomicName id="71F34D69FFB89D114BEFFB37FEAEB48D" authorityName="Pickford" authorityYear="1991" box="[145,262,1240,1261]" class="Mammalia" family="Anthracotheriidae" genus="Afromeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114BEFFB37FEAEB48D" box="[145,262,1240,1261]" italics="true" pageId="23" pageNumber="301">Afromeryx</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="71F34D69FFB89D114A41FB37FE7EB48D" box="[319,470,1240,1261]" class="Mammalia" family="Anthracotheriidae" genus="Libycosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114A41FB37FE7EB48D" box="[319,470,1240,1261]" italics="true" pageId="23" pageNumber="301">Libycosaurus</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<paragraph id="B64C36EAFFB89D114BD7FB19FA95B284" blockId="23.[145,762,197,1905]" lastBlockId="23.[809,1426,197,740]" pageId="23" pageNumber="301">
|
||
Of the two distal ridges of the protocone of P4/ of
|
||
<taxonomicName id="71F34D69FFB89D114BEFFAFAFEE4B54A" authorityName="Forster-Cooper" authorityYear="1915" box="[145,332,1301,1322]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114BEFFAFAFEE4B54A" box="[145,332,1301,1322]" italics="true" pageId="23" pageNumber="301">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
, the one that joins the distostyle can be interpreted at first as the postectoprotocrista and the other one, shorter and labially situated, as the postprotocrista, considering the convergent condition in
|
||
<taxonomicName id="71F34D69FFB89D114BD1FA7FFDD9B5C5" authority="(Scherler et al., 2018)" baseAuthorityName="Scherler" baseAuthorityYear="2018" box="[175,625,1424,1446]" class="Mammalia" family="Anthracotheriidae" genus="Anthracotherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114BD1FA7FFEDBB5C5" box="[175,371,1424,1445]" italics="true" pageId="23" pageNumber="301">Anthracotherium</emphasis>
|
||
(
|
||
<bibRefCitation id="D2624B1BFFB89D114AFDFA7FFDCFB5C5" author="Scherler L & Lihoreau F & Becker D" box="[387,615,1424,1446]" pageId="23" pageNumber="301" pagination="487 - 510" refId="ref27025" refString="Scherler L, Lihoreau F, Becker D. 2018. To split or not to split Anthracotherium? A phylogeny of Anthracotheriinae (Cetartiodactyla: Hippopotamoidea) and its palaeobiogeographical implications. Zoological Journal of the Linnean Society 185: 487 - 510." type="journal article" year="2018">
|
||
Scherler
|
||
<emphasis id="8487EAF8FFB89D114A95FA7FFD8BB5C5" box="[491,547,1424,1445]" italics="true" pageId="23" pageNumber="301">et al.</emphasis>
|
||
, 2018
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
. In the context of the basal position of
|
||
<taxonomicName id="71F34D69FFB89D114AB9FA40FD09B5A3" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[455,673,1454,1476]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFB89D114AB9FA40FD09B5A3" box="[455,673,1454,1476]" italics="true" pageId="23" pageNumber="301">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
in relation to
|
||
<taxonomicName id="71F34D69FFB89D114BA4FA22FE3BB583" box="[218,403,1485,1507]" pageId="23" pageNumber="304" rank="tribe" tribe="Merycopotamini">Merycopotamini</taxonomicName>
|
||
such as
|
||
<taxonomicName id="71F34D69FFB89D114A92FA22FD50B582" baseAuthorityName="Ducrocq & Lihoreau" baseAuthorityYear="2006" box="[492,760,1485,1506]" class="Mammalia" family="Anthracotheriidae" genus="Sivameryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="species" species="palaeindicus">
|
||
<emphasis id="8487EAF8FFB89D114A92FA22FD50B582" box="[492,760,1485,1506]" italics="true" pageId="23" pageNumber="301">SiƲameryx palaeindicus</emphasis>
|
||
</taxonomicName>
|
||
(also documented in the Bugti Hills), the hypothesis of homology involved by this terminology implies that a reduction of the postectoprotocrista – accompanied by a lingual displacement of the postprotocrista and its development until it reaches the distostyle – would have led to the occlusal pattern of the P4/ of this tribe (
|
||
<figureCitation id="2EC82A6FFFB89D1149B0F96AFF16B6D9" captionStart="Figure 10" captionStartId="24.[164,243,980,1002]" captionTargetBox="[323,1283,200,948]" captionTargetId="figure-422@24.[320,1286,197,951]" captionTargetPageId="24" captionText="Figure 10. Three hypotheses of homology for the cristae of P4/ within the clade B due to the inclusion of Parabrachyodus hyopotamoides, through scenarios sketched on asimplified topology.A, hypothesis 1, convergence with the Anthracotheriinae; B, hypothesis 2, reviewed homology of the distal crest of Telmatodon and the Merycopotamini; C, hypothesis 3 (this work), additional endoprotocrista of Par. hyopotamoides. Coloured characters: blue, postprotocrista; green, postectoprotocrista; red, endoprotocrista. Drawing of the P4/ of Elomeryx modified from Kostopoulos et al. (2012). Drawings are not to scale." figureDoi="http://doi.org/10.5281/zenodo.7927083" httpUri="https://zenodo.org/record/7927083/files/figure.png" pageId="23" pageNumber="301">Fig. 10A</figureCitation>
|
||
). In view of the configuration of the distal crest of the P4/ of
|
||
<taxonomicName id="71F34D69FFB89D114A7DF92DFEC6B6B7" baseAuthorityName="Hellmund" baseAuthorityYear="1991" box="[259,366,1730,1751]" class="Mammalia" family="Anthracotheriidae" genus="Elomeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114A7DF92DFEC6B6B7" box="[259,366,1730,1751]" italics="true" pageId="23" pageNumber="301">Elomeryx</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation id="D2624B1BFFB89D114A03F92DFD20B6B7" author="Kostopoulos DS & Koufos GD & Christanis K" box="[381,648,1730,1752]" pageId="23" pageNumber="301" pagination="303 - 315" refId="ref24912" refString="Kostopoulos DS, Koufos GD, Christanis K. 2012. On some anthracotheriid (Artiodactyla, Mammalia) remains from northern Greece: comments on the palaeozoogeography and phylogeny of Elomeryx. Saeiss Journal of Palaeontology 131: 303 - 315." type="journal article" year="2012">
|
||
Kostopoulos
|
||
<emphasis id="8487EAF8FFB89D114973F92CFDEDB6B7" box="[525,581,1730,1752]" italics="true" pageId="23" pageNumber="301">et al.</emphasis>
|
||
, 2012
|
||
</bibRefCitation>
|
||
: fig. 4) resembling those of
|
||
<taxonomicName id="71F34D69FFB89D114A25F90EFDBEB697" box="[347,534,1761,1783]" pageId="23" pageNumber="304" rank="tribe" tribe="Merycopotamini">Merycopotamini</taxonomicName>
|
||
, two hypotheses are equally parsimonious. Since
|
||
<taxonomicName id="71F34D69FFB89D114AA9F8EFFD1AB774" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[471,690,1791,1813]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFB89D114AA9F8EFFD1AB774" box="[471,690,1791,1813]" italics="true" pageId="23" pageNumber="301">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
is the first branching species of the clade excluding
|
||
<taxonomicName id="71F34D69FFB89D1149F0F8F1FD51B753" baseAuthorityName="Hellmund" baseAuthorityYear="1991" box="[654,761,1822,1843]" class="Mammalia" family="Anthracotheriidae" genus="Elomeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D1149F0F8F1FD51B753" box="[654,761,1822,1843]" italics="true" pageId="23" pageNumber="301">Elomeryx</emphasis>
|
||
</taxonomicName>
|
||
together with species of
|
||
<taxonomicName id="71F34D69FFB89D114AC2F8D2FDE1B732" baseAuthorityName="Forster-Cooper" baseAuthorityYear="1924" box="[444,585,1853,1874]" class="Mammalia" family="Anthracotheriidae" genus="Telmatodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114AC2F8D2FDE1B732" box="[444,585,1853,1874]" italics="true" pageId="23" pageNumber="301">Telmatodon</emphasis>
|
||
</taxonomicName>
|
||
, it may be expected that the
|
||
<taxonomicName id="71F34D69FFB89D114A33F8B4FDA3B710" authorityName="Forster-Cooper" authorityYear="1915" box="[333,523,1883,1904]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114A33F8B4FDA3B710" box="[333,523,1883,1904]" italics="true" pageId="23" pageNumber="301">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
-like postprotocrista has been lost and the postectoprotocrista is retained in all
|
||
<taxonomicName id="71F34D69FFB89D114813FF0CFB81B099" box="[877,1065,227,249]" pageId="23" pageNumber="304" rank="tribe" tribe="Merycopotamini">Merycopotamini</taxonomicName>
|
||
(
|
||
<figureCitation id="2EC82A6FFFB89D114F47FF0CFB31B099" box="[1081,1177,227,249]" captionStart="Figure 10" captionStartId="24.[164,243,980,1002]" captionTargetBox="[323,1283,200,948]" captionTargetId="figure-422@24.[320,1286,197,951]" captionTargetPageId="24" captionText="Figure 10. Three hypotheses of homology for the cristae of P4/ within the clade B due to the inclusion of Parabrachyodus hyopotamoides, through scenarios sketched on asimplified topology.A, hypothesis 1, convergence with the Anthracotheriinae; B, hypothesis 2, reviewed homology of the distal crest of Telmatodon and the Merycopotamini; C, hypothesis 3 (this work), additional endoprotocrista of Par. hyopotamoides. Coloured characters: blue, postprotocrista; green, postectoprotocrista; red, endoprotocrista. Drawing of the P4/ of Elomeryx modified from Kostopoulos et al. (2012). Drawings are not to scale." figureDoi="http://doi.org/10.5281/zenodo.7927083" httpUri="https://zenodo.org/record/7927083/files/figure.png" pageId="23" pageNumber="301">Fig. 10B</figureCitation>
|
||
). The postprotocrista of
|
||
<taxonomicName id="71F34D69FFB89D114839FEECFB87B177" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[839,1071,258,280]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFB89D114839FEECFB87B177" box="[839,1071,258,280]" italics="true" pageId="23" pageNumber="301">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
can be seen as an additional ‘endoprotocrista’ (formed from an enamel fold or a fossa) and the postectoprotocrista as the true postprotocrista that connects to the distostyle, inherited from a common ancestor with
|
||
<taxonomicName id="71F34D69FFB89D114F96FE92FAFDB1F2" baseAuthorityName="Hellmund" baseAuthorityYear="1991" box="[1256,1365,381,402]" class="Mammalia" family="Anthracotheriidae" genus="Elomeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114F96FE92FAFDB1F2" box="[1256,1365,381,402]" italics="true" pageId="23" pageNumber="301">Elomeryx</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation id="2EC82A6FFFB89D114E1BFE92FCF1B1D1" captionStart="Figure 10" captionStartId="24.[164,243,980,1002]" captionTargetBox="[323,1283,200,948]" captionTargetId="figure-422@24.[320,1286,197,951]" captionTargetPageId="24" captionText="Figure 10. Three hypotheses of homology for the cristae of P4/ within the clade B due to the inclusion of Parabrachyodus hyopotamoides, through scenarios sketched on asimplified topology.A, hypothesis 1, convergence with the Anthracotheriinae; B, hypothesis 2, reviewed homology of the distal crest of Telmatodon and the Merycopotamini; C, hypothesis 3 (this work), additional endoprotocrista of Par. hyopotamoides. Coloured characters: blue, postprotocrista; green, postectoprotocrista; red, endoprotocrista. Drawing of the P4/ of Elomeryx modified from Kostopoulos et al. (2012). Drawings are not to scale." figureDoi="http://doi.org/10.5281/zenodo.7927083" httpUri="https://zenodo.org/record/7927083/files/figure.png" pageId="23" pageNumber="301">Fig. 10C</figureCitation>
|
||
). Since the single known P4/ of
|
||
<taxonomicName id="71F34D69FFB89D114F81FE74FC31B1AF" baseAuthorityName="Forster-Cooper" baseAuthorityYear="1924" class="Mammalia" family="Anthracotheriidae" genus="Telmatodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="species" species="orientalis">
|
||
<emphasis id="8487EAF8FFB89D114F81FE74FC31B1AF" italics="true" pageId="23" pageNumber="301">Telmatodon orientalis</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation id="2EC82A6FFFB89D1148D4FE55FC53B1AF" box="[938,1019,442,464]" captionStart="Figure 10" captionStartId="24.[164,243,980,1002]" captionTargetBox="[323,1283,200,948]" captionTargetId="figure-422@24.[320,1286,197,951]" captionTargetPageId="24" captionText="Figure 10. Three hypotheses of homology for the cristae of P4/ within the clade B due to the inclusion of Parabrachyodus hyopotamoides, through scenarios sketched on asimplified topology.A, hypothesis 1, convergence with the Anthracotheriinae; B, hypothesis 2, reviewed homology of the distal crest of Telmatodon and the Merycopotamini; C, hypothesis 3 (this work), additional endoprotocrista of Par. hyopotamoides. Coloured characters: blue, postprotocrista; green, postectoprotocrista; red, endoprotocrista. Drawing of the P4/ of Elomeryx modified from Kostopoulos et al. (2012). Drawings are not to scale." figureDoi="http://doi.org/10.5281/zenodo.7927083" httpUri="https://zenodo.org/record/7927083/files/figure.png" pageId="23" pageNumber="301">Fig. 10</figureCitation>
|
||
;
|
||
<bibRefCitation id="D2624B1BFFB89D114F77FE55FB56B1AF" author="Forster-Cooper C" box="[1033,1278,442,463]" pageId="23" pageNumber="301" pagination="1 - 59" refId="ref24403" refString="Forster-Cooper C. 1924. The Anthracotheriidae of the Dera Bugti deposits in Baluchistan. Memoirs of the Geological SurVey of India. Palaeontologica Indica 4: 1 - 59." type="journal article" year="1924">Forster-Cooper, 1924</bibRefCitation>
|
||
: pl. 5, fig. 5) has a two-crested protocone, with one distal crest, the second proposal is the most likely in the context of the present topology (the P4/ of
|
||
<taxonomicName id="71F34D69FFB89D114FE0FDF9FAB3B24B" authorityName="Pilgrim" authorityYear="1908" box="[1182,1307,534,555]" class="Mammalia" family="Anthracotheriidae" genus="Gonotelma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114FE0FDF9FAB3B24B" box="[1182,1307,534,555]" italics="true" pageId="23" pageNumber="301">Gonotelma</emphasis>
|
||
</taxonomicName>
|
||
being unknown). This autapomorphic scenario (
|
||
<figureCitation id="2EC82A6FFFB89D114FA0FDDAFA94B22A" box="[1246,1340,565,587]" captionStart="Figure 10" captionStartId="24.[164,243,980,1002]" captionTargetBox="[323,1283,200,948]" captionTargetId="figure-422@24.[320,1286,197,951]" captionTargetPageId="24" captionText="Figure 10. Three hypotheses of homology for the cristae of P4/ within the clade B due to the inclusion of Parabrachyodus hyopotamoides, through scenarios sketched on asimplified topology.A, hypothesis 1, convergence with the Anthracotheriinae; B, hypothesis 2, reviewed homology of the distal crest of Telmatodon and the Merycopotamini; C, hypothesis 3 (this work), additional endoprotocrista of Par. hyopotamoides. Coloured characters: blue, postprotocrista; green, postectoprotocrista; red, endoprotocrista. Drawing of the P4/ of Elomeryx modified from Kostopoulos et al. (2012). Drawings are not to scale." figureDoi="http://doi.org/10.5281/zenodo.7927083" httpUri="https://zenodo.org/record/7927083/files/figure.png" pageId="23" pageNumber="301">Fig. 10C</figureCitation>
|
||
) is consistent with the overall trend towards the addition of styles and ridges, which have only been reported on the jugal teeth surrounding the P4/ of
|
||
<taxonomicName id="71F34D69FFB89D114FCFFD7EFA23B2C6" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[1201,1419,657,678]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFB89D114FCFFD7EFA23B2C6" box="[1201,1419,657,678]" italics="true" pageId="23" pageNumber="301">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
, namely the small distolingual style on the P3/ and the protostyle and ectoprotocrista of upper molars.
|
||
</paragraph>
|
||
<paragraph id="B64C36EAFFB89D1148FCFCCEFB6CB337" blockId="23.[898,1335,801,856]" pageId="23" pageNumber="301">
|
||
<smallCapsWord id="B0AAA036FFB89D1148FCFCCEFB8AB358" baselines="820,819" box="[898,1058,801,826]" lowerCaseFontSize="8" mainFontSize="10" normCase="title" normString="Implications" pageId="23" pageNumber="301">IMPLICATIONS</smallCapsWord>
|
||
<smallCapsWord id="B0AAA036FFB89D114F57FCCBFBFDB358" baselines="819" box="[1065,1109,804,824]" lowerCaseFontSize="8" mainFontSize="10" normCase="lower" normString="for" pageId="23" pageNumber="301">FOR</smallCapsWord>
|
||
<smallCapsWord id="B0AAA036FFB89D114F22FCCBFB21B358" baselines="819" box="[1116,1161,804,824]" lowerCaseFontSize="8" mainFontSize="10" normCase="lower" normString="the" pageId="23" pageNumber="301">THE</smallCapsWord>
|
||
<smallCapsWord id="B0AAA036FFB89D114FEEFCCBFABBB358" baselines="819" box="[1168,1299,804,824]" lowerCaseFontSize="8" mainFontSize="10" normCase="lower" normString="definition" pageId="23" pageNumber="301">DEFINITION</smallCapsWord>
|
||
<smallCapsWord id="B0AAA036FFB89D114E64FCCBFA9FB358" baselines="819" box="[1306,1335,804,824]" lowerCaseFontSize="8" mainFontSize="10" normCase="lower" normString="of" pageId="23" pageNumber="301">OF</smallCapsWord>
|
||
<taxonomicName id="71F34D69FFB89D11488BFCD0FB6CB337" box="[1013,1220,831,856]" pageId="23" pageNumber="304" rank="tribe" tribe="Merycopotamini">
|
||
<smallCapsWord id="B0AAA036FFB89D11488BFCD0FB6CB337" baselines="850,850" box="[1013,1220,831,856]" lowerCaseFontSize="8" mainFontSize="10" normCase="title" normString="Merycopotamini" pageId="23" pageNumber="301">MERYCOPOTAMINI</smallCapsWord>
|
||
</taxonomicName>
|
||
</paragraph>
|
||
<paragraph id="B64C36EAFFB89D114857FC87FC7EB775" blockId="23.[809,1426,872,1906]" pageId="23" pageNumber="301">
|
||
The paraphyly of
|
||
<taxonomicName id="71F34D69FFB89D114890FC87FCC6B3FC" authority="sensu Lihoreau et al. (2016)" authorityName="Lihoreau" authorityYear="2016" pageId="23" pageNumber="304" rank="tribe" tribe="Merycopotamini">
|
||
Merycopotamini
|
||
<emphasis id="8487EAF8FFB89D114FD4FC86FB40B31E" box="[1194,1256,873,894]" italics="true" pageId="23" pageNumber="301">sensu</emphasis>
|
||
<bibRefCitation id="D2624B1BFFB89D114F90FC87FCC6B3FC" author="Lihoreau F & Alloing-Seguier L & Antoine PO & Boisserie JR & Marivaux L & Metais G & Welcomme JL" pageId="23" pageNumber="301" pagination="947 - 957" refId="ref25524" refString="Lihoreau F, Alloing-Seguier L, Antoine PO, Boisserie JR, Marivaux L, Metais G, Welcomme JL. 2016. Enamel microstructure defines a major Paleogene hippopotamoid clade: the Merycopotamini (Cetartiodactyla, Hippopotamoidea). Historical Biology 29: 947 - 957." type="journal article" year="2016">
|
||
Lihoreau
|
||
<emphasis id="8487EAF8FFB89D114E24FC86FA39B31D" box="[1370,1425,872,894]" italics="true" pageId="23" pageNumber="301">et al.</emphasis>
|
||
(2016)
|
||
</bibRefCitation>
|
||
</taxonomicName>
|
||
and the weak support of the clade, assuming that
|
||
<taxonomicName id="71F34D69FFB89D114857FC49FC04B3DA" authorityName="Pilgrim" authorityYear="1908" box="[809,940,933,955]" class="Mammalia" family="Anthracotheriidae" genus="Gonotelma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="species" species="shahbazi">
|
||
<emphasis id="8487EAF8FFB89D114857FC49FC04B3DA" box="[809,940,933,955]" italics="true" pageId="23" pageNumber="301">G. shahbazi</emphasis>
|
||
</taxonomicName>
|
||
(the sister-species to
|
||
<taxonomicName id="71F34D69FFB89D114FE5FC4AFAA8B3DA" authorityName="Pickford" authorityYear="1991" box="[1179,1280,933,954]" class="Mammalia" family="Anthracotheriidae" genus="Afromeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="species" species="zelteni">
|
||
<emphasis id="8487EAF8FFB89D114FE5FC4AFAA8B3DA" box="[1179,1280,933,954]" italics="true" pageId="23" pageNumber="301">A. zelteni</emphasis>
|
||
</taxonomicName>
|
||
) is part of it, highlights the need for a more inclusive diagnosis of the tribe. Two-thirds of the non-ambiguous synapomorphies that made
|
||
<taxonomicName id="71F34D69FFB89D1148D4FBEEFBCFB477" box="[938,1127,1025,1047]" pageId="23" pageNumber="304" rank="tribe" tribe="Merycopotamini">Merycopotamini</taxonomicName>
|
||
a clade at the time of its definition were then interpreted as convergences with
|
||
<taxonomicName id="71F34D69FFB89D114857FBD1FC3DB433" baseAuthorityName="Hellmund" baseAuthorityYear="1991" box="[809,917,1086,1107]" class="Mammalia" family="Anthracotheriidae" genus="Elomeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114857FBD1FC3DB433" box="[809,917,1086,1107]" italics="true" pageId="23" pageNumber="301">Elomeryx</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="71F34D69FFB89D1148B1FBD1FAF1B434" authority="(Lihoreau et al., 2016)" baseAuthorityName="Lihoreau" baseAuthorityYear="2016" box="[975,1369,1086,1108]" class="Mammalia" family="Anthracotheriidae" genus="Bothriodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D1148B1FBD1FBF8B433" box="[975,1104,1086,1107]" italics="true" pageId="23" pageNumber="301">Bothriodon</emphasis>
|
||
(
|
||
<bibRefCitation id="D2624B1BFFB89D114F1FFBD1FAE7B434" author="Lihoreau F & Alloing-Seguier L & Antoine PO & Boisserie JR & Marivaux L & Metais G & Welcomme JL" box="[1121,1359,1086,1108]" pageId="23" pageNumber="301" pagination="947 - 957" refId="ref25524" refString="Lihoreau F, Alloing-Seguier L, Antoine PO, Boisserie JR, Marivaux L, Metais G, Welcomme JL. 2016. Enamel microstructure defines a major Paleogene hippopotamoid clade: the Merycopotamini (Cetartiodactyla, Hippopotamoidea). Historical Biology 29: 947 - 957." type="journal article" year="2016">
|
||
Lihoreau
|
||
<emphasis id="8487EAF8FFB89D114FAFFBD0FAA2B433" box="[1233,1290,1086,1108]" italics="true" pageId="23" pageNumber="301">et al.</emphasis>
|
||
, 2016
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
. The phylogenetic position of
|
||
<taxonomicName id="71F34D69FFB89D114F3FFBB2FB53B412" authorityName="Forster-Cooper" authorityYear="1915" box="[1089,1275,1117,1138]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114F3FFBB2FB53B412" box="[1089,1275,1117,1138]" italics="true" pageId="23" pageNumber="301">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
with respect to
|
||
<taxonomicName id="71F34D69FFB89D114839FB93FC1BB4F1" baseAuthorityName="Hellmund" baseAuthorityYear="1991" box="[839,947,1148,1169]" class="Mammalia" family="Anthracotheriidae" genus="Elomeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114839FB93FC1BB4F1" box="[839,947,1148,1169]" italics="true" pageId="23" pageNumber="301">Elomeryx</emphasis>
|
||
</taxonomicName>
|
||
on the one hand and
|
||
<taxonomicName id="71F34D69FFB89D114FCCFB93FAC5B4F2" box="[1202,1389,1148,1170]" pageId="23" pageNumber="304" rank="tribe" tribe="Merycopotamini">Merycopotamini</taxonomicName>
|
||
on the other hand establishes the suspected link between
|
||
<taxonomicName id="71F34D69FFB89D114857FB56FC3DB4AE" baseAuthorityName="Hellmund" baseAuthorityYear="1991" box="[809,917,1209,1230]" class="Mammalia" family="Anthracotheriidae" genus="Elomeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114857FB56FC3DB4AE" box="[809,917,1209,1230]" italics="true" pageId="23" pageNumber="301">Elomeryx</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="71F34D69FFB89D1148AFFB56FB25B4AF" box="[977,1165,1209,1231]" pageId="23" pageNumber="304" rank="tribe" tribe="Merycopotamini">Merycopotamini</taxonomicName>
|
||
(
|
||
<bibRefCitation id="D2624B1BFFB89D114FE0FB56FCF6B48D" author="Lihoreau F & Ducrocq S" pageId="23" pageNumber="301" pagination="89 - 105" refId="ref25128" refString="Lihoreau F, Ducrocq S. 2007. Family Anthracotheriidae. In: Protero DR, Foss SE, eds. The eVolution of artiodactyls. Baltimore: John Hopkins University Press, 89 - 105." type="book chapter" year="2007">Lihoreau & Ducrocq, 2007</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="D2624B1BFFB89D114819FB37FB81B48D" author="Bohme M & Aiglstorfer M & Antoine PO & Appel E & Havlik P & Metais G & Phuc LT & Schneider S & Setzer F & Tappert R & Tran DN & Uhl D & Prieto J" box="[871,1065,1240,1262]" pageId="23" pageNumber="301" pagination="121 - 167" refId="ref23806" refString="Bohme M, Aiglstorfer M, Antoine PO, Appel E, Havlik P, Metais G, Phuc LT, Schneider S, Setzer F, Tappert R, Tran DN, Uhl D, Prieto J. 2013. Na Duong (northern Vietnam) - an exceptional window into Eocene ecosystems from Southeast Asia. Zitteliana Reihe A: Mitteilungen der Bayerischen Staatssammlung fur Palaontologie und Geologie 53: 121 - 167." type="journal article" year="2013">
|
||
Böhme
|
||
<emphasis id="8487EAF8FFB89D1148C6FB37FC43B48D" box="[952,1003,1240,1261]" italics="true" pageId="23" pageNumber="301">et al.</emphasis>
|
||
, 2013
|
||
</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="D2624B1BFFB89D114F4CFB37FB5BB48D" author="Rincon AF & Bloch JI & MacFadden BJ & Jaramillo CA" box="[1074,1267,1240,1262]" pageId="23" pageNumber="301" pagination="421 - 433" refId="ref26853" refString="Rincon AF, Bloch JI, MacFadden BJ, Jaramillo CA. 2013. First Central American record of Anthracotheriidae (Mammalia, Bothriodontinae) from the early Miocene of Panama. Journal of Vertebrate Paleontology 33: 421 - 433." type="journal article" year="2013">
|
||
Rincon
|
||
<emphasis id="8487EAF8FFB89D114FFCFB37FB1DB48D" box="[1154,1205,1240,1261]" italics="true" pageId="23" pageNumber="301">et al.</emphasis>
|
||
, 2013
|
||
</bibRefCitation>
|
||
). Hence, seven out of nine characters defining
|
||
<taxonomicName id="71F34D69FFB89D114FEEFB19FBB9B54B" authority="sensu Lihoreau et al. (2016)" authorityName="Lihoreau" authorityYear="2016" pageId="23" pageNumber="304" rank="tribe" tribe="Merycopotamini">
|
||
Merycopotamini
|
||
<emphasis id="8487EAF8FFB89D114E2CFB18FA39B56C" box="[1362,1425,1271,1292]" italics="true" pageId="23" pageNumber="301">sensu</emphasis>
|
||
<bibRefCitation id="D2624B1BFFB89D114857FAFAFBB9B54B" author="Lihoreau F & Alloing-Seguier L & Antoine PO & Boisserie JR & Marivaux L & Metais G & Welcomme JL" box="[809,1041,1301,1323]" pageId="23" pageNumber="301" pagination="947 - 957" refId="ref25524" refString="Lihoreau F, Alloing-Seguier L, Antoine PO, Boisserie JR, Marivaux L, Metais G, Welcomme JL. 2016. Enamel microstructure defines a major Paleogene hippopotamoid clade: the Merycopotamini (Cetartiodactyla, Hippopotamoidea). Historical Biology 29: 947 - 957." type="journal article" year="2016">
|
||
Lihoreau
|
||
<emphasis id="8487EAF8FFB89D1148EDFAF9FC60B54A" box="[915,968,1301,1323]" italics="true" pageId="23" pageNumber="301">et al.</emphasis>
|
||
(2016)
|
||
</bibRefCitation>
|
||
</taxonomicName>
|
||
are retrieved as unambiguous synapomorphies in the branching sequence leading to the robust node comprising
|
||
<taxonomicName id="71F34D69FFB89D114F51FABCFAAAB507" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[1071,1282,1362,1384]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFB89D114F51FABCFAAAB507" box="[1071,1282,1362,1384]" italics="true" pageId="23" pageNumber="301">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation id="2EC82A6FFFB89D114E71FABDFAF0B508" box="[1295,1368,1362,1384]" captionStart="Figure 11" captionStartId="25.[147,228,987,1009]" captionTargetBox="[146,1425,200,954]" captionTargetId="figure-356@25.[143,1430,197,957]" captionTargetPageId="25" captionText="Figure 11. Part of the consensus tree mapped on the geologic time scale (Gradstein et al., 2012) and the geographic distribution of selected bothriodontines. The temporal extensions of taxa are coloured according to their geographical distribution, those in non-full lines are uncertain. Green, Indian subcontinent; blue, Europe; purple, North America; orange, Africa. The non-ambiguous synapomorphies are placed at the nodes discussed in this work, the character states are summarised in the Supporting Information (Appendix S2); the non-ambiguous synapomorphies defining Merycopotamini in the analysis of Lihoreau et al. (2016) are in red; the non-ambiguous and non-homoplasic synapomorphies are in bold. Drawings of the P/4 are (from top to bottom) Parabrachyodus hyopotamoides (this study), Afromeryx zelteni, SiƲameryx africanus, Merycopotamus medioximus and Libycosaurus anisae (Lihoreau et al., 2019). The characters highlighted in red on the P/4 are those acquired in the clade involved. Drawings are not to scale. Spatial and temporal distributions are from Lihoreau et al. (2007, 2016, 2019), Bhandari et al. (2010), Holroyd et al. (2010), Kostopoulos et al. (2012), Antoine et al. (2013), Böhme et al. (2013) and Kapur et al. (2019)." figureDoi="http://doi.org/10.5281/zenodo.7927085" httpUri="https://zenodo.org/record/7927085/files/figure.png" pageId="23" pageNumber="301">Fig. 11</figureCitation>
|
||
). For instance, the pinched loop-like hypoconulid on M/3, the parastyle issued from the preparacrista and the absence of ectocristyle on upper molars are actually inherited from a common ancestor with
|
||
<taxonomicName id="71F34D69FFB89D114F04FA22FB4BB582" baseAuthorityName="Hellmund" baseAuthorityYear="1991" box="[1146,1251,1485,1506]" class="Mammalia" family="Anthracotheriidae" genus="Elomeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114F04FA22FB4BB582" box="[1146,1251,1485,1506]" italics="true" pageId="23" pageNumber="301">Elomeryx</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="71F34D69FFB89D114E64FA22FA23B582" authorityName="Nikolov & Heissig" authorityYear="1985" box="[1306,1419,1485,1506]" class="Mammalia" family="Anthracotheriidae" genus="Bakalovia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114E64FA22FA23B582" box="[1306,1419,1485,1506]" italics="true" pageId="23" pageNumber="301">BakaloƲia</emphasis>
|
||
</taxonomicName>
|
||
. Thus, considering the first diagnosis of Merycopotami (
|
||
<bibRefCitation id="D2624B1BFFB89D11484FF9E5FB8DB640" author="Lihoreau F & Alloing-Seguier L & Antoine PO & Boisserie JR & Marivaux L & Metais G & Welcomme JL" box="[817,1061,1546,1568]" pageId="23" pageNumber="301" pagination="947 - 957" refId="ref25524" refString="Lihoreau F, Alloing-Seguier L, Antoine PO, Boisserie JR, Marivaux L, Metais G, Welcomme JL. 2016. Enamel microstructure defines a major Paleogene hippopotamoid clade: the Merycopotamini (Cetartiodactyla, Hippopotamoidea). Historical Biology 29: 947 - 957." type="journal article" year="2016">
|
||
Lihoreau
|
||
<emphasis id="8487EAF8FFB89D1148DAF9E4FC77B67F" box="[932,991,1546,1568]" italics="true" pageId="23" pageNumber="301">et al.</emphasis>
|
||
, 2016
|
||
</bibRefCitation>
|
||
),
|
||
<taxonomicName id="71F34D69FFB89D114F42F9E4FAB5B67F" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[1084,1309,1546,1568]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFB89D114F42F9E4FAB5B67F" box="[1084,1309,1546,1568]" italics="true" pageId="23" pageNumber="301">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
differs in only three out of 13 traits relative to the morphology of the crestids of the protoconid on P/3 and P/4, namely the orientation of the P/3 postprotocristid, the position of the P/4 postprotocristid and the direction of the P/4 preprotocristid. Nevertheless, the occlusal pattern of lower P/4 has proved to be of great interest for distinguishing
|
||
<taxonomicName id="71F34D69FFB89D1148EFF90EFBE0B697" box="[913,1096,1761,1783]" pageId="23" pageNumber="304" rank="tribe" tribe="Merycopotamini">Merycopotamini</taxonomicName>
|
||
from each other (
|
||
<bibRefCitation id="D2624B1BFFB89D114E6EF90EFC2EB775" author="Lihoreau F & Essid EM & Khayati Ammar H & Marivaux L & Marzougui W & Tabuce R & Temani R & Vianey-Liaud M & Merzeraud G" pageId="23" pageNumber="301" pagination="146 - 158" refId="ref25572" refString="Lihoreau F, Essid EM, Khayati Ammar H, Marivaux L, Marzougui W, Tabuce R, Temani R, Vianey-Liaud M, Merzeraud G. 2019. The Libycosaurus (Hippopotamoidea, Artiodactyla) intercontinental dispersal event at the early late Miocene revealed by new fossil remains from Kasserine area, Tunisia. Historical Biology 33: 146 - 158." type="journal article" year="2019">
|
||
Lihoreau
|
||
<emphasis id="8487EAF8FFB89D114E03F90DFCEDB775" italics="true" pageId="23" pageNumber="301">et al.</emphasis>
|
||
, 2019
|
||
</bibRefCitation>
|
||
: fig. 4).
|
||
</paragraph>
|
||
<paragraph id="B64C36EAFFB89D1E483FF8F1FB9CB6B4" blockId="23.[809,1426,872,1906]" lastBlockId="24.[827,1444,1175,1749]" lastPageId="24" lastPageNumber="302" pageId="23" pageNumber="301">
|
||
Considering the robust relationship between
|
||
<taxonomicName id="71F34D69FFB89D114857F8D2FC3EB732" baseAuthorityName="Hellmund" baseAuthorityYear="1991" box="[809,918,1853,1874]" class="Mammalia" family="Anthracotheriidae" genus="Elomeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB89D114857F8D2FC3EB732" box="[809,918,1853,1874]" italics="true" pageId="23" pageNumber="301">Elomeryx</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="71F34D69FFB89D1148AFF8D2FB26B733" box="[977,1166,1853,1875]" pageId="23" pageNumber="304" rank="tribe" tribe="Merycopotamini">Merycopotamini</taxonomicName>
|
||
through the inclusion of
|
||
<taxonomicName id="71F34D69FFB89D114838F8B3FB8DB711" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[838,1061,1884,1905]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="23" pageNumber="301" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFB89D114838F8B3FB8DB711" box="[838,1061,1884,1905]" italics="true" pageId="23" pageNumber="301">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
, and the primitive condition of its P/4 compared to
|
||
<taxonomicName id="71F34D69FFB79D1E4AEEFB78FE9EB4AB" authority="sensu Lihoreau et al. (2016)" authorityName="Lihoreau" authorityYear="2016" pageId="24" pageNumber="304" rank="tribe" tribe="Merycopotamini">
|
||
Merycopotamini
|
||
<emphasis id="8487EAF8FFB79D1E4929FB77FD30B4CD" box="[599,664,1176,1197]" italics="true" pageId="24" pageNumber="302">sensu</emphasis>
|
||
<bibRefCitation id="D2624B1BFFB79D1E49DEFB78FE9EB4AB" author="Lihoreau F & Alloing-Seguier L & Antoine PO & Boisserie JR & Marivaux L & Metais G & Welcomme JL" pageId="24" pageNumber="302" pagination="947 - 957" refId="ref25524" refString="Lihoreau F, Alloing-Seguier L, Antoine PO, Boisserie JR, Marivaux L, Metais G, Welcomme JL. 2016. Enamel microstructure defines a major Paleogene hippopotamoid clade: the Merycopotamini (Cetartiodactyla, Hippopotamoidea). Historical Biology 29: 947 - 957." type="journal article" year="2016">
|
||
Lihoreau
|
||
<emphasis id="8487EAF8FFB79D1E4BDDFB59FF48B4AA" box="[163,224,1205,1227]" italics="true" pageId="24" pageNumber="302">et al.</emphasis>
|
||
(2016)
|
||
</bibRefCitation>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation id="2EC82A6FFFB79D1E4A36FB5AFE33B4AB" box="[328,411,1205,1227]" captionStart="Figure 11" captionStartId="25.[147,228,987,1009]" captionTargetBox="[146,1425,200,954]" captionTargetId="figure-356@25.[143,1430,197,957]" captionTargetPageId="25" captionText="Figure 11. Part of the consensus tree mapped on the geologic time scale (Gradstein et al., 2012) and the geographic distribution of selected bothriodontines. The temporal extensions of taxa are coloured according to their geographical distribution, those in non-full lines are uncertain. Green, Indian subcontinent; blue, Europe; purple, North America; orange, Africa. The non-ambiguous synapomorphies are placed at the nodes discussed in this work, the character states are summarised in the Supporting Information (Appendix S2); the non-ambiguous synapomorphies defining Merycopotamini in the analysis of Lihoreau et al. (2016) are in red; the non-ambiguous and non-homoplasic synapomorphies are in bold. Drawings of the P/4 are (from top to bottom) Parabrachyodus hyopotamoides (this study), Afromeryx zelteni, SiƲameryx africanus, Merycopotamus medioximus and Libycosaurus anisae (Lihoreau et al., 2019). The characters highlighted in red on the P/4 are those acquired in the clade involved. Drawings are not to scale. Spatial and temporal distributions are from Lihoreau et al. (2007, 2016, 2019), Bhandari et al. (2010), Holroyd et al. (2010), Kostopoulos et al. (2012), Antoine et al. (2013), Böhme et al. (2013) and Kapur et al. (2019)." figureDoi="http://doi.org/10.5281/zenodo.7927085" httpUri="https://zenodo.org/record/7927085/files/figure.png" pageId="24" pageNumber="302">Fig. 11</figureCitation>
|
||
), we suggest a redefinition of
|
||
<taxonomicName id="71F34D69FFB79D1E4BDDFB3BFECBB48A" box="[163,355,1236,1258]" pageId="24" pageNumber="304" rank="tribe" tribe="Merycopotamini">Merycopotamini</taxonomicName>
|
||
encompassing the basal position of
|
||
<taxonomicName id="71F34D69FFB79D1E4BDDFB1CFECFB568" authorityName="Forster-Cooper" authorityYear="1915" box="[163,359,1267,1288]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB79D1E4BDDFB1CFECFB568" box="[163,359,1267,1288]" italics="true" pageId="24" pageNumber="302">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
and its relatives
|
||
<taxonomicName id="71F34D69FFB79D1E4938FB1CFD7CB568" baseAuthorityName="Forster-Cooper" baseAuthorityYear="1924" box="[582,724,1267,1288]" class="Mammalia" family="Anthracotheriidae" genus="Telmatodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB79D1E4938FB1CFD7CB568" box="[582,724,1267,1288]" italics="true" pageId="24" pageNumber="302">Telmatodon</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="71F34D69FFB79D1E4BDDFAFEFE88B546" authorityName="Pilgrim" authorityYear="1908" box="[163,288,1297,1318]" class="Mammalia" family="Anthracotheriidae" genus="Gonotelma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB79D1E4BDDFAFEFE88B546" box="[163,288,1297,1318]" italics="true" pageId="24" pageNumber="302">Gonotelma</emphasis>
|
||
</taxonomicName>
|
||
. Interestingly, the new traits of the larger tribe concern mainly the eruption of tubercles appearing subsequently during the development of the dP3/, recently considered as bearing strong diagnostic characters (
|
||
<bibRefCitation id="D2624B1BFFB79D1E4A4FFA63FD23B5C1" author="Gomes Rodrigues H & Lihoreau F & Orliac M & Boisserie JR" box="[305,651,1420,1442]" pageId="24" pageNumber="302" pagination="1 - 19" refId="ref24631" refString="Gomes Rodrigues H, Lihoreau F, Orliac M, Boisserie JR. 2020. Characters from the deciduous dentition and its interest for phylogenetic reconstruction in Hippopotamoidea (Cetartiodactyla: mammalia). Zoological Journal of the Linnean Society 20: 1 - 19." type="journal article" year="2020">
|
||
Gomes Rodrigues
|
||
<emphasis id="8487EAF8FFB79D1E4977FA62FDECB5C1" box="[521,580,1420,1442]" italics="true" pageId="24" pageNumber="302">et al.</emphasis>
|
||
, 2020
|
||
</bibRefCitation>
|
||
), the preferential development of the I/1 among the lower incisors, the multiplication of tubercles and crests on the P3/ and that of the number of mandibular foramen; hence different characters from the previous definition. Nonetheless, the critical role of the enamel microstructure in distinguishing this clade (
|
||
<bibRefCitation id="D2624B1BFFB79D1E49D0F9ABFE2CB618" author="Alloing-Seguier L & Lihoreau F & Boisserie JR & Charruault AL & Orliac M & Tabuce R" pageId="24" pageNumber="302" pagination="668 - 695" refId="ref23211" refString="Alloing-Seguier L, Lihoreau F, Boisserie JR, Charruault AL, Orliac M, Tabuce R. 2014. Enamel microstructure evolution in anthracotheres (Mammalia, Cetartiodactyla) and new insights on hippopotamoid phylogeny. Zoological Journal of the Linnean Society 171: 668 - 695." type="journal article" year="2014">
|
||
Alloing-Séguier
|
||
<emphasis id="8487EAF8FFB79D1E4A7AF98CFE96B618" box="[260,318,1635,1656]" italics="true" pageId="24" pageNumber="302">et al.</emphasis>
|
||
, 2014
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="D2624B1BFFB79D1E4AECF98CFD2BB619" author="Lihoreau F & Alloing-Seguier L & Antoine PO & Boisserie JR & Marivaux L & Metais G & Welcomme JL" box="[402,643,1635,1657]" pageId="24" pageNumber="302" pagination="947 - 957" refId="ref25524" refString="Lihoreau F, Alloing-Seguier L, Antoine PO, Boisserie JR, Marivaux L, Metais G, Welcomme JL. 2016. Enamel microstructure defines a major Paleogene hippopotamoid clade: the Merycopotamini (Cetartiodactyla, Hippopotamoidea). Historical Biology 29: 947 - 957." type="journal article" year="2016">
|
||
Lihoreau
|
||
<emphasis id="8487EAF8FFB79D1E497DF98CFD95B618" box="[515,573,1635,1656]" italics="true" pageId="24" pageNumber="302">et al.</emphasis>
|
||
, 2016
|
||
</bibRefCitation>
|
||
) is here reinforced, the substantial development of radial outer enamel being completed by the Schmelzmuster composed of two layers and weakly developed HSB. This diagnosis also clearly distinguishes merycopotamines from
|
||
<taxonomicName id="71F34D69FFB79D1E4BA0F913FEE1B771" baseAuthorityName="Hellmund" baseAuthorityYear="1991" box="[222,329,1788,1809]" class="Mammalia" family="Anthracotheriidae" genus="Elomeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB79D1E4BA0F913FEE1B771" box="[222,329,1788,1809]" italics="true" pageId="24" pageNumber="302">Elomeryx</emphasis>
|
||
</taxonomicName>
|
||
through the lack of connection between the protocone and the metaconule, closing the transverse valley on upper molars, except for
|
||
<taxonomicName id="71F34D69FFB79D1E490EF8D5FCA3B72E" authorityName="Geais" authorityYear="1934" box="[624,779,1849,1871]" class="Mammalia" family="Anthracotheriidae" genus="Elomeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="species" species="borbonicus">
|
||
<emphasis id="8487EAF8FFB79D1E490EF8D5FCA3B72E" box="[624,779,1849,1871]" italics="true" pageId="24" pageNumber="302">E. borbonicus</emphasis>
|
||
</taxonomicName>
|
||
(e.g.
|
||
<bibRefCitation id="D2624B1BFFB79D1E4BA3F8B7FEC2B70D" author="Geais G" box="[221,362,1880,1902]" pageId="24" pageNumber="302" pagination="1 - 54" refId="ref24459" refString="Geais G. 1934. Le Brachyodus borbonicus des argiles de St-Henri (pres Marseille). TraVaux du Laboratoire de Geologie de la Faculte de Lyon 25: 1 - 54." type="journal article" year="1934">Geais, 1934</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="D2624B1BFFB79D1E4A02F8B7FD96B70D" author="Hellmund M" box="[380,574,1880,1902]" pageId="24" pageNumber="302" pagination="1 - 101" refId="ref24782" refString="Hellmund M. 1991. Revision der europaischen Species der Gattung Elomeryx Marsh, 1894 (Anthracotheriidae, Artiodactyla, Mammalia) - odontologischeUntersuchungen. Palaeontographica Abteilung A, Palaozoologie, Stratigraphie 220: 1 - 101." type="journal article" year="1991">Hellmund, 1991</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="D2624B1BFFB79D1E4931F8B7FCDDB4CC" author="Lihoreau F & Ducrocq S & Antoine PO & Vianey-Liaud M & Rafay S & Garcia G & Valentin X" box="[591,885,1175,1902]" pageId="24" pageNumber="302" pagination="242 - 253" refId="ref25366" refString="Lihoreau F, Ducrocq S, Antoine PO, Vianey-Liaud M, Rafay S, Garcia G, Valentin X. 2009. First complete skulls of Elomeryx crispus (Gervais, 1849) and of Protaceratherium alblgense (Roman, 1912) from a new Oligocene locality near Moissac (SW France). Journal of Vertebrate Paleontology 29: 242 - 253." type="journal article" year="2009">
|
||
Lihoreau
|
||
<emphasis id="8487EAF8FFB79D1E49B9F8B6FCACB70D" box="[711,772,1880,1902]" italics="true" pageId="24" pageNumber="302">et al.</emphasis>
|
||
, 2009
|
||
</bibRefCitation>
|
||
: fig. 3), and between the premetacristid and the preprotocristid on lower molars. We note a tendency for the number of protocone crests to decrease in the extended tribe, from
|
||
<taxonomicName id="71F34D69FFB79D1E4FF5FB1CFAE5B568" authorityName="Forster-Cooper" authorityYear="1915" box="[1163,1357,1267,1288]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB79D1E4FF5FB1CFAE5B568" box="[1163,1357,1267,1288]" italics="true" pageId="24" pageNumber="302">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
with a quadricrescentic protocone to
|
||
<taxonomicName id="71F34D69FFB79D1E4FE7FAFEFAB9B546" authorityName="Pickford" authorityYear="1991" box="[1177,1297,1297,1318]" class="Mammalia" family="Anthracotheriidae" genus="Afromeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB79D1E4FE7FAFEFAB9B546" box="[1177,1297,1297,1318]" italics="true" pageId="24" pageNumber="302">Afromeryx</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="71F34D69FFB79D1E4E61FAFEFA34B546" authorityName="Pilgrim" authorityYear="1908" box="[1311,1436,1297,1318]" class="Mammalia" family="Anthracotheriidae" genus="Gonotelma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB79D1E4E61FAFEFA34B546" box="[1311,1436,1297,1318]" italics="true" pageId="24" pageNumber="302">Gonotelma</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="71F34D69FFB79D1E4845FADEFC6BB526" box="[827,963,1329,1350]" class="Mammalia" family="Anthracotheriidae" genus="Hemimeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB79D1E4845FADEFC6BB526" box="[827,963,1329,1350]" italics="true" pageId="24" pageNumber="302">Hemimeryx</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="71F34D69FFB79D1E48ACFADFFBF8B525" box="[978,1104,1328,1349]" class="Mammalia" family="Anthracotheriidae" genus="Sivameryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB79D1E48ACFADFFBF8B525" box="[978,1104,1328,1349]" italics="true" pageId="24" pageNumber="302">SiƲameryx</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="71F34D69FFB79D1E4FF1FADFFAB3B525" baseAuthorityName="Forster-Cooper" baseAuthorityYear="1924" box="[1167,1307,1328,1349]" class="Mammalia" family="Anthracotheriidae" genus="Telmatodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB79D1E4FF1FADFFAB3B525" box="[1167,1307,1328,1349]" italics="true" pageId="24" pageNumber="302">Telmatodon</emphasis>
|
||
</taxonomicName>
|
||
with three crests, and
|
||
<taxonomicName id="71F34D69FFB79D1E48B9FAA0FBCFB504" box="[967,1127,1359,1380]" class="Mammalia" family="Anthracotheriidae" genus="Libycosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB79D1E48B9FAA0FBCFB504" box="[967,1127,1359,1380]" italics="true" pageId="24" pageNumber="302">Libycosaurus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="71F34D69FFB79D1E4FD9FAA0FACBB504" box="[1191,1379,1359,1380]" class="Mammalia" family="Anthracotheriidae" genus="Merycopotamus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB79D1E4FD9FAA0FACBB504" box="[1191,1379,1359,1380]" italics="true" pageId="24" pageNumber="302">Merycopotamus</emphasis>
|
||
</taxonomicName>
|
||
with only two crests. Finally, this lineage is also characterized by a tendency to complexify the occlusal morphology of the
|
||
<taxonomicName id="71F34D69FFB79D1E48AFFA44FB3BB5A0" authorityName="Forster-Cooper" authorityYear="1915" box="[977,1171,1451,1472]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB79D1E48AFFA44FB3BB5A0" box="[977,1171,1451,1472]" italics="true" pageId="24" pageNumber="302">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
-like P/4, as suggested above. A mesial curvature of the preprotocristid and a lingual orientation of the postprotocristid are acquired in
|
||
<taxonomicName id="71F34D69FFB79D1E4826F9E8FC65B67C" authorityName="Pickford" authorityYear="1991" box="[856,973,1543,1564]" class="Mammalia" family="Anthracotheriidae" genus="Afromeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB79D1E4826F9E8FC65B67C" box="[856,973,1543,1564]" italics="true" pageId="24" pageNumber="302">Afromeryx</emphasis>
|
||
</taxonomicName>
|
||
, the hypoconid and a partial fusion of the postprotocristid and endoprotocristid in
|
||
<taxonomicName id="71F34D69FFB79D1E4E63F9C9FA0BB65B" box="[1309,1443,1574,1595]" class="Mammalia" family="Anthracotheriidae" genus="Hemimeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB79D1E4E63F9C9FA0BB65B" box="[1309,1443,1574,1595]" italics="true" pageId="24" pageNumber="302">Hemimeryx</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="71F34D69FFB79D1E480EF9ABFC45B639" box="[880,1005,1604,1625]" class="Mammalia" family="Anthracotheriidae" genus="Sivameryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB79D1E480EF9ABFC45B639" box="[880,1005,1604,1625]" italics="true" pageId="24" pageNumber="302">SiƲameryx</emphasis>
|
||
</taxonomicName>
|
||
, the distal crest of the entostylid in
|
||
<taxonomicName id="71F34D69FFB79D1E4845F98CFC45B618" box="[827,1005,1635,1656]" class="Mammalia" family="Anthracotheriidae" genus="Merycopotamus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB79D1E4845F98CFC45B618" box="[827,1005,1635,1656]" italics="true" pageId="24" pageNumber="302">Merycopotamus</emphasis>
|
||
</taxonomicName>
|
||
, and a multiplication of accessory cuspids mesially to the preprotocristid are independently developed in
|
||
<taxonomicName id="71F34D69FFB79D1E48A7F94EFBF7B6D6" box="[985,1119,1697,1718]" class="Mammalia" family="Anthracotheriidae" genus="Hemimeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB79D1E48A7F94EFBF7B6D6" box="[985,1119,1697,1718]" italics="true" pageId="24" pageNumber="302">Hemimeryx</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="71F34D69FFB79D1E4FE3F94FFA92B6D5" box="[1181,1338,1696,1717]" class="Mammalia" family="Anthracotheriidae" genus="Libycosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB79D1E4FE3F94FFA92B6D5" box="[1181,1338,1696,1717]" italics="true" pageId="24" pageNumber="302">Libycosaurus</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation id="2EC82A6FFFB79D1E4E35F94FFA34B6D5" box="[1355,1436,1696,1718]" captionStart="Figure 11" captionStartId="25.[147,228,987,1009]" captionTargetBox="[146,1425,200,954]" captionTargetId="figure-356@25.[143,1430,197,957]" captionTargetPageId="25" captionText="Figure 11. Part of the consensus tree mapped on the geologic time scale (Gradstein et al., 2012) and the geographic distribution of selected bothriodontines. The temporal extensions of taxa are coloured according to their geographical distribution, those in non-full lines are uncertain. Green, Indian subcontinent; blue, Europe; purple, North America; orange, Africa. The non-ambiguous synapomorphies are placed at the nodes discussed in this work, the character states are summarised in the Supporting Information (Appendix S2); the non-ambiguous synapomorphies defining Merycopotamini in the analysis of Lihoreau et al. (2016) are in red; the non-ambiguous and non-homoplasic synapomorphies are in bold. Drawings of the P/4 are (from top to bottom) Parabrachyodus hyopotamoides (this study), Afromeryx zelteni, SiƲameryx africanus, Merycopotamus medioximus and Libycosaurus anisae (Lihoreau et al., 2019). The characters highlighted in red on the P/4 are those acquired in the clade involved. Drawings are not to scale. Spatial and temporal distributions are from Lihoreau et al. (2007, 2016, 2019), Bhandari et al. (2010), Holroyd et al. (2010), Kostopoulos et al. (2012), Antoine et al. (2013), Böhme et al. (2013) and Kapur et al. (2019)." figureDoi="http://doi.org/10.5281/zenodo.7927085" httpUri="https://zenodo.org/record/7927085/files/figure.png" pageId="24" pageNumber="302">Fig. 11</figureCitation>
|
||
;
|
||
<bibRefCitation id="D2624B1BFFB79D1E4845F950FB8DB6B4" author="Lihoreau F & Essid EM & Khayati Ammar H & Marivaux L & Marzougui W & Tabuce R & Temani R & Vianey-Liaud M & Merzeraud G" box="[827,1061,1727,1749]" pageId="24" pageNumber="302" pagination="146 - 158" refId="ref25572" refString="Lihoreau F, Essid EM, Khayati Ammar H, Marivaux L, Marzougui W, Tabuce R, Temani R, Vianey-Liaud M, Merzeraud G. 2019. The Libycosaurus (Hippopotamoidea, Artiodactyla) intercontinental dispersal event at the early late Miocene revealed by new fossil remains from Kasserine area, Tunisia. Historical Biology 33: 146 - 158." type="journal article" year="2019">
|
||
Lihoreau
|
||
<emphasis id="8487EAF8FFB79D1E48D7F950FC49B6B4" box="[937,993,1727,1748]" italics="true" pageId="24" pageNumber="302">et al.</emphasis>
|
||
, 2019
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<caption id="E28C6662FFB79D1E4BDAFC3BFAFCB400" ID-DOI="http://doi.org/10.5281/zenodo.7927083" ID-Zenodo-Dep="7927083" httpUri="https://zenodo.org/record/7927083/files/figure.png" pageId="24" pageNumber="302" startId="24.[164,243,980,1002]" targetBox="[323,1283,200,948]" targetPageId="24" targetType="figure">
|
||
<paragraph id="B64C36EAFFB79D1E4BDAFC3BFAFCB400" blockId="24.[163,1444,980,1120]" pageId="24" pageNumber="302">
|
||
<emphasis id="8487EAF8FFB79D1E4BDAFC3BFEB1B38A" bold="true" box="[164,281,980,1002]" pageId="24" pageNumber="302">Figure 10.</emphasis>
|
||
Three hypotheses of homology for the cristae of P4/ within the clade B due to the inclusion of
|
||
<taxonomicName id="71F34D69FFB79D1E4F87FC3AFE96B468" baseAuthorityName="Lydekker" baseAuthorityYear="1883" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFB79D1E4F87FC3AFE96B468" italics="true" pageId="24" pageNumber="302">Parabrachyodus hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
, through scenarios sketched on a simplified topology.A, hypothesis 1, convergence with the
|
||
<taxonomicName id="71F34D69FFB79D1E4FA3FC1DFA08B468" box="[1245,1440,1010,1032]" class="Mammalia" family="Anthracotheriidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="subFamily" subFamily="Anthracotheriinae">Anthracotheriinae</taxonomicName>
|
||
; B, hypothesis 2, reviewed homology of the distal crest of
|
||
<taxonomicName id="71F34D69FFB79D1E4988FBE0FCD9B445" baseAuthorityName="Forster-Cooper" baseAuthorityYear="1924" box="[758,881,1039,1061]" class="Mammalia" family="Anthracotheriidae" genus="Telmatodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB79D1E4988FBE0FCD9B445" box="[758,881,1039,1061]" italics="true" pageId="24" pageNumber="302">Telmatodon</emphasis>
|
||
</taxonomicName>
|
||
and the
|
||
<taxonomicName id="71F34D69FFB79D1E48B0FBE0FBD3B445" box="[974,1147,1039,1061]" pageId="24" pageNumber="304" rank="tribe" tribe="Merycopotamini">Merycopotamini</taxonomicName>
|
||
; C, hypothesis 3 (this work), additional endoprotocrista of
|
||
<taxonomicName id="71F34D69FFB79D1E4AAAFBC2FD35B423" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[468,669,1069,1091]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFB79D1E4AAAFBC2FD35B423" box="[468,669,1069,1091]" italics="true" pageId="24" pageNumber="302">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
. Coloured characters: blue, postprotocrista; green, postectoprotocrista; red, endoprotocrista. Drawing of the P4/ of
|
||
<taxonomicName id="71F34D69FFB79D1E4948FBA5FD30B400" baseAuthorityName="Hellmund" baseAuthorityYear="1991" box="[566,664,1098,1120]" class="Mammalia" family="Anthracotheriidae" genus="Elomeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB79D1E4948FBA5FD30B400" box="[566,664,1098,1120]" italics="true" pageId="24" pageNumber="302">Elomeryx</emphasis>
|
||
</taxonomicName>
|
||
modified from
|
||
<bibRefCitation id="D2624B1BFFB79D1E4848FBA5FB9FB400" author="Kostopoulos DS & Koufos GD & Christanis K" box="[822,1079,1098,1120]" pageId="24" pageNumber="302" pagination="303 - 315" refId="ref24912" refString="Kostopoulos DS, Koufos GD, Christanis K. 2012. On some anthracotheriid (Artiodactyla, Mammalia) remains from northern Greece: comments on the palaeozoogeography and phylogeny of Elomeryx. Saeiss Journal of Palaeontology 131: 303 - 315." type="journal article" year="2012">
|
||
Kostopoulos
|
||
<emphasis id="8487EAF8FFB79D1E48C2FBA4FC58B400" box="[956,1008,1098,1120]" italics="true" pageId="24" pageNumber="302">et al.</emphasis>
|
||
(2012)
|
||
</bibRefCitation>
|
||
. Drawings are not to scale.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="B64C36EAFFB79D1E48F8F8FEFAF0B749" blockId="24.[902,1368,1809,1834]" box="[902,1368,1809,1834]" pageId="24" pageNumber="302">
|
||
<smallCapsWord id="B0AAA036FFB79D1E48F8F8FEFB1BB749" baselines="1828,1828" box="[902,1203,1809,1834]" lowerCaseFontSize="8" mainFontSize="10" normCase="title" normString="Palaeobiogeographical" pageId="24" pageNumber="302">PALAEOBIOGEOGRAPHICAL</smallCapsWord>
|
||
<smallCapsWord id="B0AAA036FFB79D1E4FC4F8FAFAF0B749" baselines="1828" box="[1210,1368,1813,1833]" lowerCaseFontSize="8" mainFontSize="10" normCase="lower" normString="implications" pageId="24" pageNumber="302">IMPLICATIONS</smallCapsWord>
|
||
</paragraph>
|
||
<paragraph id="B64C36EAFFB79D1F4845F8D5FB47B688" blockId="24.[827,1443,1850,1902]" lastBlockId="25.[809,1426,1379,1891]" lastPageId="25" lastPageNumber="303" pageId="24" pageNumber="302">
|
||
The northern distribution of
|
||
<taxonomicName id="71F34D69FFB79D1E4F05F8D5FAA9B72F" box="[1147,1281,1850,1871]" class="Mammalia" family="Anthracotheriidae" genus="Brachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="24" pageNumber="302" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB79D1E4F05F8D5FAA9B72F" box="[1147,1281,1850,1871]" italics="true" pageId="24" pageNumber="302">Brachyodus</emphasis>
|
||
</taxonomicName>
|
||
relative to the Himalaya Range during the Early to Middle Miocene (
|
||
<bibRefCitation id="D2624B1BFFB69D1F4BE7FA8CFED7B518" author="Ducrocq S & Chaimanee Y & Suteethorn V & Jaeger JJ" box="[153,383,1379,1401]" pageId="25" pageNumber="303" pagination="261 - 268" refId="ref24257" refString="Ducrocq S, Chaimanee Y, Suteethorn V, Jaeger JJ. 2003. Presence de l'anthracothere Brachyodus (Artiodactyla, Mammalia) dans le Miocene moyen de Thailande. Comptes Rendus PaleVol 2: 261 - 268." type="journal article" year="2003">
|
||
Ducrocq
|
||
<emphasis id="8487EAF8FFB69D1F4A7EFA8BFE91B518" box="[256,313,1379,1401]" italics="true" pageId="25" pageNumber="303">et al.</emphasis>
|
||
, 2003
|
||
</bibRefCitation>
|
||
) indicates that these mountains must have constituted a barrier to dispersal for this genus and vice versa for
|
||
<taxonomicName id="71F34D69FFB69D1F4ACDFA4FFDC7B5D5" authorityName="Forster-Cooper" authorityYear="1915" box="[435,623,1440,1461]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="25" pageNumber="303" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB69D1F4ACDFA4FFDC7B5D5" box="[435,623,1440,1461]" italics="true" pageId="25" pageNumber="303">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
from
|
||
<collectingCountry id="CEE4767AFFB69D1F49CAFA4FFD5BB5D6" box="[692,755,1440,1462]" name="India" pageId="25" pageNumber="303">India</collectingCountry>
|
||
, inasmuch as no specimen from
|
||
<collectingCountry id="CEE4767AFFB69D1F4A87FA50FD97B5B5" box="[505,575,1471,1493]" name="China" pageId="25" pageNumber="303">China</collectingCountry>
|
||
similar to those from the Indian subcontinent is documented. An alternative hypothesis for their non-overlap is a mutual exclusion for ecological reasons.
|
||
<taxonomicName id="71F34D69FFB69D1F4A94F9F4FD51B650" authorityName="Geais" authorityYear="1934" box="[490,761,1563,1585]" class="Mammalia" family="Anthracotheriidae" genus="Elomeryx" higherTaxonomySource="GBIF" isUncertain="true" kingdom="Animalia" order="Artiodactyla" pageId="25" pageNumber="303" phylum="Chordata" rank="species" species="borbonicus">
|
||
<emphasis id="8487EAF8FFB69D1F4A94F9F4FDFEB650" box="[490,598,1563,1584]" italics="true" pageId="25" pageNumber="303">Elomeryx</emphasis>
|
||
cf.
|
||
<emphasis id="8487EAF8FFB69D1F4903F9F4FD51B650" box="[637,761,1563,1584]" italics="true" pageId="25" pageNumber="303">borbonicus</emphasis>
|
||
</taxonomicName>
|
||
is known from two jugal teeth in the Zinda Pir Dome in Z108 locality (
|
||
<bibRefCitation id="D2624B1BFFB69D1F4A2AF9B6FD2BB60E" author="Ducrocq S & Lihoreau F" box="[340,643,1624,1646]" pageId="25" pageNumber="303" pagination="885 - 891" refId="ref24181" refString="Ducrocq S, Lihoreau F. 2006. The occurrence of bothriodontines (Artiodactyla, Mammalia) in the Paleogene of Asia with special reference to Elomeryx: paleobiogeographical implications. Journal of Asian Earth Sciences 27: 885 - 891." type="journal article" year="2006">Ducrocq & Lihoreau, 2006</bibRefCitation>
|
||
), an older locality than those that have yielded the oldest material of
|
||
<taxonomicName id="71F34D69FFB69D1F4B8BF979FE1AB6CB" authorityName="Forster-Cooper" authorityYear="1915" box="[245,434,1686,1707]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="25" pageNumber="303" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB69D1F4B8BF979FE1AB6CB" box="[245,434,1686,1707]" italics="true" pageId="25" pageNumber="303">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
(Z114 and Z154;
|
||
<bibRefCitation id="D2624B1BFFB69D1F4901F979FF59B6AA" author="Lindsay EH & Flynn LJ & Cheema IU & Barry JC & Downing K & Rajpar AR & Raza SM" pageId="25" pageNumber="303" pagination="1 - 19" refId="ref25636" refString="Lindsay EH, Flynn LJ, Cheema IU, Barry JC, Downing K, Rajpar AR, Raza SM. 2005. Will Downs and the Zinda Pir Dome. Palaeontologia Electronica 8: 1 - 19." type="journal article" year="2005">
|
||
Lindsay
|
||
<emphasis id="8487EAF8FFB69D1F499BF979FF06B6A9" italics="true" pageId="25" pageNumber="303">et al.</emphasis>
|
||
, 2005
|
||
</bibRefCitation>
|
||
: fig. 6B). The occurrence of this European species in
|
||
<collectingCountry id="CEE4767AFFB69D1F4B9BF93CFEE6B689" box="[229,334,1747,1769]" name="Pakistan" pageId="25" pageNumber="303">Pakistan</collectingCountry>
|
||
shows that the palaeobiogeography of the Late Oligocene and the Early Miocene allowed interchanges with Europe, as massively illustrated by mammalian assemblages of the Early Miocene in general (e.g.
|
||
<bibRefCitation id="D2624B1BFFB69D1F4A61F8A2FE51B703" author="Antoine PO & Downing KF & Crochet JY & Duranthon F & Flynn LJ & Marivaux L & Metais G & Rajpar AR & Roohi G" box="[287,505,1869,1891]" pageId="25" pageNumber="303" pagination="139 - 194" refId="ref23458" refString="Antoine PO, Downing KF, Crochet JY, Duranthon F, Flynn LJ, Marivaux L, Metais G, Rajpar AR, Roohi G. 2010. A revision of Aceratherium blanfordi Lydekker, 1884 (Mammalia: Rhinocerotidae) from the Early Miocene of Pakistan: postcranials as a key. Zoological Journal of the Linnean Society 160: 139 - 194." type="journal article" year="2010">
|
||
Antoine
|
||
<emphasis id="8487EAF8FFB69D1F4A01F8A1FE07B702" box="[383,431,1869,1891]" italics="true" pageId="25" pageNumber="303">et al</emphasis>
|
||
., 2010
|
||
</bibRefCitation>
|
||
, 2013). In Burdigalian times (late Early Miocene), the two concomitant dispersal events involving
|
||
<taxonomicName id="71F34D69FFB69D1F4F4CFA6DFB02B5F7" box="[1074,1194,1410,1431]" class="Mammalia" family="Anthracotheriidae" genus="Sivameryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="25" pageNumber="303" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB69D1F4F4CFA6DFB02B5F7" box="[1074,1194,1410,1431]" italics="true" pageId="25" pageNumber="303">SiƲameryx</emphasis>
|
||
</taxonomicName>
|
||
on the one hand and
|
||
<taxonomicName id="71F34D69FFB69D1F4857FA4FFC0DB5D5" authorityName="Pilgrim" authorityYear="1908" box="[809,933,1440,1461]" class="Mammalia" family="Anthracotheriidae" genus="Gonotelma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="25" pageNumber="303" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB69D1F4857FA4FFC0DB5D5" box="[809,933,1440,1461]" italics="true" pageId="25" pageNumber="303">Gonotelma</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="71F34D69FFB69D1F48A1FA4FFBFDB5D5" authorityName="Pickford" authorityYear="1991" box="[991,1109,1440,1461]" class="Mammalia" family="Anthracotheriidae" genus="Afromeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="25" pageNumber="303" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB69D1F48A1FA4FFBFDB5D5" box="[991,1109,1440,1461]" italics="true" pageId="25" pageNumber="303">Afromeryx</emphasis>
|
||
</taxonomicName>
|
||
on the other (Proboscidean Datum Events,
|
||
<figureCitation id="2EC82A6FFFB69D1F4890FA50FBEBB5B4" box="[1006,1091,1471,1493]" captionStart="Figure 11" captionStartId="25.[147,228,987,1009]" captionTargetBox="[146,1425,200,954]" captionTargetId="figure-356@25.[143,1430,197,957]" captionTargetPageId="25" captionText="Figure 11. Part of the consensus tree mapped on the geologic time scale (Gradstein et al., 2012) and the geographic distribution of selected bothriodontines. The temporal extensions of taxa are coloured according to their geographical distribution, those in non-full lines are uncertain. Green, Indian subcontinent; blue, Europe; purple, North America; orange, Africa. The non-ambiguous synapomorphies are placed at the nodes discussed in this work, the character states are summarised in the Supporting Information (Appendix S2); the non-ambiguous synapomorphies defining Merycopotamini in the analysis of Lihoreau et al. (2016) are in red; the non-ambiguous and non-homoplasic synapomorphies are in bold. Drawings of the P/4 are (from top to bottom) Parabrachyodus hyopotamoides (this study), Afromeryx zelteni, SiƲameryx africanus, Merycopotamus medioximus and Libycosaurus anisae (Lihoreau et al., 2019). The characters highlighted in red on the P/4 are those acquired in the clade involved. Drawings are not to scale. Spatial and temporal distributions are from Lihoreau et al. (2007, 2016, 2019), Bhandari et al. (2010), Holroyd et al. (2010), Kostopoulos et al. (2012), Antoine et al. (2013), Böhme et al. (2013) and Kapur et al. (2019)." figureDoi="http://doi.org/10.5281/zenodo.7927085" httpUri="https://zenodo.org/record/7927085/files/figure.png" pageId="25" pageNumber="303">Fig. 11</figureCitation>
|
||
), via the probable connection of the Indus with the Tiger–Euphrate drainage basin, provide evidence that a passageway to Africa was also open from
|
||
<collectingCountry id="CEE4767AFFB69D1F4F73F9F4FBDBB651" box="[1037,1139,1563,1585]" name="Pakistan" pageId="25" pageNumber="303">Pakistan</collectingCountry>
|
||
(e.g.
|
||
<bibRefCitation id="D2624B1BFFB69D1F4FCEF9F4FA22B650" author="Barrier E & Vrielynck B & Brouillet JF & Brunet MF" box="[1200,1418,1563,1585]" pageId="25" pageNumber="303" refId="ref23671" refString="Barrier E, Vrielynck B, Brouillet JF, Brunet MF. 2018. Paleotectonic reconstruction of the Central Tethyan Realm. Tectonono-sedimentary-palinspastic maps from Late Permian to Pliocene. Paris: Commission for the Geological Map of the World (CGMW / CCGM)." type="book" year="2018">
|
||
Barrier
|
||
<emphasis id="8487EAF8FFB69D1F4E73F9F4FAEEB650" box="[1293,1350,1563,1584]" italics="true" pageId="25" pageNumber="303">et al.</emphasis>
|
||
, 2018
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="D2624B1BFFB69D1F4857F9D5FBB6B62F" author="Grossman A & Calvo R & Lopez-Antonanzas R & Knoll F & Hartman G & Rabinovich R" box="[809,1054,1593,1615]" pageId="25" pageNumber="303" pagination="1599901" refId="ref24722" refString="Grossman A, Calvo R, Lopez-Antonanzas R, Knoll F, Hartman G, Rabinovich R. 2019. First record of SiVameryx (Cetartiodactyla, Anthracotheriidae) from the lower Miocene of Israel highlights the importance of the Levantine Corridor as a dispersal route between Eurasia and Africa. Journal of Vertebrate Paleontology 39: e 1599901." type="journal article" year="2019">
|
||
Grossman
|
||
<emphasis id="8487EAF8FFB69D1F48DDF9D5FC73B62E" box="[931,987,1593,1615]" italics="true" pageId="25" pageNumber="303">et al.</emphasis>
|
||
, 2019
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="D2624B1BFFB69D1F4F54F9D6FABBB62F" author="Lihoreau F & Essid EM & Khayati Ammar H & Marivaux L & Marzougui W & Tabuce R & Temani R & Vianey-Liaud M & Merzeraud G" box="[1066,1299,1593,1615]" pageId="25" pageNumber="303" pagination="146 - 158" refId="ref25572" refString="Lihoreau F, Essid EM, Khayati Ammar H, Marivaux L, Marzougui W, Tabuce R, Temani R, Vianey-Liaud M, Merzeraud G. 2019. The Libycosaurus (Hippopotamoidea, Artiodactyla) intercontinental dispersal event at the early late Miocene revealed by new fossil remains from Kasserine area, Tunisia. Historical Biology 33: 146 - 158." type="journal article" year="2019">
|
||
Lihoreau
|
||
<emphasis id="8487EAF8FFB69D1F4FE6F9D5FB78B62E" box="[1176,1232,1593,1615]" italics="true" pageId="25" pageNumber="303">et al.</emphasis>
|
||
, 2019
|
||
</bibRefCitation>
|
||
). An origin for
|
||
<taxonomicName id="71F34D69FFB69D1F482DF9B7FBB1B60E" box="[851,1049,1624,1646]" pageId="25" pageNumber="304" rank="tribe" tribe="Merycopotamini">Merycopotamini</taxonomicName>
|
||
, rooted by
|
||
<taxonomicName id="71F34D69FFB69D1F4FDCF9B6FA23B60D" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[1186,1419,1624,1646]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="25" pageNumber="303" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFB69D1F4FDCF9B6FA23B60D" box="[1186,1419,1624,1646]" italics="true" pageId="25" pageNumber="303">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
, in this north-western province of the Indian subcontinent (i.e.
|
||
<collectingCountry id="CEE4767AFFB69D1F48D9F97AFBA7B6CB" box="[935,1039,1685,1707]" name="Pakistan" pageId="25" pageNumber="303">Pakistan</collectingCountry>
|
||
), is therefore consistent with this second phase of dispersal in the evolutionary history of this lineage of bothriodontines (
|
||
<figureCitation id="2EC82A6FFFB69D1F4FEDF93CFB77B688" box="[1171,1247,1747,1769]" captionStart="Figure 11" captionStartId="25.[147,228,987,1009]" captionTargetBox="[146,1425,200,954]" captionTargetId="figure-356@25.[143,1430,197,957]" captionTargetPageId="25" captionText="Figure 11. Part of the consensus tree mapped on the geologic time scale (Gradstein et al., 2012) and the geographic distribution of selected bothriodontines. The temporal extensions of taxa are coloured according to their geographical distribution, those in non-full lines are uncertain. Green, Indian subcontinent; blue, Europe; purple, North America; orange, Africa. The non-ambiguous synapomorphies are placed at the nodes discussed in this work, the character states are summarised in the Supporting Information (Appendix S2); the non-ambiguous synapomorphies defining Merycopotamini in the analysis of Lihoreau et al. (2016) are in red; the non-ambiguous and non-homoplasic synapomorphies are in bold. Drawings of the P/4 are (from top to bottom) Parabrachyodus hyopotamoides (this study), Afromeryx zelteni, SiƲameryx africanus, Merycopotamus medioximus and Libycosaurus anisae (Lihoreau et al., 2019). The characters highlighted in red on the P/4 are those acquired in the clade involved. Drawings are not to scale. Spatial and temporal distributions are from Lihoreau et al. (2007, 2016, 2019), Bhandari et al. (2010), Holroyd et al. (2010), Kostopoulos et al. (2012), Antoine et al. (2013), Böhme et al. (2013) and Kapur et al. (2019)." figureDoi="http://doi.org/10.5281/zenodo.7927085" httpUri="https://zenodo.org/record/7927085/files/figure.png" pageId="25" pageNumber="303">Fig. 11</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<caption id="E28C6662FFB69D1F4BEDFC34FD00B577" ID-DOI="http://doi.org/10.5281/zenodo.7927085" ID-Zenodo-Dep="7927085" httpUri="https://zenodo.org/record/7927085/files/figure.png" pageId="25" pageNumber="303" startId="25.[147,228,987,1009]" targetBox="[146,1425,200,954]" targetPageId="25" targetType="figure">
|
||
<paragraph id="B64C36EAFFB69D1F4BEDFC34FD00B577" blockId="25.[147,1428,987,1303]" pageId="25" pageNumber="303">
|
||
<emphasis id="8487EAF8FFB69D1F4BEDFC34FEA5B390" bold="true" box="[147,269,987,1009]" pageId="25" pageNumber="303">Figure 11.</emphasis>
|
||
Part of the consensus tree mapped on the geologic time scale (
|
||
<bibRefCitation id="D2624B1BFFB69D1F48BAFC34FB1AB391" author="Gradstein FM & Ogg JG & Schmitz MD" box="[964,1202,987,1009]" pageId="25" pageNumber="303" refId="ref24677" refString="Gradstein FM, Ogg JG, Schmitz MD. 2012. The geologic time scale 2012. Amsterdam: Elsevier, 1144." type="book" year="2012">
|
||
Gradstein
|
||
<emphasis id="8487EAF8FFB69D1F4F47FC33FBD8B391" box="[1081,1136,987,1009]" italics="true" pageId="25" pageNumber="303">et al.</emphasis>
|
||
, 2012
|
||
</bibRefCitation>
|
||
) and the geographic distribution of selected bothriodontines. The temporal extensions of taxa are coloured according to their geographical distribution, those in non-full lines are uncertain. Green, Indian subcontinent; blue, Europe; purple, North America; orange, Africa. The non-ambiguous synapomorphies are placed at the nodes discussed in this work, the character states are summarised in the Supporting Information (Appendix S2); the non-ambiguous synapomorphies defining
|
||
<taxonomicName id="71F34D69FFB69D1F4F98FBBEFA39B407" box="[1254,1425,1105,1127]" pageId="25" pageNumber="304" rank="tribe" tribe="Merycopotamini">Merycopotamini</taxonomicName>
|
||
in the analysis of
|
||
<bibRefCitation id="D2624B1BFFB69D1F4A2CFB81FD90B4E4" author="Lihoreau F & Alloing-Seguier L & Antoine PO & Boisserie JR & Marivaux L & Metais G & Welcomme JL" box="[338,568,1134,1156]" pageId="25" pageNumber="303" pagination="947 - 957" refId="ref25524" refString="Lihoreau F, Alloing-Seguier L, Antoine PO, Boisserie JR, Marivaux L, Metais G, Welcomme JL. 2016. Enamel microstructure defines a major Paleogene hippopotamoid clade: the Merycopotamini (Cetartiodactyla, Hippopotamoidea). Historical Biology 29: 947 - 957." type="journal article" year="2016">
|
||
Lihoreau
|
||
<emphasis id="8487EAF8FFB69D1F4AC4FB80FE47B4E4" box="[442,495,1134,1156]" italics="true" pageId="25" pageNumber="303">et al.</emphasis>
|
||
(2016)
|
||
</bibRefCitation>
|
||
are in red; the non-ambiguous and non-homoplasic synapomorphies are in bold. Drawings of the P/4 are (from top to bottom)
|
||
<taxonomicName id="71F34D69FFB69D1F4906FB64FC63B4C1" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[632,971,1163,1185]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="25" pageNumber="303" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFB69D1F4906FB64FC63B4C1" box="[632,971,1163,1185]" italics="true" pageId="25" pageNumber="303">Parabrachyodus hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
(this study),
|
||
<taxonomicName id="71F34D69FFB69D1F4F25FB64FABDB4C1" authorityName="Pickford" authorityYear="1991" box="[1115,1301,1163,1185]" class="Mammalia" family="Anthracotheriidae" genus="Afromeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="25" pageNumber="303" phylum="Chordata" rank="species" species="zelteni">
|
||
<emphasis id="8487EAF8FFB69D1F4F25FB64FABDB4C1" box="[1115,1301,1163,1185]" italics="true" pageId="25" pageNumber="303">Afromeryx zelteni</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="71F34D69FFB69D1F4E5CFB64FF5EB4DF" class="Mammalia" family="Anthracotheriidae" genus="Sivameryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="25" pageNumber="303" phylum="Chordata" rank="species" species="africanus">
|
||
<emphasis id="8487EAF8FFB69D1F4E5CFB64FF5EB4DF" italics="true" pageId="25" pageNumber="303">SiƲameryx africanus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="71F34D69FFB69D1F4A7CFB46FD83B4DF" box="[258,555,1193,1215]" class="Mammalia" family="Anthracotheriidae" genus="Merycopotamus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="25" pageNumber="303" phylum="Chordata" rank="species" species="medioximus">
|
||
<emphasis id="8487EAF8FFB69D1F4A7CFB46FD83B4DF" box="[258,555,1193,1215]" italics="true" pageId="25" pageNumber="303">Merycopotamus medioximus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="71F34D69FFB69D1F491EFB46FB8FB4DF" authority="(Lihoreau et al., 2019)" baseAuthorityName="Lihoreau" baseAuthorityYear="2019" box="[608,1063,1193,1215]" class="Mammalia" family="Anthracotheriidae" genus="Libycosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="25" pageNumber="303" phylum="Chordata" rank="species" species="anisae">
|
||
<emphasis id="8487EAF8FFB69D1F491EFB46FC9FB4DF" box="[608,823,1193,1215]" italics="true" pageId="25" pageNumber="303">Libycosaurus anisae</emphasis>
|
||
(
|
||
<bibRefCitation id="D2624B1BFFB69D1F483BFB46FBB6B4DF" author="Lihoreau F & Essid EM & Khayati Ammar H & Marivaux L & Marzougui W & Tabuce R & Temani R & Vianey-Liaud M & Merzeraud G" box="[837,1054,1193,1215]" pageId="25" pageNumber="303" pagination="146 - 158" refId="ref25572" refString="Lihoreau F, Essid EM, Khayati Ammar H, Marivaux L, Marzougui W, Tabuce R, Temani R, Vianey-Liaud M, Merzeraud G. 2019. The Libycosaurus (Hippopotamoidea, Artiodactyla) intercontinental dispersal event at the early late Miocene revealed by new fossil remains from Kasserine area, Tunisia. Historical Biology 33: 146 - 158." type="journal article" year="2019">
|
||
Lihoreau
|
||
<emphasis id="8487EAF8FFB69D1F48D5FB46FC77B4DF" box="[939,991,1193,1215]" italics="true" pageId="25" pageNumber="303">et al.</emphasis>
|
||
, 2019
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
. The characters highlighted in red on the P/4 are those acquired in the clade involved. Drawings are not to scale. Spatial and temporal distributions are from Lihoreau
|
||
<emphasis id="8487EAF8FFB69D1F4B85FB0BFE82B499" box="[251,298,1251,1273]" italics="true" pageId="25" pageNumber="303">et al</emphasis>
|
||
. (2007, 2016, 2019),
|
||
<bibRefCitation id="D2624B1BFFB69D1F4976FB0CFD5DB499" author="Bhandari A & Mohabey DM & Bajpai S & Tiwari BN & Pickford M" box="[520,757,1251,1273]" pageId="25" pageNumber="303" pagination="69 - 97" refId="ref23717" refString="Bhandari A, Mohabey DM, Bajpai S, Tiwari BN, Pickford M. 2010. Early Miocene mammals from central Kutch (Gujarat), Western India: implications for geochronology, biogeography, eustacy and intercontinental dispersals. Neues Jahrbuch fur Geologie und Palaontologie - Abhandlungen 256: 69 - 97." type="journal article" year="2010">
|
||
Bhandari
|
||
<emphasis id="8487EAF8FFB69D1F490BFB0BFD03B499" box="[629,683,1251,1273]" italics="true" pageId="25" pageNumber="303">et al.</emphasis>
|
||
(2010)
|
||
</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="D2624B1BFFB69D1F487CFB0CFC77B499" author="Holroyd PA & Lihoreau F & Gunnell GF & Miller ER" box="[770,991,1251,1273]" pageId="25" pageNumber="303" pagination="843 - 851" refId="ref24820" refString="Holroyd PA, Lihoreau F, Gunnell GF, Miller ER. 2010. Anthracotheriidae. In: Werdelin L, Sanders WJ, eds. Cenozoic mammals of Africa. Berkeley: University of California Press, 843 - 851." type="book chapter" year="2010">
|
||
Holroyd
|
||
<emphasis id="8487EAF8FFB69D1F4821FB0BFC3DB499" box="[863,917,1251,1273]" italics="true" pageId="25" pageNumber="303">et al.</emphasis>
|
||
(2010)
|
||
</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="D2624B1BFFB69D1F4892FB0CFB5DB499" author="Kostopoulos DS & Koufos GD & Christanis K" box="[1004,1269,1251,1273]" pageId="25" pageNumber="303" pagination="303 - 315" refId="ref24912" refString="Kostopoulos DS, Koufos GD, Christanis K. 2012. On some anthracotheriid (Artiodactyla, Mammalia) remains from northern Greece: comments on the palaeozoogeography and phylogeny of Elomeryx. Saeiss Journal of Palaeontology 131: 303 - 315." type="journal article" year="2012">
|
||
Kostopoulos
|
||
<emphasis id="8487EAF8FFB69D1F4F08FB0BFB03B499" box="[1142,1195,1251,1273]" italics="true" pageId="25" pageNumber="303">et al.</emphasis>
|
||
(2012)
|
||
</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="D2624B1BFFB69D1F4E7CFB0CFF7CB577" author="Antoine PO & Metais G & Orliac MJ & Crochet JY & Flynn LJ & Marivaux L & Rajpar AR & Roohi G & Welcomme JL" pageId="25" pageNumber="303" pagination="400 - 422" refId="ref23524" refString="Antoine PO, Metais G, Orliac MJ, Crochet JY, Flynn LJ, Marivaux L, Rajpar AR, Roohi G, Welcomme JL. 2013. Mammalian Neogene biostratigraphy of the Sulaiman Province, Pakistan. In: Xiaoming W, Flynn LJ, Fortelius M, eds. Fossil mammals of Asia. New York: Columbia University Press, 400 - 422." type="book chapter" year="2013">
|
||
Antoine
|
||
<emphasis id="8487EAF8FFB69D1F4E23FB0BFA3BB499" box="[1373,1427,1251,1273]" italics="true" pageId="25" pageNumber="303">et al.</emphasis>
|
||
(2013)
|
||
</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="D2624B1BFFB69D1F4B9EFAEEFE01B577" author="Bohme M & Aiglstorfer M & Antoine PO & Appel E & Havlik P & Metais G & Phuc LT & Schneider S & Setzer F & Tappert R & Tran DN & Uhl D & Prieto J" box="[224,425,1281,1303]" pageId="25" pageNumber="303" pagination="121 - 167" refId="ref23806" refString="Bohme M, Aiglstorfer M, Antoine PO, Appel E, Havlik P, Metais G, Phuc LT, Schneider S, Setzer F, Tappert R, Tran DN, Uhl D, Prieto J. 2013. Na Duong (northern Vietnam) - an exceptional window into Eocene ecosystems from Southeast Asia. Zitteliana Reihe A: Mitteilungen der Bayerischen Staatssammlung fur Palaontologie und Geologie 53: 121 - 167." type="journal article" year="2013">
|
||
Böhme
|
||
<emphasis id="8487EAF8FFB69D1F4A51FAEEFEF5B577" box="[303,349,1281,1303]" italics="true" pageId="25" pageNumber="303">et al</emphasis>
|
||
. (2013)
|
||
</bibRefCitation>
|
||
and
|
||
<bibRefCitation id="D2624B1BFFB69D1F4AA3FAEEFD0AB577" author="Kapur VV & Pickford M & Chauhan G & Thakkar MG" box="[477,674,1281,1303]" pageId="25" pageNumber="303" pagination="595 - 615" refId="ref24862" refString="Kapur VV, Pickford M, Chauhan G, Thakkar MG. 2019. A Middle Miocene (~ 14 Ma) vertebrate assemblage from Palasava, Rapar Taluka, Kutch (Kachchh) District, Gujarat State, western India. Historical Biology 33: 595 - 615." type="journal article" year="2019">
|
||
Kapur
|
||
<emphasis id="8487EAF8FFB69D1F4959FAEEFDF3B577" box="[551,603,1281,1303]" italics="true" pageId="25" pageNumber="303">et al.</emphasis>
|
||
(2019)
|
||
</bibRefCitation>
|
||
.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="B64C36EAFFB69D1C483FF91DFE2BB18F" blockId="25.[809,1426,1379,1891]" lastBlockId="26.[163,782,197,1905]" lastPageId="26" lastPageNumber="304" pageId="25" pageNumber="303">
|
||
Unlike the small merycopotamines,
|
||
<taxonomicName id="71F34D69FFB69D1F4E22F91DFC70B745" baseAuthorityName="Lydekker" baseAuthorityYear="1883" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="25" pageNumber="303" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFB69D1F4E22F91DFC70B745" italics="true" pageId="25" pageNumber="303">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
appears to be endemic to the Indian subcontinent. The absence of dispersal towards Africa is probably linked to its extinction just before the contact between the two continents (
|
||
<figureCitation id="2EC82A6FFFB59D1C49DBFF2AFD53B0BA" box="[677,763,197,219]" captionStart="Figure 11" captionStartId="25.[147,228,987,1009]" captionTargetBox="[146,1425,200,954]" captionTargetId="figure-356@25.[143,1430,197,957]" captionTargetPageId="25" captionText="Figure 11. Part of the consensus tree mapped on the geologic time scale (Gradstein et al., 2012) and the geographic distribution of selected bothriodontines. The temporal extensions of taxa are coloured according to their geographical distribution, those in non-full lines are uncertain. Green, Indian subcontinent; blue, Europe; purple, North America; orange, Africa. The non-ambiguous synapomorphies are placed at the nodes discussed in this work, the character states are summarised in the Supporting Information (Appendix S2); the non-ambiguous synapomorphies defining Merycopotamini in the analysis of Lihoreau et al. (2016) are in red; the non-ambiguous and non-homoplasic synapomorphies are in bold. Drawings of the P/4 are (from top to bottom) Parabrachyodus hyopotamoides (this study), Afromeryx zelteni, SiƲameryx africanus, Merycopotamus medioximus and Libycosaurus anisae (Lihoreau et al., 2019). The characters highlighted in red on the P/4 are those acquired in the clade involved. Drawings are not to scale. Spatial and temporal distributions are from Lihoreau et al. (2007, 2016, 2019), Bhandari et al. (2010), Holroyd et al. (2010), Kostopoulos et al. (2012), Antoine et al. (2013), Böhme et al. (2013) and Kapur et al. (2019)." figureDoi="http://doi.org/10.5281/zenodo.7927085" httpUri="https://zenodo.org/record/7927085/files/figure.png" pageId="26" pageNumber="304">Fig. 11</figureCitation>
|
||
).
|
||
<taxonomicName id="71F34D69FFB59D1C4BDDFF0BFDE8B099" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[163,576,228,249]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="26" pageNumber="304" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFB59D1C4BDDFF0BFDE8B099" box="[163,576,228,249]" italics="true" pageId="26" pageNumber="304">Parabrachyodus hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
displaying the thickest enamel among bothriodontines, allowing more resistance to wear that could explain the strong wear gradients of molars rows (
|
||
<bibRefCitation id="D2624B1BFFB59D1C4950FEAFFEE0B114" author="Alloing-Seguier L & Lihoreau F & Boisserie JR & Charruault AL & Orliac M & Tabuce R" pageId="26" pageNumber="304" pagination="668 - 695" refId="ref23211" refString="Alloing-Seguier L, Lihoreau F, Boisserie JR, Charruault AL, Orliac M, Tabuce R. 2014. Enamel microstructure evolution in anthracotheres (Mammalia, Cetartiodactyla) and new insights on hippopotamoid phylogeny. Zoological Journal of the Linnean Society 171: 668 - 695." type="journal article" year="2014">
|
||
Alloing-Séguier
|
||
<emphasis id="8487EAF8FFB59D1C4988FEAFFF6AB113" italics="true" pageId="26" pageNumber="304">et al.</emphasis>
|
||
, 2014: 691
|
||
</bibRefCitation>
|
||
), the flattest occlusal surface among bothriodontines and molars with a selenodonty less marked than in other merycopotamines, we question if a high degree of ecological specialization may explain its extinction.
|
||
</paragraph>
|
||
<paragraph id="B64C36EAFFB59D1C4BC5FE17FE61B434" blockId="26.[163,782,197,1905]" pageId="26" pageNumber="304">
|
||
The basal position of
|
||
<taxonomicName id="71F34D69FFB59D1C4ABFFE17FD2AB26D" authorityName="Forster-Cooper" authorityYear="1915" box="[449,642,504,525]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="26" pageNumber="304" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB59D1C4ABFFE17FD2AB26D" box="[449,642,504,525]" italics="true" pageId="26" pageNumber="304">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
in relation to
|
||
<taxonomicName id="71F34D69FFB59D1C4BBBFDF9FE39B24C" box="[197,401,534,556]" pageId="26" pageNumber="304" rank="tribe" tribe="Merycopotamini">Merycopotamini</taxonomicName>
|
||
, its relatively short temporal range and atypical morphology (for a bothriodontine), as well as a fossil record limited to the Indian subcontinent, blurs its palaeobiogeographical history. Furthermore, Asian records of
|
||
<taxonomicName id="71F34D69FFB59D1C4A81FD7EFCA2B2C6" authorityName="Geais" authorityYear="1934" box="[511,778,657,679]" class="Mammalia" family="Anthracotheriidae" genus="Elomeryx" higherTaxonomySource="GBIF" isUncertain="true" kingdom="Animalia" order="Artiodactyla" pageId="26" pageNumber="304" phylum="Chordata" rank="species" species="borbonicus">
|
||
<emphasis id="8487EAF8FFB59D1C4A81FD7EFDC2B2C6" box="[511,618,657,678]" italics="true" pageId="26" pageNumber="304">Elomeryx</emphasis>
|
||
cf.
|
||
<emphasis id="8487EAF8FFB59D1C49F1FD7EFCA2B2C6" box="[655,778,657,678]" italics="true" pageId="26" pageNumber="304">borbonicus</emphasis>
|
||
</taxonomicName>
|
||
are scarce (
|
||
<bibRefCitation id="D2624B1BFFB59D1C4A58FD5FFDFDB2A6" author="Ducrocq S & Lihoreau F" box="[294,597,688,710]" pageId="26" pageNumber="304" pagination="885 - 891" refId="ref24181" refString="Ducrocq S, Lihoreau F. 2006. The occurrence of bothriodontines (Artiodactyla, Mammalia) in the Paleogene of Asia with special reference to Elomeryx: paleobiogeographical implications. Journal of Asian Earth Sciences 27: 885 - 891." type="journal article" year="2006">Ducrocq & Lihoreau, 2006</bibRefCitation>
|
||
), and there is a large gap with the record of
|
||
<taxonomicName id="71F34D69FFB59D1C4A99FD21FD66B283" box="[487,718,718,739]" class="Mammalia" family="Anthracotheriidae" genus="Bakalovia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="26" pageNumber="304" phylum="Chordata" rank="species" species="orientalis">
|
||
<emphasis id="8487EAF8FFB59D1C4A99FD21FD66B283" box="[487,718,718,739]" italics="true" pageId="26" pageNumber="304">BakaloƲia orientalis</emphasis>
|
||
</taxonomicName>
|
||
from the Late Eocene (
|
||
<bibRefCitation id="D2624B1BFFB59D1C4A0FFD02FDE3B362" author="Bohme M & Aiglstorfer M & Antoine PO & Appel E & Havlik P & Metais G & Phuc LT & Schneider S & Setzer F & Tappert R & Tran DN & Uhl D & Prieto J" box="[369,587,749,771]" pageId="26" pageNumber="304" pagination="121 - 167" refId="ref23806" refString="Bohme M, Aiglstorfer M, Antoine PO, Appel E, Havlik P, Metais G, Phuc LT, Schneider S, Setzer F, Tappert R, Tran DN, Uhl D, Prieto J. 2013. Na Duong (northern Vietnam) - an exceptional window into Eocene ecosystems from Southeast Asia. Zitteliana Reihe A: Mitteilungen der Bayerischen Staatssammlung fur Palaontologie und Geologie 53: 121 - 167." type="journal article" year="2013">
|
||
Böhme
|
||
<emphasis id="8487EAF8FFB59D1C4AB4FD01FDADB362" box="[458,517,749,771]" italics="true" pageId="26" pageNumber="304">et al.</emphasis>
|
||
, 2013
|
||
</bibRefCitation>
|
||
), which contributes to the uncertainty of the geographical origin of
|
||
<taxonomicName id="71F34D69FFB59D1C4BDDFCC5FECCB35F" authorityName="Forster-Cooper" authorityYear="1915" box="[163,356,810,831]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="26" pageNumber="304" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB59D1C4BDDFCC5FECCB35F" box="[163,356,810,831]" italics="true" pageId="26" pageNumber="304">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation id="2EC82A6FFFB59D1C4A0BFCC5FE6FB320" box="[373,455,810,832]" captionStart="Figure 11" captionStartId="25.[147,228,987,1009]" captionTargetBox="[146,1425,200,954]" captionTargetId="figure-356@25.[143,1430,197,957]" captionTargetPageId="25" captionText="Figure 11. Part of the consensus tree mapped on the geologic time scale (Gradstein et al., 2012) and the geographic distribution of selected bothriodontines. The temporal extensions of taxa are coloured according to their geographical distribution, those in non-full lines are uncertain. Green, Indian subcontinent; blue, Europe; purple, North America; orange, Africa. The non-ambiguous synapomorphies are placed at the nodes discussed in this work, the character states are summarised in the Supporting Information (Appendix S2); the non-ambiguous synapomorphies defining Merycopotamini in the analysis of Lihoreau et al. (2016) are in red; the non-ambiguous and non-homoplasic synapomorphies are in bold. Drawings of the P/4 are (from top to bottom) Parabrachyodus hyopotamoides (this study), Afromeryx zelteni, SiƲameryx africanus, Merycopotamus medioximus and Libycosaurus anisae (Lihoreau et al., 2019). The characters highlighted in red on the P/4 are those acquired in the clade involved. Drawings are not to scale. Spatial and temporal distributions are from Lihoreau et al. (2007, 2016, 2019), Bhandari et al. (2010), Holroyd et al. (2010), Kostopoulos et al. (2012), Antoine et al. (2013), Böhme et al. (2013) and Kapur et al. (2019)." figureDoi="http://doi.org/10.5281/zenodo.7927085" httpUri="https://zenodo.org/record/7927085/files/figure.png" pageId="26" pageNumber="304">Fig. 11</figureCitation>
|
||
).
|
||
<taxonomicName id="71F34D69FFB59D1C4AA1FCC5FD21B35F" authorityName="Douglass" authorityYear="1901" box="[479,649,810,831]" class="Mammalia" family="Anthracotheriidae" genus="Arretotherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="26" pageNumber="304" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB59D1C4AA1FCC5FD21B35F" box="[479,649,810,831]" italics="true" pageId="26" pageNumber="304">Arretotherium</emphasis>
|
||
</taxonomicName>
|
||
, and especially
|
||
<taxonomicName id="71F34D69FFB59D1C4B94FCA6FE3EB33E" box="[234,406,841,862]" class="Mammalia" family="Anthracotheriidae" genus="Arretotherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="26" pageNumber="304" phylum="Chordata" rank="species" species="meridionale">
|
||
<emphasis id="8487EAF8FFB59D1C4B94FCA6FE3EB33E" box="[234,406,841,862]" italics="true" pageId="26" pageNumber="304">A. meridionale</emphasis>
|
||
</taxonomicName>
|
||
from Central America, must be included in further phylogenetic analyses, since the phylogenetic position of this genus is unclear, either in an
|
||
<taxonomicName id="71F34D69FFB59D1C4B96FC4AFEFEB3DA" baseAuthorityName="Hellmund" baseAuthorityYear="1991" box="[232,342,933,954]" class="Mammalia" family="Anthracotheriidae" genus="Elomeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="26" pageNumber="304" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB59D1C4B96FC4AFEFEB3DA" box="[232,342,933,954]" italics="true" pageId="26" pageNumber="304">Elomeryx</emphasis>
|
||
</taxonomicName>
|
||
clade (
|
||
<bibRefCitation id="D2624B1BFFB59D1C4AD5FC4AFD6AB3DA" author="Kostopoulos DS & Koufos GD & Christanis K" box="[427,706,933,955]" pageId="26" pageNumber="304" pagination="303 - 315" refId="ref24912" refString="Kostopoulos DS, Koufos GD, Christanis K. 2012. On some anthracotheriid (Artiodactyla, Mammalia) remains from northern Greece: comments on the palaeozoogeography and phylogeny of Elomeryx. Saeiss Journal of Palaeontology 131: 303 - 315." type="journal article" year="2012">
|
||
Kostopoulos
|
||
<emphasis id="8487EAF8FFB59D1C493FFC49FDD4B3DA" box="[577,636,933,955]" italics="true" pageId="26" pageNumber="304">et al.</emphasis>
|
||
, 2012
|
||
</bibRefCitation>
|
||
) or in a polytomy with
|
||
<taxonomicName id="71F34D69FFB59D1C4A12FC2BFDA5B3B9" authorityName="Geais" authorityYear="1934" box="[364,525,964,985]" class="Mammalia" family="Anthracotheriidae" genus="Elomeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="26" pageNumber="304" phylum="Chordata" rank="species" species="borbonicus">
|
||
<emphasis id="8487EAF8FFB59D1C4A12FC2BFDA5B3B9" box="[364,525,964,985]" italics="true" pageId="26" pageNumber="304">E. borbonicus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="71F34D69FFB59D1C4935FC2BFCA3B3BA" box="[587,779,964,986]" pageId="26" pageNumber="304" rank="tribe" tribe="Merycopotamini">Merycopotamini</taxonomicName>
|
||
(
|
||
<bibRefCitation id="D2624B1BFFB59D1C4BD5FC0DFE7CB397" author="Lihoreau F & Ducrocq S" box="[171,468,994,1016]" pageId="26" pageNumber="304" pagination="89 - 105" refId="ref25128" refString="Lihoreau F, Ducrocq S. 2007. Family Anthracotheriidae. In: Protero DR, Foss SE, eds. The eVolution of artiodactyls. Baltimore: John Hopkins University Press, 89 - 105." type="book chapter" year="2007">Lihoreau & Ducrocq, 2007</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="D2624B1BFFB59D1C4A9EFC0DFD18B397" author="Bohme M & Aiglstorfer M & Antoine PO & Appel E & Havlik P & Metais G & Phuc LT & Schneider S & Setzer F & Tappert R & Tran DN & Uhl D & Prieto J" box="[480,688,994,1016]" pageId="26" pageNumber="304" pagination="121 - 167" refId="ref23806" refString="Bohme M, Aiglstorfer M, Antoine PO, Appel E, Havlik P, Metais G, Phuc LT, Schneider S, Setzer F, Tappert R, Tran DN, Uhl D, Prieto J. 2013. Na Duong (northern Vietnam) - an exceptional window into Eocene ecosystems from Southeast Asia. Zitteliana Reihe A: Mitteilungen der Bayerischen Staatssammlung fur Palaontologie und Geologie 53: 121 - 167." type="journal article" year="2013">
|
||
Böhme
|
||
<emphasis id="8487EAF8FFB59D1C494BFC0CFDC5B397" box="[565,621,994,1016]" italics="true" pageId="26" pageNumber="304">et al.</emphasis>
|
||
, 2013
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="D2624B1BFFB59D1C49C2FC0DFE8CB476" author="Rincon AF & Bloch JI & MacFadden BJ & Jaramillo CA" pageId="26" pageNumber="304" pagination="421 - 433" refId="ref26853" refString="Rincon AF, Bloch JI, MacFadden BJ, Jaramillo CA. 2013. First Central American record of Anthracotheriidae (Mammalia, Bothriodontinae) from the early Miocene of Panama. Journal of Vertebrate Paleontology 33: 421 - 433." type="journal article" year="2013">
|
||
Rincon
|
||
<emphasis id="8487EAF8FFB59D1C4BDDFBEDFF75B476" box="[163,221,1025,1047]" italics="true" pageId="26" pageNumber="304">et al.</emphasis>
|
||
, 2013
|
||
</bibRefCitation>
|
||
). It is a critical point for biogeographical purposes regarding the concomitant origination of merycopotamines in Asia.
|
||
</paragraph>
|
||
<paragraph id="B64C36EAFFB59D1C4BC5FBB2FD39B711" blockId="26.[163,782,197,1905]" pageId="26" pageNumber="304">
|
||
The identification of
|
||
<taxonomicName id="71F34D69FFB59D1C4AD0FBB2FD83B412" authorityName="Pilgrim" authorityYear="1908" box="[430,555,1117,1138]" class="Mammalia" family="Anthracotheriidae" genus="Gonotelma" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="26" pageNumber="304" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB59D1C4AD0FBB2FD83B412" box="[430,555,1117,1138]" italics="true" pageId="26" pageNumber="304">Gonotelma</emphasis>
|
||
</taxonomicName>
|
||
in the same stratigraphic level as
|
||
<taxonomicName id="71F34D69FFB59D1C4A22FB93FDBDB4F1" authorityName="Forster-Cooper" authorityYear="1915" box="[348,533,1148,1169]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="26" pageNumber="304" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB59D1C4A22FB93FDBDB4F1" box="[348,533,1148,1169]" italics="true" pageId="26" pageNumber="304">Parabrachyodus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="71F34D69FFB59D1C495CFB93FD32B4F1" box="[546,666,1148,1169]" class="Mammalia" family="Anthracotheriidae" genus="Sivameryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="26" pageNumber="304" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB59D1C495CFB93FD32B4F1" box="[546,666,1148,1169]" italics="true" pageId="26" pageNumber="304">SiƲameryx</emphasis>
|
||
</taxonomicName>
|
||
, probably
|
||
<taxonomicName id="71F34D69FFB59D1C4BDDFB75FDE8B4CF" authority="(Antoine et al., 2013)" baseAuthorityName="Antoine" baseAuthorityYear="2013" box="[163,576,1178,1200]" class="Mammalia" family="Anthracotheriidae" genus="Telmatodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="26" pageNumber="304" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB59D1C4BDDFB75FE99B4CF" box="[163,305,1178,1199]" italics="true" pageId="26" pageNumber="304">Telmatodon</emphasis>
|
||
(
|
||
<bibRefCitation id="D2624B1BFFB59D1C4A3AFB75FD9FB4CF" author="Antoine PO & Metais G & Orliac MJ & Crochet JY & Flynn LJ & Marivaux L & Rajpar AR & Roohi G & Welcomme JL" box="[324,567,1178,1200]" pageId="26" pageNumber="304" pagination="400 - 422" refId="ref23524" refString="Antoine PO, Metais G, Orliac MJ, Crochet JY, Flynn LJ, Marivaux L, Rajpar AR, Roohi G, Welcomme JL. 2013. Mammalian Neogene biostratigraphy of the Sulaiman Province, Pakistan. In: Xiaoming W, Flynn LJ, Fortelius M, eds. Fossil mammals of Asia. New York: Columbia University Press, 400 - 422." type="book chapter" year="2013">
|
||
Antoine
|
||
<emphasis id="8487EAF8FFB59D1C4AD0FB74FE44B4CF" box="[430,492,1178,1200]" italics="true" pageId="26" pageNumber="304">et al.</emphasis>
|
||
, 2013
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="71F34D69FFB59D1C49FFFB74FE07B4AE" authority="(Lihoreau et al., 2016)" baseAuthorityName="Lihoreau" baseAuthorityYear="2016" class="Mammalia" family="Anthracotheriidae" genus="Hemimeryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="26" pageNumber="304" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB59D1C49FFFB74FCA3B4D0" box="[641,779,1179,1200]" italics="true" pageId="26" pageNumber="304">Hemimeryx</emphasis>
|
||
(
|
||
<bibRefCitation id="D2624B1BFFB59D1C4BD5FB56FE0DB4AF" author="Lihoreau F & Alloing-Seguier L & Antoine PO & Boisserie JR & Marivaux L & Metais G & Welcomme JL" box="[171,421,1209,1231]" pageId="26" pageNumber="304" pagination="947 - 957" refId="ref25524" refString="Lihoreau F, Alloing-Seguier L, Antoine PO, Boisserie JR, Marivaux L, Metais G, Welcomme JL. 2016. Enamel microstructure defines a major Paleogene hippopotamoid clade: the Merycopotamini (Cetartiodactyla, Hippopotamoidea). Historical Biology 29: 947 - 957." type="journal article" year="2016">
|
||
Lihoreau
|
||
<emphasis id="8487EAF8FFB59D1C4A5EFB55FEF4B4AE" box="[288,348,1209,1231]" italics="true" pageId="26" pageNumber="304">et al.</emphasis>
|
||
, 2016
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
, implies that the Bugti Hills faunas simultaneously comprised at least five phylogenetically related bothriodontines. Such diversity is not surprising for the megafaunas of the region, as nine distinct species of rhinocerotids are known to co-occur from Kumbi 4 (
|
||
<bibRefCitation id="D2624B1BFFB59D1C4A83FABDFD52B507" author="Antoine PO & Downing KF & Crochet JY & Duranthon F & Flynn LJ & Marivaux L & Metais G & Rajpar AR & Roohi G" box="[509,762,1362,1384]" pageId="26" pageNumber="304" pagination="139 - 194" refId="ref23458" refString="Antoine PO, Downing KF, Crochet JY, Duranthon F, Flynn LJ, Marivaux L, Metais G, Rajpar AR, Roohi G. 2010. A revision of Aceratherium blanfordi Lydekker, 1884 (Mammalia: Rhinocerotidae) from the Early Miocene of Pakistan: postcranials as a key. Zoological Journal of the Linnean Society 160: 139 - 194." type="journal article" year="2010">
|
||
Antoine
|
||
<emphasis id="8487EAF8FFB59D1C4915FABCFD04B507" box="[619,684,1362,1384]" italics="true" pageId="26" pageNumber="304">et al.</emphasis>
|
||
, 2010
|
||
</bibRefCitation>
|
||
), the lateral equivalent of SAM 4. The diversity of merycopotamines from the Early Miocene of the Bugti Hills is unique in that no homotaxic assemblages are known elsewhere. The available craniomandibular material for
|
||
<taxonomicName id="71F34D69FFB59D1C4A4EFA03FDA2B661" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[304,522,1516,1537]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="26" pageNumber="304" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFB59D1C4A4EFA03FDA2B661" box="[304,522,1516,1537]" italics="true" pageId="26" pageNumber="304">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
did neither allow us to define a sexual dimorphism in the studied populations, nor a semi-aquatic lifestyle, as for
|
||
<taxonomicName id="71F34D69FFB59D1C494DF9C6FE84B63D" authority="(Rowan et al., 2015)" baseAuthorityName="Rowan" baseAuthorityYear="2015" class="Mammalia" family="Anthracotheriidae" genus="Sivameryx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="26" pageNumber="304" phylum="Chordata" rank="genus">
|
||
<emphasis id="8487EAF8FFB59D1C494DF9C6FD05B65E" box="[563,685,1577,1598]" italics="true" pageId="26" pageNumber="304">SiƲameryx</emphasis>
|
||
(
|
||
<bibRefCitation id="D2624B1BFFB59D1C49C2F9C6FE8BB63D" author="Rowan J & Adrian B & Grossman A" pageId="26" pageNumber="304" pagination="92830541" refId="ref26954" refString="Rowan J, Adrian B, Grossman A. 2015. The first skull of SiVameryx africanus (Anthracotheriidae, Bothriodontinae) from the early Miocene of East Africa. Journal of Vertebrate Paleontology 35: e 92830537 - e 92830541." type="journal article" year="2015">
|
||
Rowan
|
||
<emphasis id="8487EAF8FFB59D1C4BDDF9A7FF75B63D" box="[163,221,1608,1629]" italics="true" pageId="26" pageNumber="304">et al.</emphasis>
|
||
, 2015
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
and contrary to what is observed in certain bothriodontines with a proven semi-aquatic lifestyle (
|
||
<bibRefCitation id="D2624B1BFFB59D1C4B99F96AFE06B6FA" author="Orliac MJ & Antoine PO & Charruault AL & Hervet S & Prodeo F & Duranthon F" box="[231,430,1669,1691]" pageId="26" pageNumber="304" pagination="265 - 278" refId="ref26366" refString="Orliac MJ, Antoine PO, Charruault AL, Hervet S, Prodeo F, Duranthon F. 2013. Specialization for amphibiosis in Brachyodus onoideus (Artiodactyla, Hippopotamoidea) from the Early Miocene of France. Saeiss Journal of Geosciences 106: 265 - 278." type="journal article" year="2013">
|
||
Orliac
|
||
<emphasis id="8487EAF8FFB59D1C4A4AF969FEC3B6FA" box="[308,363,1669,1691]" italics="true" pageId="26" pageNumber="304">et al.</emphasis>
|
||
, 2013
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="D2624B1BFFB59D1C4AC4F96AFD0AB6FA" author="Lihoreau F & Boisserie JR & Blondel C & Jacques L & Likius A & Mackaye HT & Vignaud P & Brunet M" box="[442,674,1669,1691]" pageId="26" pageNumber="304" pagination="761 - 798" refId="ref25431" refString="Lihoreau F, Boisserie JR, Blondel C, Jacques L, Likius A, Mackaye HT, Vignaud P, Brunet M. 2014. Description and palaeobiology of a new species of Libycosaurus (Cetartiodactyla, Anthracotheriidae) from the Late Miocene of Toros-Menalla, northern Chad. Journal of Systematic Palaeontology 12: 761 - 798." type="journal article" year="2014">
|
||
Lihoreau
|
||
<emphasis id="8487EAF8FFB59D1C4956F969FDF7B6FA" box="[552,607,1669,1691]" italics="true" pageId="26" pageNumber="304">et al.</emphasis>
|
||
, 2014
|
||
</bibRefCitation>
|
||
). Yet, the predominance of the remains of
|
||
<taxonomicName id="71F34D69FFB59D1C4958F94BFCA3B6D9" baseAuthorityName="Lydekker" baseAuthorityYear="1883" box="[550,779,1700,1721]" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="26" pageNumber="304" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFB59D1C4958F94BFCA3B6D9" box="[550,779,1700,1721]" italics="true" pageId="26" pageNumber="304">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
over other Bugti anthracotheres (
|
||
<bibRefCitation id="D2624B1BFFB59D1C495FF92DFD6DB6B7" author="Pickford M" box="[545,709,1730,1752]" pageId="26" pageNumber="304" pagination="289 - 350" refId="ref26447" refString="Pickford M. 1987. Revision des suiformes (Artiodactyla, Mammalia) de Bugti (Pakistan). Annales de Paleontologie 73: 289 - 350." type="journal article" year="1987">Pickford, 1987</bibRefCitation>
|
||
: table 6), and the proportion of unicuspidate P4/ for
|
||
<taxonomicName id="71F34D69FFB59D1C49A3F90DFEE4B774" baseAuthorityName="Lydekker" baseAuthorityYear="1883" class="Mammalia" family="Anthracotheriidae" genus="Parabrachyodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="26" pageNumber="304" phylum="Chordata" rank="species" species="hyopotamoides">
|
||
<emphasis id="8487EAF8FFB59D1C49A3F90DFEE4B774" italics="true" pageId="26" pageNumber="304">Par. hyopotamoides</emphasis>
|
||
</taxonomicName>
|
||
, show that these megaherbivores must have lived in sufficiently isolated and small populations for such dental variations to become common (
|
||
<bibRefCitation id="D2624B1BFFB59D1C4BD5F8B3FE22B711" author="Ducrocq S & Chaimanee Y & Suteethorn V & Jaeger JJ" box="[171,394,1883,1905]" pageId="26" pageNumber="304" pagination="355 - 360" refId="ref24223" refString="Ducrocq S, Chaimanee Y, Suteethorn V, Jaeger JJ. 1995. Dental anomalies in Upper Eocene Anthracotheriidae: a possible case of inbreeding. Lethaia 28: 355 - 360." type="journal article" year="1995">
|
||
Ducrocq
|
||
<emphasis id="8487EAF8FFB59D1C4A71F8B3FE97B710" box="[271,319,1883,1905]" italics="true" pageId="26" pageNumber="304">et al</emphasis>
|
||
., 1995
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="D2624B1BFFB59D1C4AE9F8B4FD29B711" author="Lihoreau F & Boisserie JR & Viriot L & Coppens Y & Likius A & Mackaye HT & Tafforeau P & Vignaud P & Brunet M" box="[407,641,1883,1905]" pageId="26" pageNumber="304" pagination="8763 - 8767" refId="ref25261" refString="Lihoreau F, Boisserie JR, Viriot L, Coppens Y, Likius A, Mackaye HT, Tafforeau P, Vignaud P, Brunet M. 2006. Anthracothere dental anatomy reveals a late Miocene Chado- Libyan bioprovince. Proceedings of the National Academy of Sciences of the USA 103: 8763 - 8767." type="journal article" year="2006">
|
||
Lihoreau
|
||
<emphasis id="8487EAF8FFB59D1C4978F8B3FD96B710" box="[518,574,1883,1905]" italics="true" pageId="26" pageNumber="304">et al.</emphasis>
|
||
, 2006
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |