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<document id="D805CB217256639F35C816C8334E478D" ID-DOI="10.1111/j.1096-3642.2004.00138.x" ID-ISSN="0024-4082" ID-Zenodo-Dep="5431832" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1630429132170" checkinUser="diego" docAuthor="Boisserie, Jean-Renaud" docDate="2005" docId="354487FCFF8EFFC6FC3B3CD3A98CFAFE" docLanguage="en" docName="j.1096-3642.2004.00138.x.pdf" docOrigin="Zoological Journal of the Linnean Society 143 (1)" docSource="https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2004.00138.x" docStyle="DocumentStyle:0DD8C314D74634CE09062A86991413F8.2:ZoolJLinnSoc.2002-2009.journal_article" docStyleId="0DD8C314D74634CE09062A86991413F8" docStyleName="ZoolJLinnSoc.2002-2009.journal_article" docStyleVersion="2" docTitle="Saotherium Boisserie 2005, GEN. NOV." docType="treatment" docVersion="3" lastPageNumber="15" masterDocId="C97DFF84FF85FFC8FF903A26A878FFE5" masterDocTitle="The phylogeny and taxonomy of Hippopotamidae (Mammalia: Artiodactyla): a review based on morphology and cladistic analysis" masterLastPageNumber="26" masterPageNumber="1" pageNumber="12" updateTime="1699266389593" updateUser="plazi" zenodo-license-document="CC-BY-4.0" zenodo-license-figures="CC-BY-4.0">
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<mods:title id="4C898BD9EA244BE86426EA45B3E86BD6">The phylogeny and taxonomy of Hippopotamidae (Mammalia: Artiodactyla): a review based on morphology and cladistic analysis</mods:title>
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<mods:namePart id="A813370E7AABD18ED849372181154B35">Boisserie, Jean-Renaud</mods:namePart>
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<mods:title id="55FF7E316468DC5EC696557EDB373592">Zoological Journal of the Linnean Society</mods:title>
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<mods:date id="50EBE5BDBB45EEC48356821E4C83B2E9">2005</mods:date>
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<subSubSection id="F5F76561FF8EFFC3FC3B3CD3AD5AF8E8" box="[939,1314,1781,1805]" pageId="11" pageNumber="12" type="nomenclature">
<paragraph id="BD5236EAFF8EFFC3FC3B3CD3AD5AF8E8" blockId="11.[907,1346,1685,1805]" box="[939,1314,1781,1805]" pageId="11" pageNumber="12">
<heading id="E61A8186FF8EFFC3FC3B3CD3AD5AF8E8" box="[939,1314,1781,1805]" centered="true" fontSize="9" level="2" pageId="11" pageNumber="12" reason="2">
GENUS
<taxonomicName id="7AED4D69FF8EFFC3FB943CD3ACD9F8E9" authority="Boisserie, 2005" authorityName="Boisserie" authorityYear="2005" box="[1028,1185,1781,1805]" class="Mammalia" family="Hippopotamidae" genus="Saotherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="11" pageNumber="12" phylum="Chordata" rank="genus" status="gen. nov.">SAOTHERIUM</taxonomicName>
<emphasis id="8F99EAF8FF8EFFC3FB393CDFAD5AF8E8" bold="true" box="[1193,1314,1781,1805]" pageId="11" pageNumber="12">
<taxonomicNameLabel id="94AA5783FF8EFFC3FB393CDFAD5AF8E8" box="[1193,1314,1781,1805]" pageId="11" pageNumber="12" rank="genus">GEN. NOV.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
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<subSubSection id="F5F76561FF8EFFC6FCA33D38A98CFAFE" lastPageId="14" lastPageNumber="15" pageId="11" pageNumber="12" type="description">
<paragraph id="BD5236EAFF8EFFC3FCA33D38ABC9F8D6" blockId="11.[819,1433,1822,1905]" box="[819,945,1822,1843]" pageId="11" pageNumber="12">
<emphasis id="8F99EAF8FF8EFFC3FCA33D38ABC9F8D6" box="[819,945,1822,1843]" italics="true" pageId="11" pageNumber="12">Description</emphasis>
</paragraph>
<paragraph id="BD5236EAFF8EFFC3FCA33D1BADE1F894" blockId="11.[819,1433,1822,1905]" pageId="11" pageNumber="12">
<emphasis id="8F99EAF8FF8EFFC3FCA33D1BABD2F8B7" box="[819,938,1853,1874]" italics="true" pageId="11" pageNumber="12">Diagnosis:</emphasis>
Hexaprotodont, with the following apomorphies: cranial roof showing an antorbital angle
</paragraph>
<caption id="E9926662FF89FFC4FF1E3C30AD2CF889" ID-DOI="http://doi.org/10.5281/zenodo.5431850" ID-Zenodo-Dep="5431850" httpUri="https://zenodo.org/record/5431850/files/figure.png" pageId="12" pageNumber="13" startId="12.[142,220,1558,1577]" targetBox="[232,1332,193,1524]" targetPageId="12">
<paragraph id="BD5236EAFF89FFC4FF1E3C30AD2CF889" blockId="12.[142,1422,1558,1900]" pageId="12" pageNumber="13">
<emphasis id="8F99EAF8FF89FFC4FF1E3C30A88EF9CC" bold="true" box="[142,246,1558,1577]" pageId="12" pageNumber="13">Figure 9.</emphasis>
Mandibular anatomy within the
<taxonomicName id="7AED4D69FF89FFC4FDCC3C30AB74F9CC" authorityName="GRAY" authorityYear="1821" box="[604,780,1558,1577]" class="Mammalia" family="Hippopotamidae" kingdom="Animalia" order="Artiodactyla" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Hippopotamidae</taxonomicName>
. This figure shows the new taxonomic divisions of the family
<taxonomicName id="7AED4D69FF89FFC4FF1E3C15A943F9A3" authorityName="GRAY" authorityYear="1821" box="[142,315,1587,1606]" class="Mammalia" family="Hippopotamidae" kingdom="Animalia" order="Artiodactyla" pageId="12" pageNumber="13" phylum="Chordata" rank="family">Hippopotamidae</taxonomicName>
and, for each discussed taxon, some of the mandibular characters that provided additional support to the clades identified in the parsimony analysis (boxes in this figure). These features include: the general shape of the mandible, with expansion of the canine processes and relative length of the symphysis (seen in the dorsal outlines); the shape of the symphysis sagittal cross section; the length of the premolar row relative to the length of the molar row. The figure shows the following features for the taxa listed under each genus name:
<taxonomicName id="7AED4D69FF89FFC4FC873C8FABEAF959" authorityName="Boisserie" authorityYear="2005" box="[791,914,1705,1724]" class="Mammalia" family="Hippopotamidae" genus="Saotherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="8F99EAF8FF89FFC4FC873C8FABEAF959" box="[791,914,1705,1724]" italics="true" pageId="12" pageNumber="13">Saotherium</emphasis>
</taxonomicName>
, very inclined symphysis with thin cross-section and poorly developed canine processes;
<taxonomicName id="7AED4D69FF89FFC4FDAD3CE0AA92F93C" authorityName="Boisserie" authorityYear="2005" box="[573,746,1734,1753]" class="Mammalia" family="Hippopotamidae" genus="Archaeopotamus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="8F99EAF8FF89FFC4FDAD3CE0AA92F93C" box="[573,746,1734,1753]" italics="true" pageId="12" pageNumber="13">Archaeopotamus</emphasis>
</taxonomicName>
, relatively long and shallow symphysis with poorly developed canine processes and longer premolar rows than in any other clade;
<taxonomicName id="7AED4D69FF89FFC4FCFD3CC5AB87F913" authorityName="Falconer &amp; Cautley" authorityYear="1836" box="[877,1023,1763,1782]" class="Mammalia" family="Hippopotamidae" genus="Hexaprotodon" kingdom="Animalia" order="Artiodactyla" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="8F99EAF8FF89FFC4FCFD3CC5AB87F913" box="[877,1023,1763,1782]" italics="true" pageId="12" pageNumber="13">Hexaprotodon</emphasis>
</taxonomicName>
, wide symphysis but with poorly differentiated canine processes, very robust symphysis in cross section;
<taxonomicName id="7AED4D69FF89FFC4FCC63D27ABB0F8F1" authorityName="LEIDY" authorityYear="1853" box="[854,968,1793,1812]" class="Mammalia" family="Hippopotamidae" genus="Choeropsis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="8F99EAF8FF89FFC4FCC63D27ABB0F8F1" box="[854,968,1793,1812]" italics="true" pageId="12" pageNumber="13">Choeropsis</emphasis>
</taxonomicName>
, very short symphysis globular in cross section and poorly developed canine processes;
<taxonomicName id="7AED4D69FF89FFC4FDE13D39AB72F8D4" authorityName="Linnaeus" authorityYear="1758" box="[625,778,1823,1841]" class="Mammalia" family="Hippopotamidae" genus="Hippopotamus" kingdom="Animalia" order="Artiodactyla" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="8F99EAF8FF89FFC4FDE13D39AB72F8D4" box="[625,778,1823,1841]" italics="true" pageId="12" pageNumber="13">Hippopotamus</emphasis>
</taxonomicName>
and aff.
<taxonomicName id="7AED4D69FF89FFC4FCE03D39AC71F8D4" authorityName="Linnaeus" authorityYear="1758" box="[880,1033,1823,1841]" class="Mammalia" family="Hippopotamidae" genus="Hippopotamus" kingdom="Animalia" order="Artiodactyla" pageId="12" pageNumber="13" phylum="Chordata" rank="genus">
<emphasis id="8F99EAF8FF89FFC4FCE03D39AC71F8D4" box="[880,1033,1823,1841]" italics="true" pageId="12" pageNumber="13">Hippopotamus</emphasis>
</taxonomicName>
, short symphysis globular in crosssection (lacking a projected incisor alveolar process) and strong extension of the canine processes the latter feature being not salient in the Afar species (aff.
<emphasis id="8F99EAF8FF89FFC4FD933D7FAAE4F889" box="[515,668,1881,1900]" italics="true" pageId="12" pageNumber="13">Hip. coryndoni</emphasis>
, aff.
<taxonomicName id="7AED4D69FF89FFC4FD423D7FAB1BF889" authorityName="Geze" authorityYear="1985" box="[722,867,1881,1900]" class="Mammalia" family="Hippopotamidae" genus="Trilobophorus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="afarensis">
<emphasis id="8F99EAF8FF89FFC4FD423D7FAB1BF889" box="[722,867,1881,1900]" italics="true" pageId="12" pageNumber="13">Hip. afarensis</emphasis>
</taxonomicName>
) and aff.
<taxonomicName id="7AED4D69FF89FFC4FC593D7FACCCF889" box="[969,1204,1881,1900]" class="Mammalia" family="Hippopotamidae" genus="Hexaprotodon" kingdom="Animalia" order="Artiodactyla" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="protamphibius">
<emphasis id="8F99EAF8FF89FFC4FC593D7FAB8FF88E" box="[969,1015,1881,1899]" italics="true" pageId="12" pageNumber="13">Hip.</emphasis>
cf.
<emphasis id="8F99EAF8FF89FFC4FB8A3D7FACCCF889" box="[1050,1204,1881,1900]" italics="true" pageId="12" pageNumber="13">protamphibius</emphasis>
</taxonomicName>
from Kanapoi.
</paragraph>
</caption>
<paragraph id="BD5236EAFF88FFC5FF0B3E26A892F9FA" lastBlockId="13.[155,771,1238,1567]" pageId="13" pageNumber="14">10Mya in lateral view; skull very high above the molars; slender mandibular symphysis in the sagittal plane. Also exhibiting these plesiomorphic or convergent features: orbits below the cranial roof; slender zygomatic arches; cylindrical braincase; slender and low sagittal crest; laterally developed occipital plate; maxillary process of the frontal separating the nasal and the lachrymal bones; short extension of the canine processes; lingual border of the lower cheek tooth alveolar process lower than the labial border.</paragraph>
<caption id="E9926662FF88FFC5FF0B3E14ADE3FB65" ID-DOI="http://doi.org/10.5281/zenodo.5431852" ID-Zenodo-Dep="5431852" httpUri="https://zenodo.org/record/5431852/files/figure.png" pageId="13" pageNumber="14" startId="13.[155,233,1074,1093]" targetBox="[163,1432,206,1028]" targetPageId="13">
<paragraph id="BD5236EAFF88FFC5FF0B3E14ADE3FB65" blockId="13.[155,1435,1071,1153]" pageId="13" pageNumber="14">
<emphasis id="8F99EAF8FF88FFC5FF0B3E14A968FBA0" bold="true" box="[155,272,1074,1093]" pageId="13" pageNumber="14">Figure 10.</emphasis>
A new phylogeny for the
<taxonomicName id="7AED4D69FF88FFC5FD893E14AAB1FBA0" authorityName="GRAY" authorityYear="1821" box="[537,713,1074,1093]" class="Mammalia" family="Hippopotamidae" kingdom="Animalia" order="Artiodactyla" pageId="13" pageNumber="14" phylum="Chordata" rank="family">Hippopotamidae</taxonomicName>
. Geographical distribution:
<superScript id="4A989BA2FF88FFC5FC773E09AB97FBDE" attach="right" box="[999,1007,1071,1083]" fontSize="5" pageId="13" pageNumber="14">a</superScript>
not Eastern African, but from Abu Dhabi, the Arab United Emirates, the Arabic Peninsula (see
<bibRefCitation id="D97C4B1BFF88FFC5FD533E76AB32FB86" author="Gentry AW" box="[707,842,1104,1123]" pageId="13" pageNumber="14" pagination="271 - 289" refId="ref16641" refString="Gentry AW. 1999. A fossil hippopotamus from the Emirate of Abu Dhabi, United Arab Emirates. In: Whybrow PJ, Hill A, eds. Fossil vertebrates of Arabia. New Haven, CT: Yale University Press, 271 - 289." type="book chapter" year="1999">Gentry, 1999</bibRefCitation>
);
<superScript id="4A989BA2FF88FFC5FCCA3E6AAB1AFBBD" attach="right" box="[858,866,1100,1112]" fontSize="5" pageId="13" pageNumber="14">b</superScript>
known in Eastern Africa but also in Oubeidiyeh, Israel (see
<bibRefCitation id="D97C4B1BFF88FFC5FF593E48A93DFB65" author="Faure M" box="[201,325,1133,1153]" pageId="13" pageNumber="14" pagination="107 - 142" refId="ref16314" refString="Faure M. 1986. Les Hippopotamides du Pleistocene ancien d'Oubeidiyeh (Israel). Memoires et Travaux du Centre de Recherche Francais de Jerusalem 5: 107 - 142." type="journal article" year="1986">Faure, 1986</bibRefCitation>
) and maybe in Algeria (
<bibRefCitation id="D97C4B1BFF88FFC5FDAE3E4BAAADFB65" author="Geraads D" box="[574,725,1133,1152]" pageId="13" pageNumber="14" pagination="65 - 94" refId="ref16729" refString="Geraads D. 1980. La faune des sites a ' Homo erectus ' des carrieres Thomas (Casablanca, Maroc). Quaternaria 22: 65 - 94." type="journal article" year="1980">Geraads, 1980</bibRefCitation>
);
<superScript id="4A989BA2FF88FFC5FD763E4CAA94FB93" attach="right" box="[742,748,1130,1142]" fontSize="5" pageId="13" pageNumber="14">c</superScript>
known in Africa but also in continental Europe (see
<bibRefCitation id="D97C4B1BFF88FFC5FA9A3E48ADE8FB65" author="Mazza P" box="[1290,1424,1133,1153]" pageId="13" pageNumber="14" pagination="61 - 241" refId="ref17721" refString="Mazza P. 1995. New evidence on the Pleistocene hippopotamuses of Western Europe. Geologica Romana 31: 61 - 241." type="journal article" year="1995">Mazza, 1995</bibRefCitation>
).
</paragraph>
</caption>
<paragraph id="BD5236EAFF88FFC5FF0B3C1BAA20F994" blockId="13.[155,770,1596,1649]" pageId="13" pageNumber="14">
<materialsCitation id="0D853CB7FF88FFC5FF0B3C1BAA2CF994" country="Chad" location="Kolle" municipality="Lower Pliocene" pageId="13" pageNumber="14" specimenCount="1" typeStatus="holotype">
<emphasis id="8F99EAF8FF88FFC5FF0B3C1BAA62F9B4" box="[155,538,1596,1618]" italics="true" pageId="13" pageNumber="14">
<typeStatus id="62568848FF88FFC5FF0B3C1BA8A8F9B7" box="[155,208,1597,1618]" pageId="13" pageNumber="14">Type</typeStatus>
species:
<taxonomicName id="7AED4D69FF88FFC5FEA93C1AAA62F9B4" authorityName="Boisserie" authorityYear="2005" box="[313,538,1596,1617]" class="Mammalia" family="Hippopotamidae" genus="Saotherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="mingoz">Saotherium mingoz</taxonomicName>
</emphasis>
, from
<location id="B8326031FF88FFC5FDF93C1AAADDF9B7" LSID="urn:lsid:plazi:treatment:354487FCFF8EFFC6FC3B3CD3A98CFAFE:B8326031FF88FFC5FDF93C1AAADDF9B7" box="[617,677,1596,1618]" country="Chad" municipality="Lower Pliocene" name="Kolle" pageId="13" pageNumber="14">Kollé</location>
(
<collectingCountry id="C5FA767AFF88FFC5FD263C1AAA8EF9B7" box="[694,758,1596,1618]" name="Chad" pageId="13" pageNumber="14">Chad</collectingCountry>
),
<collectingMunicipality id="5D36AC90FF88FFC5FF0B3C7DA931F995" box="[155,329,1626,1648]" pageId="13" pageNumber="14">Lower Pliocene</collectingMunicipality>
(
<bibRefCitation id="D97C4B1BFF88FFC5FEC93C7CAA34F994" author="Boisserie JR &amp; Brunet M &amp; Likius A &amp; Vignaud P" box="[345,588,1626,1649]" pageId="13" pageNumber="14" pagination="15 - 27" refId="ref15268" refString="Boisserie JR, Brunet M, Likius A, Vignaud P. 2003. Hippopotamids from the Djurab Pliocene faunas, Chad, Central Africa. Journal of African Earth Sciences 36 (1 - 2): 15 - 27." type="journal article" year="2003">
Boisserie
<emphasis id="8F99EAF8FF88FFC5FE5B3C7DAA7BF98A" box="[459,515,1626,1648]" italics="true" pageId="13" pageNumber="14">et al.</emphasis>
, 2003
</bibRefCitation>
)
</materialsCitation>
.
</paragraph>
<paragraph id="BD5236EAFF88FFC5FF0B3CA8A899F904" blockId="13.[155,771,1678,1761]" pageId="13" pageNumber="14">
<emphasis id="8F99EAF8FF88FFC5FF0B3CA8A934F946" box="[155,332,1678,1699]" italics="true" pageId="13" pageNumber="14">Other material:</emphasis>
<taxonomicName id="7AED4D69FF88FFC5FEC93CA8AA23F946" authorityName="Boisserie" authorityYear="2005" box="[345,603,1678,1700]" class="Mammalia" family="Hippopotamidae" genus="Saotherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="mingoz">
<emphasis id="8F99EAF8FF88FFC5FEC93CA8A9A7F946" box="[345,479,1678,1699]" italics="true" pageId="13" pageNumber="14">Saotherium</emphasis>
cf.
<emphasis id="8F99EAF8FF88FFC5FD993CA8AA23F946" box="[521,603,1678,1699]" italics="true" pageId="13" pageNumber="14">mingoz</emphasis>
</taxonomicName>
, from Kossom Bougoudi (
<collectingCountry id="C5FA767AFF88FFC5FE8F3C8AA927F927" box="[287,351,1708,1730]" name="Chad" pageId="13" pageNumber="14">Chad</collectingCountry>
), Lower Pliocene (
<bibRefCitation id="D97C4B1BFF88FFC5FDDA3C8AA8AEF904" author="Boisserie JR &amp; Brunet M &amp; Likius A &amp; Vignaud P" pageId="13" pageNumber="14" pagination="15 - 27" refId="ref15268" refString="Boisserie JR, Brunet M, Likius A, Vignaud P. 2003. Hippopotamids from the Djurab Pliocene faunas, Chad, Central Africa. Journal of African Earth Sciences 36 (1 - 2): 15 - 27." type="journal article" year="2003">
Boisserie
<emphasis id="8F99EAF8FF88FFC5FD543C8BAA84F924" box="[708,764,1708,1730]" italics="true" pageId="13" pageNumber="14">et al.</emphasis>
, 2003
</bibRefCitation>
).
</paragraph>
<paragraph id="BD5236EAFF88FFC5FF0B3CD8A89BF895" blockId="13.[155,770,1790,1904]" pageId="13" pageNumber="14">
<emphasis id="8F99EAF8FF88FFC5FF0B3CD8A97EF8F6" box="[155,262,1790,1811]" italics="true" pageId="13" pageNumber="14">Remarks:</emphasis>
The diagnosis is the same as for the
<typeStatus id="62568848FF88FFC5FD403CD9AB79F8F1" box="[720,769,1791,1812]" pageId="13" pageNumber="14">type</typeStatus>
species, excluding those characters that differentiate the two known taxa, or that have not been seen in both forms.
</paragraph>
<paragraph id="BD5236EAFF88FFC5FCA33EF0AD11FAEE" blockId="13.[819,1433,1238,1291]" pageId="13" pageNumber="14">
<emphasis id="8F99EAF8FF88FFC5FCA33EF0ABC9FB0E" box="[819,945,1238,1259]" italics="true" pageId="13" pageNumber="14">Etymology:</emphasis>
From the Sao, an enigmatic medieval civilization known in the
<collectingCountry id="C5FA767AFF88FFC5FBA93ED3AC0EFAEE" box="[1081,1142,1269,1291]" name="Chad" pageId="13" pageNumber="14">Chad</collectingCountry>
basin (
<bibRefCitation id="D97C4B1BFF88FFC5FB5C3ED3AD25FAEE" author="Lange D" box="[1228,1373,1269,1291]" pageId="13" pageNumber="14" pagination="189 - 210" refId="ref17411" refString="Lange D. 1989. Preliminaires pour un histoire des Sao. The Journal of African History 30 (2): 189 - 210." type="journal article" year="1989">Lange, 1989</bibRefCitation>
).
</paragraph>
<paragraph id="BD5236EAFF88FFC5FCA33F05ACF8FABD" blockId="13.[819,1434,1315,1368]" pageId="13" pageNumber="14">
<emphasis id="8F99EAF8FF88FFC5FCA33F05AC18FADD" box="[819,1120,1315,1336]" italics="true" pageId="13" pageNumber="14">Geographical distribution:</emphasis>
Djurab desert, Lake
<collectingCountry id="C5FA767AFF88FFC5FACD3F05ADE2FADC" box="[1373,1434,1315,1337]" name="Chad" pageId="13" pageNumber="14">Chad</collectingCountry>
basin (
<collectingCountry id="C5FA767AFF88FFC5FC103F64ABC7FABD" box="[896,959,1346,1368]" name="Chad" pageId="13" pageNumber="14">Chad</collectingCountry>
, Central Africa).
</paragraph>
<paragraph id="BD5236EAFF88FFC5FCA33F56ACF0FA26" blockId="13.[819,1435,1392,1475]" pageId="13" pageNumber="14">
<emphasis id="8F99EAF8FF88FFC5FCA33F56AC4CFA60" box="[819,1076,1392,1413]" italics="true" pageId="13" pageNumber="14">Temporal distribution:</emphasis>
early Pliocene, between the Mio-Pliocene boundary and 4.0 Mya (see
<bibRefCitation id="D97C4B1BFF88FFC5FA943FA9AB14FA27" author="Brunet M &amp; Beauvilain A &amp; Geraads D &amp; Guy F &amp; Kasser M &amp; Mackaye HT &amp; Maclatchy LM &amp; Mouchelin G &amp; Sudre J &amp; Vignaud P" pageId="13" pageNumber="14" pagination="153 - 158" refId="ref15379" refString="Brunet M, Beauvilain A, Geraads D, Guy F, Kasser M, Mackaye HT, Maclatchy LM, Mouchelin G, Sudre J, Vignaud P. 1998. Tchad: decouverte d'une faune de Mammiferes du Pliocene inferieur. Comptes Rendus de l'Academie des Sciences Paris 326 (2 a): 153 - 158." type="journal article" year="1998">
Brunet
<emphasis id="8F99EAF8FF88FFC5FACC3FA9ADECFA41" box="[1372,1428,1423,1444]" italics="true" pageId="13" pageNumber="14">et al.</emphasis>
, 1998
</bibRefCitation>
;
<bibRefCitation id="D97C4B1BFF88FFC5FCE83F88AC05FA26" author="Brunet M &amp; MPFT" box="[888,1149,1453,1475]" pageId="13" pageNumber="14" pagination="205 - 209" refId="ref15344" refString="Brunet M, MPFT. 2000. Chad: discovery of a vertebrate fauna close to the Mio - Pliocene boundary. Journal of Vertebrate Paleontology 20 (1): 205 - 209." type="journal article" year="2000">Brunet &amp; MPFT, 2000</bibRefCitation>
).
</paragraph>
<paragraph id="BD5236EAFF88FFC5FCA33C2FABD2F9FB" blockId="13.[819,1434,1545,1905]" box="[819,938,1545,1566]" pageId="13" pageNumber="14">
<emphasis id="8F99EAF8FF88FFC5FCA33C2FABD2F9FB" box="[819,938,1545,1566]" italics="true" pageId="13" pageNumber="14">Discussion</emphasis>
</paragraph>
<paragraph id="BD5236EAFF88FFC6FCA33C0EA95EFD62" blockId="13.[819,1434,1545,1905]" lastBlockId="14.[142,759,196,1138]" lastPageId="14" lastPageNumber="15" pageId="13" pageNumber="14">
In their description (
<bibRefCitation id="D97C4B1BFF88FFC5FBBE3C0EAD5FF9DB" author="Boisserie JR &amp; Brunet M &amp; Likius A &amp; Vignaud P" box="[1070,1319,1576,1598]" pageId="13" pageNumber="14" pagination="15 - 27" refId="ref15268" refString="Boisserie JR, Brunet M, Likius A, Vignaud P. 2003. Hippopotamids from the Djurab Pliocene faunas, Chad, Central Africa. Journal of African Earth Sciences 36 (1 - 2): 15 - 27." type="journal article" year="2003">
Boisserie
<emphasis id="8F99EAF8FF88FFC5FB343C0FACA4F9D8" box="[1188,1244,1576,1598]" italics="true" pageId="13" pageNumber="14">et al.</emphasis>
, 2003
</bibRefCitation>
), the two Djurab Pliocene hippos are shown to possess an association of original cranial features: the antorbital angle of the cranial roof and the correlated anterior convergence of the nasal toward the palate; the important relative height of the skull above the molars; the elongated braincase with a rounded transversal section and a weak postorbital constriction (cylindrical aspect). In this respect, these hippopotamids differ considerably from the other known hippos. For this reason,
<bibRefCitation id="D97C4B1BFF88FFC5FBDA3D7CAD3BF894" author="Boisserie JR &amp; Brunet M &amp; Likius A &amp; Vignaud P" box="[1098,1347,1882,1905]" pageId="13" pageNumber="14" pagination="15 - 27" refId="ref15268" refString="Boisserie JR, Brunet M, Likius A, Vignaud P. 2003. Hippopotamids from the Djurab Pliocene faunas, Chad, Central Africa. Journal of African Earth Sciences 36 (1 - 2): 15 - 27." type="journal article" year="2003">
Boisserie
<emphasis id="8F99EAF8FF88FFC5FB2B3D7DAC8BF88A" box="[1211,1267,1882,1904]" italics="true" pageId="13" pageNumber="14">et al.</emphasis>
(2003)
</bibRefCitation>
evoked an independent hippo lineage in central Africa originating at the Mio-Pliocene boundary if not before. This opinion is confirmed by the position of these forms in the parsimony analysis (
<figureCitation id="25D62A6FFF8BFFC6FD8B3B06AA1BFED3" box="[539,611,288,310]" captionStart="Figure 6" captionStartId="8.[142,220,840,859]" captionTargetBox="[226,675,199,807]" captionTargetId="graphics-775@8.[230,670,345,808]" captionTargetPageId="8" captionText="Figure 6. Strict consensus tree of the three most parsimonious trees obtained from the cladistic analysis (length = 81, CI = 0.6420, RI = 0.7752). In italic-bold, the bootstrap results are given as an indication of clade robustness." figureDoi="http://doi.org/10.5281/zenodo.5431844" httpUri="https://zenodo.org/record/5431844/files/figure.png" pageId="14" pageNumber="15">Figs 6</figureCitation>
,
<figureCitation id="25D62A6FFF8BFFC6FDE23B06AAF9FED0" box="[626,641,288,309]" captionStart="Figure 7" captionStartId="9.[155,233,1397,1416]" captionTargetBox="[377,1209,196,1364]" captionTargetId="graphics-265@9.[402,1209,412,1364]" captionTargetPageId="9" captionText="Figure 7. Second of the three most parsimonious tree obtained from the cladistic analysis. The bold numbers indicate the nodes. The numbered boxes indicate the ACCTRAN character state changes (white boxes indicate reversions and convergences)." figureDoi="http://doi.org/10.5281/zenodo.5431846" httpUri="https://zenodo.org/record/5431846/files/figure.png" pageId="14" pageNumber="15">7</figureCitation>
), showing also many primitive traits in this morphology. The mandibular morphology, especially the symphysial sagittal section between the central incisors (see
<figureCitation id="25D62A6FFF8BFFC6FF1E3BBDA8B6FE54" box="[142,206,411,433]" captionStart="Figure 9" captionStartId="12.[142,220,1558,1577]" captionTargetBox="[232,1332,193,1524]" captionTargetId="graphics-250@12.[771,1332,193,1523]" captionTargetPageId="12" captionText="Figure 9. Mandibular anatomy within the Hippopotamidae. This figure shows the new taxonomic divisions of the family Hippopotamidae and, for each discussed taxon, some of the mandibular characters that provided additional support to the clades identified in the parsimony analysis (boxes in this figure). These features include: the general shape of the mandible, with expansion of the canine processes and relative length of the symphysis (seen in the dorsal outlines); the shape of the symphysis sagittal cross section; the length of the premolar row relative to the length of the molar row. The figure shows the following features for the taxa listed under each genus name: Saotherium, very inclined symphysis with thin cross-section and poorly developed canine processes; Archaeopotamus, relatively long and shallow symphysis with poorly developed canine processes and longer premolar rows than in any other clade; Hexaprotodon, wide symphysis but with poorly differentiated canine processes, very robust symphysis in cross section; Choeropsis, very short symphysis globular in cross section and poorly developed canine processes; Hippopotamus and aff. Hippopotamus, short symphysis globular in crosssection (lacking a projected incisor alveolar process) and strong extension of the canine processes the latter feature being not salient in the Afar species (aff. Hip. coryndoni, aff. Hip. afarensis) and aff. Hip. cf. protamphibius from Kanapoi." figureDoi="http://doi.org/10.5281/zenodo.5431850" httpUri="https://zenodo.org/record/5431850/files/figure.png" pageId="14" pageNumber="15">Fig. 9</figureCitation>
), reinforces this position; the association of a general thinness and of a very inclined main axis differs from the conditions seen in the other
<taxonomicName id="7AED4D69FF8BFFC6FDE53BFEA8AEFDE8" authorityName="GRAY" authorityYear="1821" class="Mammalia" family="Hippopotamidae" kingdom="Animalia" order="Artiodactyla" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Hippopotamidae</taxonomicName>
. Finally, it appears that the two Djurab Pliocene hippopotamids constitute a peculiar lineage. Following the above discussion, this lineage is separated here from all other
<taxonomicName id="7AED4D69FF8BFFC6FE4C3875AAE1FD8C" authorityName="GRAY" authorityYear="1821" box="[476,665,595,617]" class="Mammalia" family="Hippopotamidae" kingdom="Animalia" order="Artiodactyla" pageId="14" pageNumber="15" phylum="Chordata" rank="family">Hippopotamidae</taxonomicName>
at the generic level.
</paragraph>
<paragraph id="BD5236EAFF8BFFC6FF3738B6A893FB97" blockId="14.[142,759,196,1138]" pageId="14" pageNumber="15">
On the other hand, the parsimony analysis relates these Pliocene hippopotamids to the extant Liberian hippo. However, the long list of convergences accumulated by the latter taxon, its apomorphies and autapomorphies (see below) and the absence of the peculiar cranial structure of
<taxonomicName id="7AED4D69FF8BFFC6FE66390FAA04FCDB" authorityName="Boisserie" authorityYear="2005" box="[502,636,809,830]" class="Mammalia" family="Hippopotamidae" genus="Saotherium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Artiodactyla" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="8F99EAF8FF8BFFC6FE66390FAA04FCDB" box="[502,636,809,830]" italics="true" pageId="14" pageNumber="15">Saotherium</emphasis>
</taxonomicName>
obviously differentiate these animals. In fact, these taxa mainly share character states that are plesiomorphic or convergent with other taxa in the analysis, with the exception of the enlarged orbit size (character 8, state 1, see
<figureCitation id="25D62A6FFF8BFFC6FF4D39E5A964FC3D" box="[221,284,963,985]" captionStart="Figure 3" captionStartId="5.[155,233,1248,1267]" captionTargetBox="[387,780,781,1214]" captionTargetPageId="5" captionText="Figure 3. Cranial character states. A, lateral view of a Hexaprotodon liberiensis skull. B, Lateral view of a Hippopotamus amphibius skull. C, lateral view of a Hex. mingoz skull. D, three dorsal views of the braincase (from bottom to top: in Hex. harvardi, in Hex. mingoz, in Hip. amphibius). E, four schematic anterior views of the left orbit (from right to left: in Hex. protamphibius, in Hex. harvardi, in Hip. gorgops, in Hex. sivalensis)." figureDoi="http://doi.org/10.5281/zenodo.5431838" httpUri="https://zenodo.org/record/5431838/files/figure.png" pageId="14" pageNumber="15">Fig. 3</figureCitation>
and the above results). However, given the available data, it is difficult to define the most probable primitive state of this feature and hence its probable evolutionary trend. Therefore, this relationship must be carefully envisaged, but not completely ignored.
</paragraph>
<paragraph id="BD5236EAFF8BFFC6FF1E3EACA98CFAFE" blockId="14.[142,758,1162,1307]" pageId="14" pageNumber="15">
<emphasis id="8F99EAF8FF8BFFC6FF1E3EACA900FB7A" box="[142,376,1162,1183]" italics="true" pageId="14" pageNumber="15">Evolutionary trends:</emphasis>
The comparison of the Kossom Bougoudi material and the younger Kollé material led
<bibRefCitation id="D97C4B1BFF8BFFC6FF1E3EE1A9FEFB38" author="Boisserie JR &amp; Brunet M &amp; Likius A &amp; Vignaud P" box="[142,390,1223,1245]" pageId="14" pageNumber="15" pagination="15 - 27" refId="ref15268" refString="Boisserie JR, Brunet M, Likius A, Vignaud P. 2003. Hippopotamids from the Djurab Pliocene faunas, Chad, Central Africa. Journal of African Earth Sciences 36 (1 - 2): 15 - 27." type="journal article" year="2003">
Boisserie
<emphasis id="8F99EAF8FF8BFFC6FF6E3EEEA94EFB39" box="[254,310,1223,1245]" italics="true" pageId="14" pageNumber="15">et al.</emphasis>
(2003)
</bibRefCitation>
to propose some possible evolutionary trends: a relative shortening of the premolar row and a global size decrease.
</paragraph>
</subSubSection>
</treatment>
</document>