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<document id="7FF28D4BE092C4366FD83FDE554319FC" ID-DOI="10.1093/zoolinnean/zlad063" ID-ISSN="0024-4082" ID-Zenodo-Dep="10470258" IM.bibliography_approvedBy="felipe" IM.illustrations_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" IM.treatmentCitations_approvedBy="felipe" IM.treatments_approvedBy="felipe" checkinTime="1704718315156" checkinUser="plazi" docAuthor="Mizerovská, Daniela, Martynov, Aleksey A., Mikula, Ondřej, Bryjová, Anna, Meheretu, Yonas, Lavrenchenko, Leonid A. &amp; Bryja, Josef" docDate="2023" docId="853D1219FFFCFFADE909ED1EB7E5FDA1" docLanguage="en" docName="zlad063.pdf" docOrigin="Zoological Journal of the Linnean Society 199 (4)" docSource="http://dx.doi.org/10.1093/zoolinnean/zlad063" docStyle="DocumentStyle:4F230B9370E98E256D973D6DFB57F36C.5:ZoolJLinnSoc.2023-.journal_article" docStyleId="4F230B9370E98E256D973D6DFB57F36C" docStyleName="ZoolJLinnSoc.2023-.journal_article" docStyleVersion="4" docTitle="Otomys fortior Thomas 1906" docType="treatment" docVersion="3" lastPageNumber="1073" masterDocId="79046A61FFF2FFA3E839E84DB432FFBB" masterDocTitle="Genomic diversity, evolutionary history, and species limits of the endemic Ethiopian laminate-toothed rats (genus Otomys, Rodentia: Muridae)" masterLastPageNumber="1077" masterPageNumber="1059" pageNumber="1073" updateTime="1704748301190" updateUser="felipe">
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<mods:title id="16E83CED6913C2A57A9BDDA48219E584">Genomic diversity, evolutionary history, and species limits of the endemic Ethiopian laminate-toothed rats (genus Otomys, Rodentia: Muridae)</mods:title>
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<mods:namePart id="28C9082BDE5A8A11EC02FB97C81A4A5C">Mizerovská, Daniela</mods:namePart>
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<mods:namePart id="925EFC6281E74524FD58640D676CA378">Martynov, Aleksey A.</mods:namePart>
<mods:affiliation id="7DCCD359D194165D852E90445A4B5433">A. N. Severtsov Institute of Ecology and Evolution of the Russian Academy of Sciences, Moscow, Russia</mods:affiliation>
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<mods:namePart id="000B74112A564E08641DB9214B99104A">Mikula, Ondřej</mods:namePart>
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<mods:namePart id="E24C4F64BAB54165DF617329B40B9193">Bryjová, Anna</mods:namePart>
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<mods:namePart id="6217BE77320C9CE9FAAEE8ABED5BF36F">Meheretu, Yonas</mods:namePart>
<mods:affiliation id="4E2E8BCE9F1BCC70E4E0E26C450DCECA">Institute of Vertebrate Biology of the Czech Academy of Sciences, Brno, Czech Republic &amp; Department of Biology and Institute of Mountain Research and Development, Mekelle University, Mekelle, Tigray, Ethiopia &amp; Department of Wildlife, Fish, and Environmental Studies, Swedish University of Agricultural Sciences, Umeå, Sweden</mods:affiliation>
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<mods:namePart id="73B7F51B3E74E1A143AEAFDE2BA8AF56">Lavrenchenko, Leonid A.</mods:namePart>
<mods:affiliation id="B45C2FF63BF17FA64CD7068F9A642CD2">A. N. Severtsov Institute of Ecology and Evolution of the Russian Academy of Sciences, Moscow, Russia &amp; Institute of Vertebrate Biology of the Czech Academy of Sciences, Brno, Czech Republic &amp; Department of Botany and Zoology, Faculty of Science, Masaryk University, Brno, Czech Republic</mods:affiliation>
<mods:nameIdentifier id="6605888083F75979608999F19AACFE44" type="email">bryja@ivb.cz</mods:nameIdentifier>
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<mods:namePart id="8AF9AE5E8903B00C0C2A5315C6CDB8F7">Bryja, Josef</mods:namePart>
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<mods:date id="AAB39244275420ED73701E06B9542B41">2023</mods:date>
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<heading id="56631463FFFCFFADE909ED1EB66BFAD6" bold="true" box="[304,601,1362,1389]" centered="true" fontSize="11" level="2" pageId="14" pageNumber="1073" reason="2">
<taxonomicName id="CA94D88CFFFCFFADE909ED1EB66BFAD6" ID-CoL="4B7KK" authority="Thomas 1906" authorityName="Thomas" authorityYear="1906" box="[304,601,1362,1389]" class="Mammalia" family="Muridae" genus="Otomys" kingdom="Animalia" order="Rodentia" pageId="14" pageNumber="1073" phylum="Chordata" rank="species" species="fortior">
<emphasis id="3FE07F1DFFFCFFADE909ED1EB66BFAD6" bold="true" box="[304,601,1362,1389]" pageId="14" pageNumber="1073">
<emphasis id="3FE07F1DFFFCFFADE909ED1EB5F6FAD7" bold="true" box="[304,452,1362,1389]" italics="true" pageId="14" pageNumber="1073">Otomys fortior</emphasis>
Thomas 1906
</emphasis>
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</heading>
</paragraph>
<paragraph id="0D2BA30FFFFCFFADE8B8ED34B502FA2A" blockId="14.[129,778,1401,1676]" box="[129,304,1401,1425]" pageId="14" pageNumber="1073">Charada Vlei Rat</paragraph>
</subSubSection>
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<paragraph id="0D2BA30FFFFCFFADE8B8EDD4B4E8F995" blockId="14.[129,778,1401,1676]" pageId="14" pageNumber="1073">
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<taxonomicName id="CA94D88CFFFCFFADE8B8EDD4B577FA0A" ID-CoL="4B7L9" authorityName="Thomas" authorityYear="1906" box="[129,325,1433,1457]" class="Mammalia" family="Muridae" genus="Otomys" kingdom="Animalia" order="Rodentia" pageId="14" pageNumber="1073" phylum="Chordata" rank="species" species="typus">
<emphasis id="3FE07F1DFFFCFFADE8B8EDD4B577FA0A" box="[129,325,1433,1457]" italics="true" pageId="14" pageNumber="1073">Otomys typus fortior</emphasis>
</taxonomicName>
: Thomas 1906:302 (
<typeStatus id="D22F1DADFFFCFFADEA2AEDD4B676FA0A" box="[531,580,1433,1457]" pageId="14" pageNumber="1073">type</typeStatus>
locality—
<collectingCountry id="7583E39FFFFCFFADEA94EDD4B737FA0A" box="[685,773,1433,1457]" name="Ethiopia" pageId="14" pageNumber="1073">Ethiopia</collectingCountry>
, Kaffa, Charada Forest,
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);
<treatmentCitation id="8C35851EFFFCFFADE9E9EDF5B642FA6B" author="Allen GM" box="[464,624,1464,1488]" page="348" pageId="14" pageNumber="1073" year="1939">
<bibRefCitation id="6905DEFEFFFCFFADE9E9EDF5B642FA6B" author="Allen GM" box="[464,624,1464,1488]" pageId="14" pageNumber="1073" pagination="1 - 763" refId="ref15278" refString="Allen GM. A checklist of African mammals. Bulletin of the Museum of Comparative Zoology at Harvard College 1939; 83: 1 - 763." type="journal article" year="1939">Allen 1939:348</bibRefCitation>
</treatmentCitation>
(listed as valid subspecies);
<treatmentCitation id="8C35851EFFFCFFADE933ED9AB5E4FA4B" author="Ellerman JR" box="[266,470,1495,1520]" page="321" pageId="14" pageNumber="1073" year="1941">
<bibRefCitation id="6905DEFEFFFCFFADE933ED9AB5E4FA4B" author="Ellerman JR" box="[266,470,1495,1520]" pageId="14" pageNumber="1073" refId="ref16142" refString="Ellerman JR. The Families and Genera of Living Rodents. Vol II - Muridae. London: British Museum, 1941." type="book" year="1941">Ellerman 1941:321</bibRefCitation>
</treatmentCitation>
(listed as valid subspecies);
<treatmentCitation id="8C35851EFFFCFFADE8B8EDBAB576F9B4" author="Bohmann LV" box="[129,324,1527,1551]" page="45" pageId="14" pageNumber="1073" year="1952">
<bibRefCitation id="6905DEFEFFFCFFADE8B8EDBAB576F9B4" author="Bohmann LV" box="[129,324,1527,1551]" pageId="14" pageNumber="1073" pagination="1 - 80" refId="ref15489" refString="Bohmann LV. Die afrikanische Nagergauung Otomys F. Cuvier. Zeitschriss fur Saugetierkunde 1952; 18: 1 - 80." type="journal article" year="1952">Bohmann 1952:45</bibRefCitation>
</treatmentCitation>
(taxonomic revision, retained as valid subspecies).
</treatmentCitationGroup>
</paragraph>
<paragraph id="0D2BA30FFFFCFFADE8B8EE7BB650F9F6" blockId="14.[129,778,1401,1676]" box="[129,610,1589,1613]" pageId="14" pageNumber="1073">
<taxonomicName id="CA94D88CFFFCFFADE8B8EE7BB59AF9F6" ID-CoL="4B7KK" authority="Dollman 1915" authorityName="Dollman" authorityYear="1915" box="[129,424,1589,1613]" class="Mammalia" family="Muridae" genus="Otomys" kingdom="Animalia" order="Rodentia" pageId="14" pageNumber="1073" phylum="Chordata" rank="species" species="fortior">
<emphasis id="3FE07F1DFFFCFFADE8B8EE7BB53DF9F6" box="[129,271,1589,1613]" italics="true" pageId="14" pageNumber="1073">Otomys fortior</emphasis>
<bibRefCitation id="6905DEFEFFFCFFADE92DEE78B59AF9F6" author="Dollman G" box="[276,424,1589,1613]" pageId="14" pageNumber="1073" pagination="149 - 70" refId="ref16065" refString="Dollman G. IX. - On the swamp-rats (Otomys) of East Africa. Annals and Magazine of Natural History 1915; 15: 149 - 70." type="journal article" year="1915">Dollman 1915</bibRefCitation>
</taxonomicName>
—Kaffa,
<collectingCountry id="7583E39FFFFCFFADEA3DEE78B66CF9F6" box="[516,606,1589,1613]" name="Ethiopia" pageId="14" pageNumber="1073">Ethiopia</collectingCountry>
.
</paragraph>
<paragraph id="0D2BA30FFFFCFFADE8B8EE18B68FF937" blockId="14.[129,778,1401,1676]" pageId="14" pageNumber="1073">
<taxonomicName id="CA94D88CFFFCFFADE8B8EE18B538F9D7" ID-CoL="4B7L9" authorityName="Thomas" authorityYear="1906" box="[129,266,1621,1644]" class="Mammalia" family="Muridae" genus="Otomys" kingdom="Animalia" order="Rodentia" pageId="14" pageNumber="1073" phylum="Chordata" rank="species" species="typus">
<emphasis id="3FE07F1DFFFCFFADE8B8EE18B538F9D7" box="[129,266,1621,1644]" italics="true" pageId="14" pageNumber="1073">Otomys typus</emphasis>
</taxonomicName>
(
<taxonomicName id="CA94D88CFFFCFFADE926EE18B56FF9D6" ID-CoL="4B7KK" authorityName="Dollman" authorityYear="1915" box="[287,349,1621,1645]" class="Mammalia" family="Muridae" genus="Otomys" kingdom="Animalia" order="Rodentia" pageId="14" pageNumber="1073" phylum="Chordata" rank="species" species="fortior">
<emphasis id="3FE07F1DFFFCFFADE926EE18B56FF9D6" box="[287,349,1621,1645]" italics="true" pageId="14" pageNumber="1073">fortior</emphasis>
</taxonomicName>
)
<bibRefCitation id="6905DEFEFFFCFFADE94BEE18B639F9D6" author="Misonne X" box="[370,523,1621,1645]" pageId="14" pageNumber="1073" refId="ref17143" refString="Misonne X. Part 6, Order Rodentia. In: Meester J, Setzer HW (eds.), The Mammals of Affica: An Identification Manual. Washington, D. C: Smithsonian Institution Press, 1974." type="book" year="1974">Misonne 1974</bibRefCitation>
,
<bibRefCitation id="6905DEFEFFFCFFADEA23EE18B6EBF9D6" author="Yalden DW &amp; Largen MJ &amp; Kock D" box="[538,729,1621,1645]" pageId="14" pageNumber="1073" pagination="1 - 118" refId="ref18779" refString="Yalden DW, Largen MJ, Kock D. Catalogue of the mammals of Ethiopia. 2. Insectivora and Rodentia. Monitore Zoologico Italiano, Supplemento 1976; 8: 1 - 118." type="journal article" year="1976">
Yalden
<emphasis id="3FE07F1DFFFCFFADEA50EE18B6AEF9D6" box="[617,668,1621,1645]" italics="true" pageId="14" pageNumber="1073">et al.</emphasis>
1976
</bibRefCitation>
and Musser and Carleton 1993—without indication of rank.
</paragraph>
</subSubSection>
<subSubSection id="458EF084FFFCFFADE8B8EEE0B7E5FDA1" pageId="14" pageNumber="1073" type="materials_examined">
<paragraph id="0D2BA30FFFFCFFADE8B8EEE0B592F97E" blockId="14.[129,416,1709,1733]" box="[129,416,1709,1733]" pageId="14" pageNumber="1073">
<materialsCitation id="BDFCA952FFFCFFADE8B8EEE0B592F97E" box="[129,416,1709,1733]" collectionCode="BMNH" pageId="14" pageNumber="1073" specimenCode="BMNH 6.11" specimenCount="1" typeStatus="holotype">
<emphasis id="3FE07F1DFFFCFFADE8B8EEE0B4D3F97E" box="[129,225,1709,1733]" italics="true" pageId="14" pageNumber="1073">
<typeStatus id="D22F1DADFFFCFFADE8B8EEE0B4EEF97E" box="[129,220,1709,1733]" pageId="14" pageNumber="1073" type="holotype">Holotype</typeStatus>
:
</emphasis>
<specimenCode id="5D320B74FFFCFFADE8DFEEE3B558F97E" box="[230,362,1709,1733]" collectionCode="BMNH" pageId="14" pageNumber="1073">BMNH 6.11</specimenCode>
.1.29.
</materialsCitation>
</paragraph>
<paragraph id="0D2BA30FFFFCFFADE8B8EEAAB6E5F944" blockId="14.[129,727,1767,1791]" box="[129,727,1767,1791]" pageId="14" pageNumber="1073">
<emphasis id="3FE07F1DFFFCFFADE8B8EEAAB531F944" box="[129,259,1767,1791]" italics="true" pageId="14" pageNumber="1073">Type locality:</emphasis>
<collectingCountry id="7583E39FFFFCFFADE931EEAAB553F944" box="[264,353,1767,1791]" name="Ethiopia" pageId="14" pageNumber="1073">Ethiopia</collectingCountry>
, Charada Forest, Kaffa,
<quantity id="CA6C0EEAFFFCFFADEA6AEEAAB693F945" box="[595,673,1767,1791]" metricMagnitude="3" metricUnit="m" metricValue="1.829" pageId="14" pageNumber="1073" unit="m" value="1829.0">1829 m</quantity>
a.s.l..
</paragraph>
<paragraph id="0D2BA30FFFFCFFADE8B8EF6CB7FFFF5C" blockId="14.[128,778,1824,1973]" lastBlockId="14.[825,1474,144,231]" pageId="14" pageNumber="1073">
<emphasis id="3FE07F1DFFFCFFADE8B8EF6CB5F6F883" box="[129,452,1824,1848]" italics="true" pageId="14" pageNumber="1073">
Taxonomic notes:
<taxonomicName id="CA94D88CFFFCFFADE90DEF6DB5F6F883" authorityName="Dollman" authorityYear="1915" box="[308,452,1824,1848]" class="Mammalia" family="Muridae" genus="Otomys" kingdom="Animalia" order="Rodentia" pageId="14" pageNumber="1073" phylum="Chordata" rank="species" species="fortior">Otomys fortior</taxonomicName>
</emphasis>
shares a common karyotype of 2n = 58, NFa = 58 with
<taxonomicName id="CA94D88CFFFCFFADE943EF0DB5FEF8EC" box="[378,460,1856,1879]" class="Mammalia" family="Muridae" genus="Otomys" kingdom="Animalia" order="Rodentia" pageId="14" pageNumber="1073" phylum="Chordata" rank="species" species="typus">
<emphasis id="3FE07F1DFFFCFFADE943EF0DB5BFF8EC" box="[378,397,1856,1879]" italics="true" pageId="14" pageNumber="1073">O</emphasis>
.
<emphasis id="3FE07F1DFFFCFFADE9A1EF0DB5FEF8EC" box="[408,460,1856,1879]" italics="true" pageId="14" pageNumber="1073">
<typeStatus id="D22F1DADFFFCFFADE9A1EF0DB5FEF8EC" box="[408,460,1856,1879]" pageId="14" pageNumber="1073">typus</typeStatus>
</emphasis>
</taxonomicName>
and
<taxonomicName id="CA94D88CFFFCFFADE9C6EF0DB66BF8EC" box="[511,601,1855,1879]" class="Mammalia" family="Muridae" genus="Otomys" kingdom="Animalia" order="Rodentia" pageId="14" pageNumber="1073" phylum="Chordata" rank="species" species="helleri">
<emphasis id="3FE07F1DFFFCFFADE9C6EF0DB620F8EC" box="[511,530,1856,1879]" italics="true" pageId="14" pageNumber="1073">O</emphasis>
.
<emphasis id="3FE07F1DFFFCFFADEA24EF72B66BF8EC" box="[541,601,1855,1879]" italics="true" pageId="14" pageNumber="1073">helleri</emphasis>
</taxonomicName>
, as well as higher number of eight laminae in M3. This is the most distinct species in the
<typeStatus id="D22F1DADFFFCFFADE886EF33B522F82D" box="[191,272,1918,1942]" pageId="14" pageNumber="1073">
<collectionCode id="6B853BCAFFFCFFADE886EF33B522F82D" box="[191,272,1918,1942]" pageId="14" pageNumber="1073">TYPUS</collectionCode>
</typeStatus>
group, both genomically (
<figureCitation id="95AFBF8AFFFCFFADEA25EF33B640F82D" box="[540,626,1918,1942]" captionStart-0="Figure 3" captionStart-1="Figure 4" captionStart-2="Figure 5" captionStart-3="Figure 6" captionStartId-0="7.[113,178,1133,1157]" captionStartId-1="8.[129,194,818,842]" captionStartId-2="8.[129,194,1586,1610]" captionStartId-3="9.[113,178,1166,1190]" captionTargetBox-0="[115,1458,146,1104]" captionTargetBox-1="[131,1473,146,789]" captionTargetBox-2="[242,1363,1032,1558]" captionTargetBox-3="[116,1456,575,1136]" captionTargetId-0="figure-433@7.[114,1458,144,1105]" captionTargetId-1="figure-169@8.[129,1473,144,790]" captionTargetId-2="figure-277@8.[240,1363,1030,1559]" captionTargetId-3="figure-261@9.[114,1458,572,1138]" captionTargetPageId-0="7" captionTargetPageId-1="8" captionTargetPageId-2="8" captionTargetPageId-3="9" captionText-0="Figure 3. ML tree of concatenated SNPs loci developed by ddÞD approach. Numbers above the branches indicate the bootstrap support; coloured bars correspond to six presumed Ethiopian species (for their distribution see Fig. 2B and C). The tree is based on the dataset of 5587 loci and 48 200 SNPs with a minimum 10% occupancy, but the trees from more filtered data had concordant topologies, only with progressively lower support of clades (not shown)." captionText-1="Figure 4. The result of sNMF analysis based on 80% occupancy dataset, i.e. 3141 biallelic SNPs (one SNP per locus).Vertical bars show individual genotypes as composed of ancestral population gene pools, whose proportional contributions are indicated by coloured segments. Horizontal lines at the boưom delimit clusters defined by the dominant ancestral contributions. The associated labels identify clusters with named species or their geographically restricted populations (B. S. = Borena Saynt, D. S. = Debre Sina).Note that specimens from the populations in Borena Saynt and Aboye Gara, identified as O. simiensis by mtDNA and concatenated ddÞD data, cluster with O. cf. yaldeni from Ankober (purple cluster)." captionText-2="Figure 5. Co-ancestry matrix and delimitation of gene pools by two different methods: fineSTRUCTURE (18 clusters shown by grey squares) and Infomap (three clusters shown by black squares). The intensity of the colour (from yellow to blue) indicates the level of co-ancestry among individual genomes." captionText-3="Figure 6. Species trees obtained from the MSC analyses in BPP. The MCC tree from A11 analysis (A) is shown with clade PPs in its branch labels, while the tree from A00 analysis (B) shows gdi values. In circles, there are high gdi values (≥0.70) obtained for the extant or ancestral species in the original A00 analysis; in diamonds, there are high or low gdi values (white and grey background, respectively) that were obtained by the iterative merging procedure. Colour bars indicate six nominal Ethiopian species." figureDoi-0="http://doi.org/10.5281/zenodo.10470264" figureDoi-1="http://doi.org/10.5281/zenodo.10470266" figureDoi-2="http://doi.org/10.5281/zenodo.10470268" figureDoi-3="http://doi.org/10.5281/zenodo.10470270" httpUri-0="https://zenodo.org/record/10470264/files/figure.png" httpUri-1="https://zenodo.org/record/10470266/files/figure.png" httpUri-2="https://zenodo.org/record/10470268/files/figure.png" httpUri-3="https://zenodo.org/record/10470270/files/figure.png" pageId="14" pageNumber="1073">Figs 36</figureCitation>
) and ecologically (the only species living in forest clearings at relatively low elevation in south-western
<collectingCountry id="7583E39FFFFCFFADEC70E8DDB096FF13" box="[1097,1188,144,168]" name="Ethiopia" pageId="14" pageNumber="1073">Ethiopia</collectingCountry>
;
<figureCitation id="95AFBF8AFFFCFFADEC94E8DDB0D6FF13" box="[1197,1252,144,168]" captionStart="Figure 2" captionStartId="6.[129,194,1297,1321]" captionTargetBox="[131,1473,147,1268]" captionTargetId="figure-73@6.[129,1473,144,1269]" captionTargetPageId="6" captionText="Figure 2. A, Cytb phylogeny of two groups (the TYPUS and SIMIENSIS groups) of endemic Ethiopian taxa from the O. typus complex. The numbers above the branches indicate BI posterior probability/ML bootstrap support.Maps show the distribution of the mitochondrial diversity in the Ethiopian SIMIENSIS group (B) and TYPUS group (C). Coloured circles indicate the localities with genetically characterized individuals; different colours correspond to six Ethiopian species marked by coloured bars on the right side of the phylogenetic tree.Stars of particular colours show the type localities (there are two stars for O. cheesmani; Dangila is the type locality, the paratype specimen was collected from Gojam). White dots are records of non-genotyped Otomys [based on Yalden et al. (1976, 1996) and the GBIF database; see more details in Bryja et al. (2019a)]. The shades of grey on the map indicate elevation." figureDoi="http://doi.org/10.5281/zenodo.10470262" httpUri="https://zenodo.org/record/10470262/files/figure.png" pageId="14" pageNumber="1073">Fig. 2</figureCitation>
). Morphological analysis did not show any special ecological adaptations to different environments.
</paragraph>
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<emphasis id="3FE07F1DFFFCFFADEB00E945B785FE9B" box="[825,951,264,288]" italics="true" pageId="14" pageNumber="1073">Distribution:</emphasis>
The species lives in tropical forests in south-western
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, west of the Omo River, from where we provide numerous new data [compared to
<bibRefCitation id="6905DEFEFFFCFFADECB0E90AB167FEE4" author="Taylor PJ &amp; Lavrenchenko LA &amp; Carleton MD" box="[1161,1365,327,351]" pageId="14" pageNumber="1073" pagination="1 - 66" refId="ref18434" refString="Taylor PJ, Lavrenchenko LA, Carleton MD, et al. Specific limits and emerging diversity pauerns in East African populations of laminate-toothed rats, genus Otomys (Muridae: Murinae: Otomyini): revision of the Otomys typus complex. Zootaxa 2011; 3024: 1 - 66." type="journal article" year="2011">
Taylor
<emphasis id="3FE07F1DFFFCFFADECEAE90AB136FEE4" box="[1235,1284,327,351]" italics="true" pageId="14" pageNumber="1073">et al.</emphasis>
(2011)
</bibRefCitation>
]. The elevational range of genetically confirmed records is
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a.s.l.. The specimens from the Jimma area were assigned to
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<emphasis id="3FE07F1DFFFCFFADEB00E9E8B77EFE07" box="[825,844,421,444]" italics="true" pageId="14" pageNumber="1073">O</emphasis>
.
<emphasis id="3FE07F1DFFFCFFADEB62E9E8B7ABFE06" box="[859,921,421,445]" italics="true" pageId="14" pageNumber="1073">fortior</emphasis>
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based on geographic and ecological data. However, based on cranial characteristics, they are related to
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<emphasis id="3FE07F1DFFFCFFADED78E989B166FE60" box="[1345,1364,452,475]" italics="true" pageId="14" pageNumber="1073">O</emphasis>
.
<emphasis id="3FE07F1DFFFCFFADED66E989B1F0FE67" box="[1375,1474,452,476]" italics="true" pageId="14" pageNumber="1073">cheesmani</emphasis>
(
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Taylor
<emphasis id="3FE07F1DFFFCFFADEBB5E9A9B789FE40" box="[908,955,483,507]" italics="true" pageId="14" pageNumber="1073">et al.</emphasis>
2011
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)
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, which might be just an artefact of morphological analysis.
</paragraph>
</subSubSection>
</treatment>
</document>