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<taxonomicName id="4C4CDAFCFFD3FF8F2A9F04C8FD1AFB0E" authority="(VON MEYER, 1848)" baseAuthorityName="VON MEYER" baseAuthorityYear="1848" box="[192,658,1188,1213]" class="Mammalia" family="Sciuridae" genus="Spermophilinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="2" pageNumber="176" phylum="Chordata" rank="species" species="bredai">
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<emphasis id="B9387D6DFFD3FF8F2A9F04C8FE33FB0E" bold="true" box="[192,443,1188,1212]" italics="true" pageId="2" pageNumber="176">Spermophilinus bredai</emphasis>
(VON MEYER, 1848)
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<paragraph id="8BF3A17FFFD3FF8F2B3C04A9FE67FB6F" blockId="2.[355,495,1221,1245]" box="[355,495,1221,1245]" pageId="2" pageNumber="176">
<figureCitation id="1377BDFAFFD3FF8F2B3C04A9FE5DFB6F" box="[355,469,1221,1245]" captionStart="Text-fig" captionStartId="1.[147,228,2019,2040]" captionTargetBox="[149,1460,969,1981]" captionTargetId="figure-312@1.[146,900,966,1613]" captionTargetPageId="1" captionText="Text-fig. 1. Spermophilinus bredai (VON MEYER, 1848): 3D models of a right upper jaw with (abnormal) P3, P4 and all molars (mirrored; NMA-2019-1/2352). a – the specimen in occlusal view showing the relative position of the three roots in place of P3 (yellow) compared to P4 (green); b – same view, with focus on P3–P4; c – lingual view; d – mesial view. Scale bars 2mm." figureDoi="http://doi.org/10.5281/zenodo.5383455" httpUri="https://zenodo.org/record/5383455/files/figure.png" pageId="2" pageNumber="176">Text-figs 1</figureCitation>
, 
<figureCitation id="1377BDFAFFD3FF8F2BBD04A9FE67FB6F" box="[482,495,1221,1245]" captionStart="Text-fig" captionStartId="2.[125,206,1075,1096]" captionTargetBox="[125,1440,151,1053]" captionTargetId="figure-579@2.[781,1441,150,786]" captionTargetPageId="2" captionText="Text-fig. 2. Spermophilinus bredai (VON MEYER, 1848): left upper jaw with P3–M1 (NMA-2019-2/2352). a – the specimen in occlusal view; b – mesial view; c – labial view. Scale bars 1mm." figureDoi="http://doi.org/10.5281/zenodo.5383457" httpUri="https://zenodo.org/record/5383457/files/figure.png" pageId="2" pageNumber="176">2</figureCitation>
</paragraph>
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M a t e r i a l a n d m e a s u r e m e n t s. Right upper jaw with (abnormal) P3, P4 and all molars (NMA-2019- 1/2352; 
<figureCitation id="1377BDFAFFD3FF8F2A9E0559FEA6FAFC" box="[193,302,1333,1358]" captionStart="Text-fig" captionStartId="1.[147,228,2019,2040]" captionTargetBox="[149,1460,969,1981]" captionTargetId="figure-312@1.[146,900,966,1613]" captionTargetPageId="1" captionText="Text-fig. 1. Spermophilinus bredai (VON MEYER, 1848): 3D models of a right upper jaw with (abnormal) P3, P4 and all molars (mirrored; NMA-2019-1/2352). a – the specimen in occlusal view showing the relative position of the three roots in place of P3 (yellow) compared to P4 (green); b – same view, with focus on P3–P4; c – lingual view; d – mesial view. Scale bars 2mm." figureDoi="http://doi.org/10.5281/zenodo.5383455" httpUri="https://zenodo.org/record/5383455/files/figure.png" pageId="2" pageNumber="176">Text-fig. 1</figureCitation>
). Measurements (L × W in mm): P4: 1.38 × 1.74; M1: 1.64 × 2.06; M2: 1.71 × 2.13; M3: 2.08 × 2.05.
</paragraph>
<paragraph id="8BF3A17FFFD3FF8F2ACF05E9FED3FA4C" blockId="2.[106,745,1413,1535]" pageId="2" pageNumber="176">C o m p a r a t i v e m a t e r i a l. Left upper jaw with P3–M1 from the same locality (NMA-2019-2/2352; Textfig. 2). Measurements: P3: Di: 0.54, H: ~0.38; P4: 1.44 × 1.65; M1: 1.68 × 2.05.</paragraph>
<paragraph id="8BF3A17FFFD3FF8F2ACF067AFACAFAAC" blockId="2.[106,745,1558,2100]" lastBlockId="2.[820,1459,1189,1504]" pageId="2" pageNumber="176">
D e s c r i p t i o n. 
<emphasis id="B9387D6DFFD3FF8F2B03067AFDA0F99C" bold="true" box="[348,552,1558,1582]" pageId="2" pageNumber="176">NMA-2019-1/2352</emphasis>
. The fragmentary maxilla conserves the P3–M 
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situ. The specimen is adult as shown by the wear stage of the teeth.The P3 is not recognizable as such. The crown of the tooth or teeth is/are broken. This break reflects the original condition of the specimen and does not result from damage due to, for example, screen washing of the fossil-enriched sediment. Two posteriorly-directed roots are developed in front of the P4, between the two anterior roots of the last premolar. A very small third root emerges in front of the anterolabial root of the P4. The lingual root is the strongest. The morphology of 
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<emphasis id="B9387D6DFFD3FF8F2B850736FDF6F8C0" box="[474,638,1882,1906]" italics="true" pageId="2" pageNumber="176">Spermophilinus</emphasis>
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molars is well known and, thus, deserves no detailed description. We only notice that the parastyle is not significantly developed in the P4. P4–M3 have a strong, crest-shaped mesostyle. It forms a bridge between the paracone and metacone (P4), or is directed rather towards the labial part of the tooth (M1, M2). The crest is observed in the M3, but included in the expanded posterior part of the tooth. The M1 differs from M2 by the metacone which is more clearly labially differentiated, the protocone which has a more central position, as well as the less well-achieved metalophid/protocone junction.
</paragraph>
<paragraph id="8BF3A17FFFD3FF8F2905054AFA1DFA52" blockId="2.[820,1459,1189,1504]" pageId="2" pageNumber="176">
<emphasis id="B9387D6DFFD3FF8F2905054AFBA3FA8C" bold="true" box="[858,1067,1318,1342]" pageId="2" pageNumber="176">NMA-2019-2/2352</emphasis>
. The specimen shows less trace of tooth wear. The P4 has a parastyle completely included in the anteroloph. The P3 is complete, uniradicular, posteriorly directed, and placed between the anterolabial root and lingual root of the P4. The crown of the tooth is conical and comes into contact with the P4 under the hypothetical parastyle.
</paragraph>
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R e m a r k s. The molars show the characteristics of 
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<emphasis id="B9387D6DFFD3FF8F296B067BFC52F99D" box="[820,986,1559,1583]" italics="true" pageId="2" pageNumber="176">Spermophilinus</emphasis>
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, a genus relatively common in the German middle Miocene (see details in 
<bibRefCitation id="EFDDDC8EFFD3FF8F2E280654FA88F9E2" author="Ziegler, R." box="[1143,1280,1592,1616]" pageId="2" pageNumber="176" pagination="273 - 312" refId="ref5471" refString="Ziegler, R. (2005): The squirrels (Sciuridae, Mammalia) of the Miocene Fossil-Lagerstaette Sandelzhausen (Bavaria, S Germany). - Neues Jahrbuch fur Geologie und Palaontologie, Abhandlungen, 237: 273 - 312. https: // doi. org / 10.1127 / njgpa / 237 / 2005 / 273" type="journal article" year="2005">Ziegler 2005</bibRefCitation>
, for instance). To be brief, four known species of 
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<emphasis id="B9387D6DFFD3FF8F2ED10634FABCF9C2" box="[1166,1332,1624,1648]" italics="true" pageId="2" pageNumber="176">Spermophilinus</emphasis>
</taxonomicName>
(
<taxonomicName id="4C4CDAFCFFD3FF8F2F1A0634FA24F9C2" box="[1349,1452,1624,1648]" class="Mammalia" family="Sciuridae" genus="Spermophilinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="2" pageNumber="176" phylum="Chordata" rank="species" species="besana">
<emphasis id="B9387D6DFFD3FF8F2F1A0634FA24F9C2" box="[1349,1452,1624,1648]" italics="true" pageId="2" pageNumber="176">S. besana</emphasis>
</taxonomicName>
, 
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<emphasis id="B9387D6DFFD3FF8F296B0615FC1BF922" box="[820,915,1656,1680]" italics="true" pageId="2" pageNumber="176">S. bredai</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C4CDAFCFFD3FF8F29FF0615FBAAF922" authorityName="de Bruijn &amp; Mein" authorityYear="1968" box="[928,1058,1656,1680]" class="Mammalia" family="Sciuridae" genus="Spermophilinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="2" pageNumber="176" phylum="Chordata" rank="species" species="turolensis">
<emphasis id="B9387D6DFFD3FF8F29FF0615FBAAF922" box="[928,1058,1656,1680]" italics="true" pageId="2" pageNumber="176">S. turolensis</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C4CDAFCFFD3FF8F2E090615FB50F922" box="[1110,1240,1657,1680]" class="Mammalia" family="Sciuridae" genus="Spermophilinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="2" pageNumber="176" phylum="Chordata" rank="species" species="giganteus">
<emphasis id="B9387D6DFFD3FF8F2E090615FB50F922" box="[1110,1240,1657,1680]" italics="true" pageId="2" pageNumber="176">S. giganteus</emphasis>
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) differ essentially in size. It has long been accepted that these species represent a lineage characterized by an increase in molar size. 
<bibRefCitation id="EFDDDC8EFFD3FF8F2F0E06D5FC12F943" author="Bosma, A. A. &amp; Bruijn, H. de &amp; Wessels, W." pageId="2" pageNumber="176" refId="ref3869" refString="Bosma, A. A., Bruijn, H. de, Wessels, W. (2018): Early and middle Miocene Sciuridae (Mammalia, Rodentia) from Anatolia, Turkey. - Journal of Vertebrate Paleontology, 38 (6): e 1537281 (25 pp.). https: // doi. org / 10.1080 / 02724634.2018.1537281" type="book" year="2018">Bosma et al. (2018)</bibRefCitation>
have recently changed this view by demonstrating the presence of two coexisting species of 
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<emphasis id="B9387D6DFFD3FF8F2EAE0695FA1FF8A3" box="[1265,1431,1785,1809]" italics="true" pageId="2" pageNumber="176">Spermophilinus</emphasis>
</taxonomicName>
in a single site in 
<collectingCountry id="F35BE1EFFFD3FF8F29870776FBBEF880" box="[984,1078,1818,1842]" name="Turkey" pageId="2" pageNumber="176">Anatolia</collectingCountry>
. It is out of the scope of this paper to review the German record in order to discuss the specific homogeneity, and we, thus, consider the Bavarian fossil populations to be monospecific in each locality.
</paragraph>
<paragraph id="8BF3A17FFFD3FF8E290507F7FDD5FDBD" blockId="2.[820,1459,1527,2100]" lastBlockId="3.[129,768,151,624]" lastPageId="3" lastPageNumber="177" pageId="2" pageNumber="176">
The early and middle Miocene 
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<emphasis id="B9387D6DFFD3FF8F2EC607F7FAB5F801" box="[1177,1341,1947,1971]" italics="true" pageId="2" pageNumber="176">Spermophilinus</emphasis>
</taxonomicName>
collections from the Upper Freshwater Molasse were considered to be assigned to 
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<emphasis id="B9387D6DFFD3FF8F298207B0FC62F841" box="[989,1002,2012,2035]" italics="true" pageId="2" pageNumber="176">S</emphasis>
. aff. 
<emphasis id="B9387D6DFFD3FF8F2E7607B7FBE6F841" box="[1065,1134,2011,2035]" italics="true" pageId="2" pageNumber="176">bredai</emphasis>
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for the older localities while 
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<emphasis id="B9387D6DFFD3FF8F296B0790FC15F7A1" box="[820,925,2043,2067]" italics="true" pageId="2" pageNumber="176">S. bredai</emphasis>
</taxonomicName>
is found in younger ones (e.g., 
<bibRefCitation id="EFDDDC8EFFD3FF8F2F700797FC6BF786" author="Ziegler, R. &amp; Fahlbusch, V." pageId="2" pageNumber="176" pagination="3 - 58" refId="ref5530" refString="Ziegler, R., Fahlbusch, V. (1986): Kleinsauger-Faunen aus der basalen Oberen Susswasser-Molasse Niederbayerns. - Zitteliana, A, 14: 3 - 58." type="journal article" year="1986">Ziegler and Fahlbusch 1986</bibRefCitation>
, 
<bibRefCitation id="EFDDDC8EFFD3FF8F29AF0870FBE6F786" author="Kalin, D." box="[1008,1134,2076,2100]" pageId="2" pageNumber="176" refId="ref4630" refString="Kalin, D. (1993): Stratigraphie und Saugetierfaunen der oberen Susswassermolasse der Nordwestschweiz; PhD Thesis - MS, Eidgenossische Technische Hochschule Zurich, Zurich, Swittzerland, 238 pp. (copy in Bibliothek Erdwissenschaften, ETH Zurich)" type="book" year="1993">Kälin 1993</bibRefCitation>
). 
<bibRefCitation id="EFDDDC8EFFD3FF8F2EDC0870FA93F786" author="Ziegler, R." box="[1155,1307,2076,2100]" pageId="2" pageNumber="176" pagination="273 - 312" refId="ref5471" refString="Ziegler, R. (2005): The squirrels (Sciuridae, Mammalia) of the Miocene Fossil-Lagerstaette Sandelzhausen (Bavaria, S Germany). - Neues Jahrbuch fur Geologie und Palaontologie, Abhandlungen, 237: 273 - 312. https: // doi. org / 10.1127 / njgpa / 237 / 2005 / 273" type="journal article" year="2005">Ziegler (2005</bibRefCitation>
; following de 
<bibRefCitation id="EFDDDC8EFFD2FF8E2ADE00FBFE8DFF1D" author="Bruijn, H. de" box="[129,261,151,175]" pageId="3" pageNumber="177" pagination="87 - 102" refId="ref3934" refString="Bruijn, H. de (1995): The Vertebrate Locality Maramena (Macedonia, Greece) at the Turolian-Ruscinian Boundary (Neogene) 8. Sciuridae, Petauristidae and Eomyidae (Rodentia, Mammalia). - Munchner Geowissenschaftliche Abhandlungen, A, 28: 87 - 102." type="journal article" year="1995">Bruijn 1995</bibRefCitation>
) considered that the smaller and older species belong to 
<taxonomicName id="4C4CDAFCFFD2FF8E2AB500D4FED9FF7D" box="[234,337,183,207]" class="Mammalia" family="Sciuridae" genus="Spermophilinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="3" pageNumber="177" phylum="Chordata" rank="species" species="besana">
<emphasis id="B9387D6DFFD2FF8E2AB500D4FED9FF7D" box="[234,337,183,207]" italics="true" pageId="3" pageNumber="177">S. besana</emphasis>
</taxonomicName>
. The measurements of M1 and M2 from selected localities in 
<collectingCountry id="F35BE1EFFFD2FF8E2B0000BBFE56FF5D" box="[351,478,215,239]" name="Switzerland" pageId="3" pageNumber="177">Switzerland</collectingCountry>
and southern 
<collectingCountry id="F35BE1EFFFD2FF8E282A00B4FD50FF42" box="[629,728,216,240]" name="Germany" pageId="3" pageNumber="177">Germany</collectingCountry>
are shown in the 
<figureCitation id="1377BDFAFFD2FF8E2B50009BFE09FEA2" box="[271,385,247,272]" captionStart="Text-fig" captionStartId="3.[147,228,1931,1952]" captionTargetBox="[149,1430,871,1905]" captionTargetId="figure-350@3.[146,900,867,1438]" captionTargetPageId="3" captionText="Text-fig. 3. Scatter diagram of mean L/W values of selected Spermophilinus upper first molars (M1 and M2) from Switzerland and south Germany. The crosses correspond to the minimum and maximum values of the length and width. The intersection gives the value of the mean. Comparative data are after Ziegler and Fahlbusch (1986), Bolliger (1992), Kälin (1993), Ziegler (1995, 2005), Kälin and Engesser (2001), Prieto (2007), Prieto et al. (2009, 2017), Seehuber (2009). Blue: localities younger than ~14 Ma (S. bredai); Green: localities ranging from ~14 Ma to ~15Ma (S. bredai); Red: localities ranging from ~15 Ma to ~16 Ma (S. besana); Yellow: older localities (most S. besana)." figureDoi="http://doi.org/10.5281/zenodo.5383459" httpUri="https://zenodo.org/record/5383459/files/figure.png" pageId="3" pageNumber="177">Text-fig. 3</figureCitation>
. It emerges that it becomes difficult to clearly separate the species in a stratigraphic framework. The use of the average tooth size for all selected localities would certainly have added more clarity to present the evolution of the genus, but would not have allowed us to discuss in detail the size of our specimens (some molar samples are indeed too small). Pending further study of the Petersbuch 136 material, the studied specimens do not oppose a taxonomic assignment to 
<taxonomicName id="4C4CDAFCFFD2FF8E2BA80194FDDEFDBD" baseAuthorityName="VON MEYER" baseAuthorityYear="1848" box="[503,598,503,527]" class="Mammalia" family="Sciuridae" genus="Spermophilinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rodentia" pageId="3" pageNumber="177" phylum="Chordata" rank="species" species="bredai">
<emphasis id="B9387D6DFFD2FF8E2BA80194FDDEFDBD" box="[503,598,503,527]" italics="true" pageId="3" pageNumber="177">S. bredai</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BF3A17FFFD2FF8E2AF90274FE6EFDDD" blockId="3.[129,768,151,624]" pageId="3" pageNumber="177">
It is important to note for the following discussion that we consider the tooth shown in 
<figureCitation id="1377BDFAFFD2FF8E2B8F025BFDB5FDE2" box="[464,573,567,592]" captionStart="Text-fig" captionStartId="2.[125,206,1075,1096]" captionTargetBox="[125,1440,151,1053]" captionTargetId="figure-579@2.[781,1441,150,786]" captionTargetPageId="2" captionText="Text-fig. 2. Spermophilinus bredai (VON MEYER, 1848): left upper jaw with P3–M1 (NMA-2019-2/2352). a – the specimen in occlusal view; b – mesial view; c – labial view. Scale bars 1mm." figureDoi="http://doi.org/10.5281/zenodo.5383457" httpUri="https://zenodo.org/record/5383457/files/figure.png" pageId="3" pageNumber="177">Text-fig. 2</figureCitation>
as a P3 and not as a DP3 that has not been replaced.
</paragraph>
</subSubSection>
</treatment>
</document>