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<document id="ADDA64ADF3848607F3F2EA2CDF12E6D6" ID-DOI="10.2478/if-2019-0028" ID-ISSN="2533-4069" ID-Zenodo-Dep="5383901" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1630297739932" checkinUser="diego" docAuthor="Vianey-Liaud, Monique, Marivaux, Laurent &amp; Lehmann, Thomas" docDate="2019" docId="0394CF6EFFE5C239FF73FC754C93B595" docLanguage="en" docName="11_Vianey-Liaud.pdf" docOrigin="Fossil Imprint 75 (3 - 4)" docSource="http://dx.doi.org/10.2478/if-2019-0028" docStyle="DocumentStyle:0EC54BF35E227CA4627F894F6BA30634.4:FossilImprint.2016-.journal_article" docStyleId="0EC54BF35E227CA4627F894F6BA30634" docStyleName="FossilImprint.2016-.journal_article" docStyleVersion="4" docTitle="Masillamys TOBIEN 1954" docType="treatment" docVersion="3" lastPageNumber="471" masterDocId="FFADB716FFEAC228FF9EFF8E4F03B426" masterDocTitle="A Reevaluation Of The Taxonomic Status Of The Rodent Masillamys Tobien, 1954 From Messel (Germany, Late Early To Early Middle Eocene, 48 - 47 M. Y.)" masterLastPageNumber="483" masterPageNumber="454" pageNumber="469" updateTime="1699219439774" updateUser="plazi" zenodo-license-document="CC-BY-NC-ND-3.0">
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<mods:title id="555EB86CAFD53E264B6A03F2B3B12853">A Reevaluation Of The Taxonomic Status Of The Rodent Masillamys Tobien, 1954 From Messel (Germany, Late Early To Early Middle Eocene, 48 - 47 M. Y.)</mods:title>
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<paragraph id="8B827E78FFE5C227FF73FC754D66B035" blockId="15.[237,613,1019,1043]" box="[237,613,1019,1043]" pageId="15" pageNumber="469">
<heading id="D0CAC914FFE5C227FF73FC754D66B035" box="[237,613,1019,1043]" centered="true" fontSize="8" level="2" pageId="15" pageNumber="469" reason="2">
<emphasis id="B949A26AFFE5C227FF73FC754D66B035" bold="true" box="[237,613,1019,1043]" pageId="15" pageNumber="469">
<emphasis id="B949A26AFFE5C227FF73FC754EBFB034" bold="true" box="[237,444,1019,1043]" italics="true" pageId="15" pageNumber="469">
<taxonomicName id="4C3D05FBFFE5C227FF73FC754E68B035" authorityName="TOBIEN" authorityYear="1954" box="[237,363,1019,1043]" class="Mammalia" family="Ischyromyidae" genus="Masillamys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="15" pageNumber="469" phylum="Chordata" rank="genus">Masillamys</taxonomicName>
parvus
</emphasis>
(
<bibRefCitation id="EFAC0389FFE5C227FE52FC754D1AB037" author="Tobien, H." box="[460,537,1019,1043]" pageId="15" pageNumber="469" pagination="13 - 29" refId="ref23624" refString="Tobien, H. (1954): Nagerreste aus dem Mitteleozan von Messel bei Darmstadt. - Notizblatt des Hessischen Landesamtes fur Bodenforschung, 82: 13 - 29, 2 pls." type="journal article" year="1954">TOBIEN</bibRefCitation>
, 1954)
</emphasis>
</heading>
</paragraph>
<paragraph id="8B827E78FFE5C227FEECFB954EE3B015" blockId="15.[370,480,1051,1075]" box="[370,480,1051,1075]" pageId="15" pageNumber="469">
<figureCitation id="130662FDFFE5C227FEECFB954EE3B015" box="[370,480,1051,1075]" captionStart="Text-fig" captionStartId="16.[147,228,1499,1520]" captionTargetBox="[154,1431,188,1472]" captionTargetId="figure-219@16.[147,1462,151,1543]" captionTargetPageId="16" captionText="Text-fig. 7. Teeth of Masillamys parvus TOBIEN, from Messel (Hesse, Germany, MP 11), HLMD-Me 625, holotype. a) left upper M2, protocone damaged, occlusal view. b) left upper M3, occlusal view. c) left lower p4; c1 occlusal view, c2 buccal aspect, c3 lingual aspect. d) left lower m2, metaconid damaged; d1 occlusal view, d2 buccal aspect, d3 lingual aspect. e) right m3, occlusal view. f) left m3; f1 occlusal view, f2 buccal aspect, base of crown lacking, f3 lingual aspect; g) SMF-ME 2099A: right lower tooth row with dp4 to m3, g1 occlusal view, g2 lingual aspect. Scale bar 1 mm." figureDoi="http://doi.org/10.5281/zenodo.5383915" httpUri="https://zenodo.org/record/5383915/files/figure.png" pageId="15" pageNumber="469">Text-fig. 7</figureCitation>
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<paragraph id="8B827E78FFE5C227FF0EFBC54E20B085" blockId="15.[106,745,1098,1187]" pageId="15" pageNumber="469">H o l o t y p e. HLMD-Me 625, poorly preserved skeleton with extracted teeth: left upper M1M3 and lower p4m3, right m3;</paragraph>
<paragraph id="8B827E78FFE5C227FF0EFB354DEAB0F4" blockId="15.[106,745,1210,1267]" box="[144,745,1210,1235]" pageId="15" pageNumber="469">
Ty p e l o c a l i t y. Messel Fossil Pit (
<collectingCountry id="F32A3EE8FFE5C227FDEAFB344DDDB0F4" box="[628,734,1210,1234]" name="Germany" pageId="15" pageNumber="469">Germany</collectingCountry>
).
</paragraph>
<paragraph id="8B827E78FFE5C227FFF4FB544DDEB0D5" blockId="15.[106,745,1210,1267]" box="[106,733,1242,1267]" pageId="15" pageNumber="469">
<materialsCitation id="3B557425FFE5C227FFF4FB544DC8B0D5" box="[106,715,1242,1267]" collectionCode="MP" elevation="48" pageId="15" pageNumber="469" specimenCode="MP 11" specimenCount="48">
Transition late Ypresian-early Lutetian,
<specimenCode id="DB9BD603FFE5C227FD8CFB554D5AB0D4" box="[530,601,1242,1266]" collectionCode="MP" country="0" name="Mohonk Preserve, Inc." pageId="15" pageNumber="469">MP 11</specimenCode>
, 47
<specimenCount id="9D3BB5F1FFE5C227FD10FB544DC8B0D5" box="[654,715,1242,1267]" pageId="15" pageNumber="469" type="generic">
<quantity id="4CC5D39DFFE5C227FD10FB544DC4B0D5" box="[654,711,1242,1267]" metricMagnitude="1" metricUnit="m" metricValue="4.8" pageId="15" pageNumber="469" unit="m" value="48.0">
<elevation id="0010994BFFE5C227FD10FB544DC4B0D5" box="[654,711,1242,1267]" metricMagnitude="1" metricUnit="m" metricValue="4.8" pageId="15" pageNumber="469" unit="m" value="48.0">48 m</elevation>
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.
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y.
</paragraph>
<paragraph id="8B827E78FFE5C227FF0EFA844DCBB245" blockId="15.[106,745,1289,1635]" pageId="15" pageNumber="469">
O r i g i n a l d i a g n o s i s. Definitely smaller than
<emphasis id="B949A26AFFE5C227FD52FA844FB9B167" italics="true" pageId="15" pageNumber="469">M. beegeri</emphasis>
and
<emphasis id="B949A26AFFE5C227FF72FAA44E44B167" box="[236,327,1321,1345]" italics="true" pageId="15" pageNumber="469">M. krugi</emphasis>
. On lower molars, the mesial cingulum is separated from the protoconid by a deep incision. The only remain of the metalophid is a posterior arm of the protoconid, whereas a lingual metalophid is missing. The hypolophid [= entolophid] is more sharply defined than in the other two species, proceeding from the entoconid, and joining the ectolophid between the mesoconid and the hypoconid. Entoconid opposite to the hypoconid on the p4, whereas it is distinctly more mesial on the lower molars. (
<bibRefCitation id="EFAC0389FFE5C227FFEFF9C54E4FB245" author="Tobien, H." box="[113,332,1611,1635]" pageId="15" pageNumber="469" pagination="13 - 29" refId="ref23624" refString="Tobien, H. (1954): Nagerreste aus dem Mitteleozan von Messel bei Darmstadt. - Notizblatt des Hessischen Landesamtes fur Bodenforschung, 82: 13 - 29, 2 pls." type="journal article" year="1954">Tobien 1954: 2324</bibRefCitation>
, translation adapted from German).
</paragraph>
<paragraph id="8B827E78FFE5C227FF0EF9F44EDDBC12" blockId="15.[106,746,1658,2100]" pageId="15" pageNumber="469">
E m e n d e d d i a g n o s i s. Relatively small sized rodent (
<typeStatus id="5486C0DAFFE5C227FFEFF9144FD1B294" box="[113,210,1690,1714]" pageId="15" pageNumber="469" type="holotype">holotype</typeStatus>
: p4 = 1.75 × 1.42; m2 = 1.93 × 1.84; m3 = 2.02 × 1.75), slighly larger than
<emphasis id="B949A26AFFE5C227FE0DF9344DE1B2F4" box="[403,738,1722,1746]" italics="true" pageId="15" pageNumber="469">
<taxonomicName id="4C3D05FBFFE5C227FE0DF9344D55B2F4" baseAuthorityName="Escarguel" baseAuthorityYear="1999" box="[403,598,1722,1746]" class="Mammalia" family="Ischyromyidae" genus="Hartenbergeromys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="15" pageNumber="469" phylum="Chordata" rank="genus">Hartenbergeromys</taxonomicName>
hautefeuillei
</emphasis>
. On lower molars, lingual main cuspids more mesial than buccal ones; p4 with protruding metaconid, mediomesial, and no protoconid; posterior lobe of m3 not strongly reduced. On molars, lingual metalophulid I incomplete, buccal metalophulid I ending at midwidth; entolophid complete, attached to the distal ectolophid/prehypocristid junction; ectomesolophid present and hypoconulid reduced. On upper M2 and M3, mesostyle present and positioned only slightly more buccally than the paracone and metacone; hypocone well-developed, with strong pre- and post-hypocristae; shallow sinus present on both teeth.
</paragraph>
<paragraph id="8B827E78FFE5C227FCC4FF164B08B4F6" blockId="15.[820,1459,151,499]" pageId="15" pageNumber="469">
D i f f e r e n t i a l d i a g n o s i s.Thespecies
<emphasis id="B949A26AFFE5C227FAA6FF194C7EB4F6" italics="true" pageId="15" pageNumber="469">
<taxonomicName id="4C3D05FBFFE5C227FAA6FF194AB0B489" authorityName="TOBIEN" authorityYear="1954" box="[1336,1459,151,175]" class="Mammalia" family="Ischyromyidae" genus="Masillamys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="15" pageNumber="469" phylum="Chordata" rank="genus">Masillamys</taxonomicName>
parvus
</emphasis>
differs from:
</paragraph>
<paragraph id="8B827E78FFE5C227FCAAFF574B0CB577" blockId="15.[820,1459,151,499]" pageId="15" pageNumber="469">
<emphasis id="B949A26AFFE5C227FCC4FF574CCCB4D6" box="[858,975,216,240]" italics="true" pageId="15" pageNumber="469">M. beegeri</emphasis>
and
<emphasis id="B949A26AFFE5C227FB9BFF574B62B4D6" box="[1029,1121,216,240]" italics="true" pageId="15" pageNumber="469">M. krugi</emphasis>
in its smaller size; the absence of mesiobuccal cingulid and the reduction of the protocristid on p4; the absence of lingual metalophulid on lower molars.
</paragraph>
<paragraph id="8B827E78FFE5C227FCAAFED44A8BB5D5" blockId="15.[820,1459,151,499]" pageId="15" pageNumber="469">
<emphasis id="B949A26AFFE5C227FCC4FED54B02B554" box="[858,1025,346,370]" italics="true" pageId="15" pageNumber="469">H. hautefeuillei</emphasis>
in the absence of mesiobuccal cingulid, the reduction of the protocristid and the metaconid lower relatively to the entoconid on p4; the entolophid more regularly continuous on molars, the less reduced hypocone relative to the protocone on upper molars.
</paragraph>
<paragraph id="8B827E78FFE5C227FCC4FD824A54B6A2" blockId="15.[820,1460,523,774]" pageId="15" pageNumber="469">
R e f e r r e d m a t e r i a l.
<emphasis id="B949A26AFFE5C227FB1EFD854B23B665" bold="true" pageId="15" pageNumber="469">Hessisches Landesmuseum Darmstadt (HLMD).</emphasis>
<emphasis id="B949A26AFFE5C227FBBAFDA54BEEB665" box="[1060,1261,555,579]" italics="true" pageId="15" pageNumber="469">
<taxonomicName id="4C3D05FBFFE5C227FBBAFDA54B9CB665" authorityName="TOBIEN" authorityYear="1954" box="[1060,1183,555,579]" class="Mammalia" family="Ischyromyidae" genus="Masillamys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="15" pageNumber="469" phylum="Chordata" rank="genus">Masillamys</taxonomicName>
parvus
</emphasis>
<typeStatus id="5486C0DAFFE5C227FB6CFDA54A57B665" box="[1266,1364,555,579]" pageId="15" pageNumber="469" type="holotype">holotype</typeStatus>
: HLMD- Me 625: a poorly preserved skeleton with teeth extracted: left upper M1M3 and lower p4m3, and right m3.
</paragraph>
<paragraph id="8B827E78FFE5C227FCC4FD024AB0B720" blockId="15.[820,1460,523,774]" pageId="15" pageNumber="469">
<emphasis id="B949A26AFFE5C227FCC4FD024B6EB6E3" bold="true" pageId="15" pageNumber="469">Senckenberg Research Institute and Natural History Museum Frankfurt (SMF).</emphasis>
<taxonomicName id="4C3D05FBFFE5C227FBEBFD234BF3B6E3" authorityName="TOBIEN" authorityYear="1954" box="[1141,1264,685,709]" class="Mammalia" family="Ischyromyidae" genus="Masillamys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="15" pageNumber="469" phylum="Chordata" rank="genus">
<emphasis id="B949A26AFFE5C227FBEBFD234BF3B6E3" box="[1141,1264,685,709]" italics="true" pageId="15" pageNumber="469">Masillamys</emphasis>
</taxonomicName>
?
<emphasis id="B949A26AFFE5C227FA9BFD204A4DB6E3" box="[1285,1358,686,709]" italics="true" pageId="15" pageNumber="469">parvus</emphasis>
juvenile: SMF-ME 2099: a skull badly crushed; a left row of lower teeth exposed in nearly occlusal view (dp4, m1, m2 and m3).
</paragraph>
<paragraph id="8B827E78FFE5C227FCC4FC904CDFB03E" blockId="15.[820,1459,797,1081]" pageId="15" pageNumber="469">
M e a s u r e m e n t s. The only known skeleton is that of the
<typeStatus id="5486C0DAFFE5C227FC22FCB04B1AB770" box="[956,1049,830,854]" pageId="15" pageNumber="469" type="holotype">holotype</typeStatus>
(HLMD-Me 625), which is poorly preserved. Almost all its articulations are crushed, distorted or unrecognizable. Nonetheless, its estimated body weight of
<quantity id="4CC5D39DFFE5C227FCC8FC114CB2B791" box="[854,945,927,951]" metricMagnitude="-2" metricUnit="kg" metricValue="5.75" metricValueMax="6.0" metricValueMin="5.5" pageId="15" pageNumber="469" unit="g" value="57.5" valueMax="60.0" valueMin="55.0">5560 g</quantity>
(
<bibRefCitation id="EFAC0389FFE5C227FC5DFC114B71B791" author="Escarguel, G." box="[963,1138,927,951]" pageId="15" pageNumber="469" pagination="89 - 351" refId="ref22759" refString="Escarguel, G. (1999): Les rongeurs de l'Eocene inferieur et moyen d'Europe Occidentale. Systematique, phylogenie, biochronologie et paleobiogeographie des niveaux- - reperes MP 7 a MP 14. - Palaeovertebrata, 28: 89 - 351." type="journal article" year="1999">Escarguel 1999</bibRefCitation>
), and estimated trunk length (measured along the cervical-thoracic-lumbar spine) of less than
<quantity id="4CC5D39DFFE5C227FCF6FC6E4CB2B7D1" box="[872,945,992,1016]" metricMagnitude="-2" metricUnit="m" metricValue="8.0" pageId="15" pageNumber="469" unit="mm" value="80.0">80 mm</quantity>
makes
<emphasis id="B949A26AFFE5C227FB9CFC6F4B69B7DE" box="[1026,1130,993,1016]" italics="true" pageId="15" pageNumber="469">H. parvus</emphasis>
a smaller animal than
<emphasis id="B949A26AFFE5C227FAC6FC6F4AB0B7DE" box="[1368,1459,992,1016]" italics="true" pageId="15" pageNumber="469">M. krugi</emphasis>
and
<emphasis id="B949A26AFFE5C227FCFFFB8F4CD6B03E" box="[865,981,1024,1048]" italics="true" pageId="15" pageNumber="469">M. beegeri</emphasis>
.
</paragraph>
<paragraph id="8B827E78FFE5C227FCC4FBAF4A70B01F" blockId="15.[820,1459,797,1081]" box="[858,1395,1057,1081]" pageId="15" pageNumber="469">
For teeth measurements see
<tableCitation id="C6BF4BC3FFE5C227FB16FBAF4BC9B01F" box="[1160,1226,1057,1081]" captionStart="Table 1" captionStartId="7.[106,162,152,173]" captionText="Table 1. List of material and measurements (L = length, W = width; in mm) of Masillamys mattaueri from Mas de Gimel (Hérault, France) (MGL) stored in the Instiut des Sciences de lEvolution, Montpellier University (ISE-M), and Masillamys beegeri, M. krugi and M. parvus from Messel (Hesse, Germany), stored in the Senckenberg Research Institute and Natural History Museum Frankfurt (SMF) and in Hessisches Landesmuseum Darmstadt (HLMD)." pageId="15" pageNumber="469">Tab. 1</tableCitation>
and
<figureCitation id="130662FDFFE5C227FB63FBAF4A70B01F" box="[1277,1395,1057,1081]" captionStart="Text-fig" captionStartId="6.[129,210,1146,1167]" captionTargetBox="[142,745,194,1120]" captionTargetId="figure-834@6.[130,761,189,1181]" captionTargetPageId="6" captionText="Text-fig. 4. Bivariate graph (length/width) of the teeth of the three species of Masillamys from Messel." figureDoi="http://doi.org/10.5281/zenodo.5383909" httpUri="https://zenodo.org/record/5383909/files/figure.png" pageId="15" pageNumber="469">Text-fig. 4.</figureCitation>
</paragraph>
<paragraph id="8B827E78FFE5C227FCC4FBDF4A78B04E" blockId="15.[820,1459,1104,2100]" box="[858,1403,1104,1128]" pageId="15" pageNumber="469">
D e s c r i p t i o n.
<emphasis id="B949A26AFFE5C227FB80FBDE4BAFB04E" bold="true" box="[1054,1196,1104,1128]" pageId="15" pageNumber="469">Upper teeth.</emphasis>
See
<figureCitation id="130662FDFFE5C227FB41FBDE4A78B04E" box="[1247,1403,1104,1128]" captionStart="Text-fig" captionStartId="16.[147,228,1499,1520]" captionTargetBox="[154,1431,188,1472]" captionTargetId="figure-219@16.[147,1462,151,1543]" captionTargetPageId="16" captionText="Text-fig. 7. Teeth of Masillamys parvus TOBIEN, from Messel (Hesse, Germany, MP 11), HLMD-Me 625, holotype. a) left upper M2, protocone damaged, occlusal view. b) left upper M3, occlusal view. c) left lower p4; c1 occlusal view, c2 buccal aspect, c3 lingual aspect. d) left lower m2, metaconid damaged; d1 occlusal view, d2 buccal aspect, d3 lingual aspect. e) right m3, occlusal view. f) left m3; f1 occlusal view, f2 buccal aspect, base of crown lacking, f3 lingual aspect; g) SMF-ME 2099A: right lower tooth row with dp4 to m3, g1 occlusal view, g2 lingual aspect. Scale bar 1 mm." figureDoi="http://doi.org/10.5281/zenodo.5383915" httpUri="https://zenodo.org/record/5383915/files/figure.png" pageId="15" pageNumber="469">Text-fig. 7a, b.</figureCitation>
</paragraph>
<paragraph id="8B827E78FFE5C227FCC4FBFE4C71B10C" blockId="15.[820,1459,1104,2100]" pageId="15" pageNumber="469">
<emphasis id="B949A26AFFE5C227FCC4FBFE4CB8B0AE" bold="true" box="[858,955,1136,1160]" pageId="15" pageNumber="469">
M
<figureCitation id="130662FDFFE5C227FCEAFBFE4CAAB0AE" box="[884,937,1136,1160]" captionStart="Text-fig" captionStartId="3.[125,204,1381,1402]" captionTargetBox="[125,1440,151,1361]" captionTargetId="figure-340@3.[125,1440,151,1361]" captionTargetPageId="3" captionText="Text-fig. 2. Species of Masillamys considered on the phylogenetic tree of theridomorphs (Vianey-Liaud and Marivaux 2017: fig. 7), within the basal Theridomorpha, before the polyphyletic genus Protadelomys. Position inferred from their dental features (see text)." figureDoi="http://doi.org/10.5281/zenodo.5383905" httpUri="https://zenodo.org/record/5383905/files/figure.png" pageId="15" pageNumber="469">2M</figureCitation>
<figureCitation id="130662FDFFE5C227FC37FBFE4CB5B0AE" box="[937,950,1136,1160]" captionStart="Text-fig" captionStartId="4.[129,210,898,919]" captionTargetBox="[135,762,152,879]" captionTargetId="figure-848@4.[133,763,151,879]" captionTargetPageId="4" captionText="Text-fig. 3. Front views of hemi-maxillaries from Prémontré (Paris Basin, MP 10), showing the large infraorbitary foramen ofa) SLP29PR-1312,storedintheMontpellierUniversitycollections (ISE-M), Hartenbergeromys hautefeuillei; b) SLP29PR-960, Pantrogna marandati. Scale bar 1 mm." figureDoi="http://doi.org/10.5281/zenodo.5383907" httpUri="https://zenodo.org/record/5383907/files/figure.png" pageId="15" pageNumber="469">3</figureCitation>
.
</emphasis>
Two fragmentary left teeth have been described and illustrated by
<bibRefCitation id="EFAC0389FFE5C227FB98FB1F4BA4B08F" author="Tobien, H." box="[1030,1191,1169,1193]" pageId="15" pageNumber="469" pagination="13 - 29" refId="ref23624" refString="Tobien, H. (1954): Nagerreste aus dem Mitteleozan von Messel bei Darmstadt. - Notizblatt des Hessischen Landesamtes fur Bodenforschung, 82: 13 - 29, 2 pls." type="journal article" year="1954">Tobien (1954)</bibRefCitation>
. The left M2 lacks the protocone and its anterior arm plus a part of the anteroloph, and the posterobuccal corner of the crown. The left M3 is less incomplete, lacking only the posterolingual edge of the tooth.
</paragraph>
<paragraph id="8B827E78FFE5C227FCC4FABD4A2FB3D5" blockId="15.[820,1459,1104,2100]" pageId="15" pageNumber="469">On M2, the buccal part of the anteroloph lacks a parastyle. The anteroloph is regular, thin and low, leaving a well-developed but narrow anterosyncline. The paracone and metacone have equal size. The postparacrista and premetacrista are equally developed, both attached to the mesostyle. The latter is low and weakly stretched mesiodistally; it is connected to the mesoloph, which occupies alone the bottom of the small mesosyncline. The buccal part of the protoloph is straight and, after the protruding paraconule, it joins the protocone. From the buccal part of the protoloph, there is one extra-ridge running mesially into the anterosyncline, and three very low extra-ridges descending to the centre of the basin. One linguobuccal extra-ridge starts from the lingual protoloph to the basin. The buccal metalophule II is weak and thin, turning mesially to join the metaconule, which is low and stronger than the paraconule. It is also linked to the posteroloph. Two small low extra-ridges are present along the mesial base of the metaconule. The hypocone is small and the endoloph short; below, the sinus is not well defined. A posterostyle marks the posthypocrista/ posteroloph junction. The posteroloph is higher than the anteroloph, ending at the base of the metacone.</paragraph>
<paragraph id="8B827E78FFE5C238FCC4F8724B25B36C" blockId="15.[820,1459,1104,2100]" lastBlockId="16.[843,1482,1708,2100]" lastPageId="16" lastPageNumber="470" pageId="15" pageNumber="469">On the weakly worn M3, the parastyle appears slightly swollen at the lingual end of the anteroloph, which is more worn. The mesial end of the preprotocrista is swollen at the place of the anterostyle. The two arms of the protocone make a wide open angle. The buccal half of the protoloph is high and plunging to the level of the low extra-ridges. It bears a stretched and protruding paraconule, and then continues to a swelling before being weakly attached to the thick protocrista. Three extra-ridges are present on the distal slope of the buccal protoloph, parallel to the postparacrista. Another ridge runs from the level of the postparacrista to the basin. The mesostyle is low and stretched mesiodistally; it is prolonged in one short, low and thick mesoloph. The metaconule is present as a low bulged ridge facing the endoloph. The hypocone is reduced; distally, the area between the posthypocrista + the posteroloph + the metacone form a cingulum elevated above the basin. Its posterior edge being broken, precise relationships between these structures cannot be described.</paragraph>
<caption id="DF422EF0FFFAC238FF0DFA554B0CB243" ID-DOI="http://doi.org/10.5281/zenodo.5383915" ID-Zenodo-Dep="5383915" httpUri="https://zenodo.org/record/5383915/files/figure.png" pageId="16" pageNumber="470" startId="16.[147,228,1499,1520]" targetBox="[154,1431,188,1472]" targetPageId="16">
<paragraph id="8B827E78FFFAC238FF0DFA554B0CB243" blockId="16.[147,1463,1499,1637]" pageId="16" pageNumber="470">
<emphasis id="B949A26AFFFAC238FF0DFA554D71B1C9" bold="true" box="[147,626,1499,1520]" pageId="16" pageNumber="470">
Text-fig. 7. Teeth of
<emphasis id="B949A26AFFFAC238FEF6FA554D27B1D6" bold="true" box="[360,548,1499,1520]" italics="true" pageId="16" pageNumber="470">
<taxonomicName id="4C3D05FBFFFAC238FEF6FA554EDAB1D6" authorityName="TOBIEN" authorityYear="1954" box="[360,473,1499,1520]" class="Mammalia" family="Ischyromyidae" genus="Masillamys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="16" pageNumber="470" phylum="Chordata" rank="genus">Masillamys</taxonomicName>
parvus
</emphasis>
TOBIEN
</emphasis>
, from Messel (Hesse, Germany, MP 11), HLMD-Me 625, holotype. a) left upper M2, protocone damaged, occlusal view. b) left upper M3, occlusal view. c) left lower p4; c1 occlusal view, c2 buccal aspect, c3 lingual aspect. d) left lower m2, metaconid damaged; d1 occlusal view, d2 buccal aspect, d3 lingual aspect. e) right m3, occlusal view. f) left m3; f1 occlusal view, f2 buccal aspect, base of crown lacking, f3 lingual aspect; g) SMF-ME 2099A: right lower tooth row with dp4 to m3, g1 occlusal view, g2 lingual aspect. Scale bar 1 mm.
</paragraph>
</caption>
<paragraph id="8B827E78FFFAC238FCEFF8DD4BE2B388" blockId="16.[843,1482,1708,2100]" pageId="16" pageNumber="470">
<emphasis id="B949A26AFFFAC238FCEFF8DD4B08B34D" bold="true" box="[881,1035,1875,1899]" pageId="16" pageNumber="470">Lower teeth.</emphasis>
See
<figureCitation id="130662FDFFFAC238FBCBF8DD4BFAB34A" box="[1109,1273,1875,1900]" captionStart="Text-fig" captionStartId="16.[147,228,1499,1520]" captionTargetBox="[154,1431,188,1472]" captionTargetId="figure-219@16.[147,1462,151,1543]" captionTargetPageId="16" captionText="Text-fig. 7. Teeth of Masillamys parvus TOBIEN, from Messel (Hesse, Germany, MP 11), HLMD-Me 625, holotype. a) left upper M2, protocone damaged, occlusal view. b) left upper M3, occlusal view. c) left lower p4; c1 occlusal view, c2 buccal aspect, c3 lingual aspect. d) left lower m2, metaconid damaged; d1 occlusal view, d2 buccal aspect, d3 lingual aspect. e) right m3, occlusal view. f) left m3; f1 occlusal view, f2 buccal aspect, base of crown lacking, f3 lingual aspect; g) SMF-ME 2099A: right lower tooth row with dp4 to m3, g1 occlusal view, g2 lingual aspect. Scale bar 1 mm." figureDoi="http://doi.org/10.5281/zenodo.5383915" httpUri="https://zenodo.org/record/5383915/files/figure.png" pageId="16" pageNumber="470">Text-fig. 7cg</figureCitation>
. The crowns are relatively low and basined, with moderately prominent main cuspids, showing acute lingual ridges.
</paragraph>
<paragraph id="8B827E78FFFAC239FCEFF8394DBDB4F6" blockId="16.[843,1482,1708,2100]" lastBlockId="17.[106,746,151,2100]" lastPageId="17" lastPageNumber="471" pageId="16" pageNumber="470">
<emphasis id="B949A26AFFFAC238FCEFF8394CA1B3E9" bold="true" box="[881,930,1975,1999]" pageId="16" pageNumber="470">dp4.</emphasis>
SMF-ME 2099: As it is heavily worn, extraridges are not visible. The metaconid is the unique mesial cuspid, from which a faint mesiodistal ridge descends. The protoconid ridge is low and worn. At the opposite, the lingual postmetacristid is long and high. The entoconid is isolated from the posterolophid, and the entolophid seems absent.
</paragraph>
<paragraph id="8B827E78FFFBC239FF0EFF594D59B6B7" blockId="17.[106,746,151,2100]" pageId="17" pageNumber="471">
<emphasis id="B949A26AFFFBC239FF0EFF594FB1B4C9" bold="true" box="[144,178,215,239]" pageId="17" pageNumber="471">p4.</emphasis>
HLMD-Me 625: This tooth is quite smaller than m1. The metaconid, mesiomedian, is the highest and strongest cuspid. Its posterior arm descends gently towards the mesosynclinid lingual opening. The tooth is slightly damaged, but it is possible to see one main low mesiodistal ridge descending from the apex of the metaconid to the centre of the talonid basin. The postmetacristid is gently sploping. The protoconid is absent and the protocristid ridge hardly visible. A swelling indicates the presence of a mesoconid. There is a break between a spur at the junction between the ectolophid anterior arm of the hypoconid and the thin and continuous entolophid. The posthypocristid makes an angle with the long posterolophid, which joins the distal slope of the entoconid. The hypoconulid is not distinct.
</paragraph>
<paragraph id="8B827E78FFFBC239FF0EFD174D97B7D4" blockId="17.[106,746,151,2100]" pageId="17" pageNumber="471">
<emphasis id="B949A26AFFFBC239FF0EFD174FE8B697" bold="true" box="[144,235,665,689]" pageId="17" pageNumber="471">m2m3.</emphasis>
HLMD-Me 625: Only one fragmentary m1, one slightly damaged m2, and the two m3s are known. On the well-preserved surfaces of teeth, the enamel of the crown outline is rough. Like in
<emphasis id="B949A26AFFFBC239FE1FFD744EDCB737" box="[385,479,761,785]" italics="true" pageId="17" pageNumber="471">M. krugi</emphasis>
, the lingual cuspids are less high than the buccal ones. But there is neither an isolated mesostylid nor a wide lingual opening for the mesosynclinid in front of the entoconid. On m2, the postmetacristid descends first gently, then more abruptly at the end, from where starts the relatively long mesolophulid (
<figureCitation id="130662FDFFFBC239FDC5FC174DDEB797" box="[603,733,921,945]" captionStart="Text-fig" captionStartId="16.[147,228,1499,1520]" captionTargetBox="[154,1431,188,1472]" captionTargetId="figure-219@16.[147,1462,151,1543]" captionTargetPageId="16" captionText="Text-fig. 7. Teeth of Masillamys parvus TOBIEN, from Messel (Hesse, Germany, MP 11), HLMD-Me 625, holotype. a) left upper M2, protocone damaged, occlusal view. b) left upper M3, occlusal view. c) left lower p4; c1 occlusal view, c2 buccal aspect, c3 lingual aspect. d) left lower m2, metaconid damaged; d1 occlusal view, d2 buccal aspect, d3 lingual aspect. e) right m3, occlusal view. f) left m3; f1 occlusal view, f2 buccal aspect, base of crown lacking, f3 lingual aspect; g) SMF-ME 2099A: right lower tooth row with dp4 to m3, g1 occlusal view, g2 lingual aspect. Scale bar 1 mm." figureDoi="http://doi.org/10.5281/zenodo.5383915" httpUri="https://zenodo.org/record/5383915/files/figure.png" pageId="17" pageNumber="471">Text-fig. 7d</figureCitation>
). On m3, the end of the postmetacristid makes a kind of flat, from which the mesolophulid starts (
<figureCitation id="130662FDFFFBC239FE6FFC544D84B7D4" box="[497,647,986,1010]" captionStart="Text-fig" captionStartId="16.[147,228,1499,1520]" captionTargetBox="[154,1431,188,1472]" captionTargetId="figure-219@16.[147,1462,151,1543]" captionTargetPageId="16" captionText="Text-fig. 7. Teeth of Masillamys parvus TOBIEN, from Messel (Hesse, Germany, MP 11), HLMD-Me 625, holotype. a) left upper M2, protocone damaged, occlusal view. b) left upper M3, occlusal view. c) left lower p4; c1 occlusal view, c2 buccal aspect, c3 lingual aspect. d) left lower m2, metaconid damaged; d1 occlusal view, d2 buccal aspect, d3 lingual aspect. e) right m3, occlusal view. f) left m3; f1 occlusal view, f2 buccal aspect, base of crown lacking, f3 lingual aspect; g) SMF-ME 2099A: right lower tooth row with dp4 to m3, g1 occlusal view, g2 lingual aspect. Scale bar 1 mm." figureDoi="http://doi.org/10.5281/zenodo.5383915" httpUri="https://zenodo.org/record/5383915/files/figure.png" pageId="17" pageNumber="471">Text-fig. 7e, f</figureCitation>
).
</paragraph>
<paragraph id="8B827E78FFFBC239FF0EFC744D07B255" blockId="17.[106,746,151,2100]" pageId="17" pageNumber="471">The anterolophid is well-preserved only on the m3: it is slender, without an anteroconid and it ends lingually at a short mesial and linguobuccal premetacristid. It does not fuse with the protoconid, leaving a narrow buccal opening of the anteroflexid. The anterolophid is higher than the other ridges, which run towards the anteroflexid and mesoflexid. From the metaconid and anterolophid, short and low extraridges descend on the centre of the tooth. Starting from the protoconid, the short buccal metalophulid is the only component of the metalophid being well developed. The thick and oblique postprotocristid bears a premesoconid spur, before the shrinkage of the mesial ectolophid. The mesoconid is bulged (horizontally worn like the protocone and hypocone) and buccally bears the ectomesolophid, as well as lingually a long mesolophid, nearly merged with the mesolophulid. The angled entolophid connects to the distal ectolophid, where it thickens (postmesoconid swelling?). There are two extra-ridges in the anterosynclinid and a few along the lingual slope of the metaconid to the basin and some granules in the posterosynclinid.</paragraph>
<paragraph id="8B827E78FFFBC239FF0EF9F54C93B595" blockId="17.[106,746,151,2100]" lastBlockId="17.[820,1459,152,435]" pageId="17" pageNumber="471">
<emphasis id="B949A26AFFFBC239FF0EF9F54E1FB2B5" bold="true" box="[144,284,1659,1683]" pageId="17" pageNumber="471">m1m2m3.</emphasis>
SMF-ME 2099 (
<figureCitation id="130662FDFFFBC239FE6DF9F54D6CB2B5" box="[499,623,1659,1683]" captionStart="Text-fig" captionStartId="16.[147,228,1499,1520]" captionTargetBox="[154,1431,188,1472]" captionTargetId="figure-219@16.[147,1462,151,1543]" captionTargetPageId="16" captionText="Text-fig. 7. Teeth of Masillamys parvus TOBIEN, from Messel (Hesse, Germany, MP 11), HLMD-Me 625, holotype. a) left upper M2, protocone damaged, occlusal view. b) left upper M3, occlusal view. c) left lower p4; c1 occlusal view, c2 buccal aspect, c3 lingual aspect. d) left lower m2, metaconid damaged; d1 occlusal view, d2 buccal aspect, d3 lingual aspect. e) right m3, occlusal view. f) left m3; f1 occlusal view, f2 buccal aspect, base of crown lacking, f3 lingual aspect; g) SMF-ME 2099A: right lower tooth row with dp4 to m3, g1 occlusal view, g2 lingual aspect. Scale bar 1 mm." figureDoi="http://doi.org/10.5281/zenodo.5383915" httpUri="https://zenodo.org/record/5383915/files/figure.png" pageId="17" pageNumber="471">
Text-fig.
<quantity id="4CC5D39DFFFBC239FDC8F9F54D6CB2B5" box="[598,623,1659,1683]" metricMagnitude="-3" metricUnit="kg" metricValue="7.0" pageId="17" pageNumber="471" unit="g" value="7.0">7g</quantity>
</figureCitation>
): Like on the
<typeStatus id="5486C0DAFFFBC239FF06F9154FFBB295" box="[152,248,1691,1715]" pageId="17" pageNumber="471" type="holotype">holotype</typeStatus>
, the lingual main cusps are more mesially positioned than the buccal ones, but only slighly more than in
<emphasis id="B949A26AFFFBC239FF12F9524E04B2D5" box="[140,263,1755,1779]" italics="true" pageId="17" pageNumber="471">M. beegeri</emphasis>
. The complete anterolophid is transverse, aligned with the plunging mesial premetacristid, without an individualized anteroconid, as on the type specimen; as the teeth are nearly unworn, the buccal end of the anterosynclinid remains open. This feature was argued to differentiate
<emphasis id="B949A26AFFFBC239FD52F8D34FB0B3B2" italics="true" pageId="17" pageNumber="471">M. parvus</emphasis>
from
<emphasis id="B949A26AFFFBC239FF6DF8F34E64B3B2" box="[243,359,1916,1940]" italics="true" pageId="17" pageNumber="471">M. beegeri</emphasis>
and
<emphasis id="B949A26AFFFBC239FE04F8F34EF6B3B2" box="[410,501,1916,1940]" italics="true" pageId="17" pageNumber="471">M. krugi</emphasis>
but this opening is not deep. It is found also on the m1 of
<emphasis id="B949A26AFFFBC239FE7DF8134D3CB392" box="[483,575,1948,1972]" italics="true" pageId="17" pageNumber="471">M. krugi</emphasis>
, less worn than the m2, and on the m3 of
<emphasis id="B949A26AFFFBC239FEE4F8334EEEB3F2" box="[378,493,1980,2004]" italics="true" pageId="17" pageNumber="471">M. beegeri</emphasis>
, less worn than m2 and m1. This feature cannot be used in a differential diagnosis. The lingual metalophulid is stronger than on the type, and weakly joins the buccal metalophulid I on m1 and m2. The postmetacristid has the same organization as on the type with a narrow lingual opening of the mesosynclinid, but it ends into a shorter low lingual mesolophulid. Like on the type, the entolophid is complete, attached to the distal ectolophid junction with the prehypocristid. The ectomesolophid is present and the hypoconulid is reduced. The extra-ridges are better marked (but the teeth are less worn). Like for the type, the width of the posterior lobe of the m3 is only slightly reduced.
</paragraph>
</subSubSection>
</treatment>
</document>