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<mods:title id="D40E46BB26F1A4927589B7913E9581F7">A revision of African helmeted terrapins (Testudines: Pelomedusidae: Pelomedusa), with descriptions of six new species</mods:title>
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<mods:affiliation id="FBFFB9F313425234883640A3161B91DB">Department of Herpetology, Port Elizabeth Museum, P. O. Box 13147, Humewood 6013, South Africa &amp; Department of Zoology, P. O. Box 77000, Nelson Mandela Metropolitan University, Port Elizabeth 6031, South Africa</mods:affiliation>
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<mods:namePart id="665FCE7254D1EE10DBB4E476D78A58D0">Schleicher, Alfred</mods:namePart>
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<mods:affiliation id="61EA3C068232E76C9E20421FBE6B3B30">Department of Biology and Wildlife Diseases, Faculty of Veterinary Hygiene and Ecology, University of Veterinary and Pharmaceutical Sciences, Palackého tř. 1 / 3, 612 42 Brno, Czech Republic</mods:affiliation>
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<treatment id="CB02879FF923FFD5FF74F8B6FBDCFD4F" ID-DOI="http://doi.org/10.5281/zenodo.5082937" ID-GBIF-Taxon="183403852" ID-Zenodo-Dep="5082937" LSID="urn:lsid:plazi:treatment:CB02879FF923FFD5FF74F8B6FBDCFD4F" httpUri="http://treatment.plazi.org/id/CB02879FF923FFD5FF74F8B6FBDCFD4F" lastPageId="21" lastPageNumber="544" pageId="20" pageNumber="543">
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<paragraph id="43143689F923FFD4FF74F8B6FDECF897" blockId="20.[151,549,1806,1832]" box="[151,549,1806,1832]" pageId="20" pageNumber="543">
<heading id="185C81E5F923FFD4FF74F8B6FDECF897" bold="true" box="[151,549,1806,1832]" fontSize="11" level="1" pageId="20" pageNumber="543" reason="1">
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<taxonomicName id="84AB4D0AF923FFD4FF74F8B6FEE3F897" authorityName="Wagler" authorityYear="1830" box="[151,298,1806,1832]" class="Reptilia" family="Pelomedusidae" genus="Pelomedusa" kingdom="Animalia" order="Testudines" pageId="20" pageNumber="543" phylum="Chordata" rank="genus">
<emphasis id="71DFEA9BF923FFD4FF74F8B6FEE3F897" bold="true" box="[151,298,1806,1832]" italics="true" pageId="20" pageNumber="543">Pelomedusa</emphasis>
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candidate species A
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<paragraph id="43143689F923FFD5FF74F8EDFB94FE12" blockId="20.[151,1436,1877,2010]" lastBlockId="21.[151,1440,151,752]" lastPageId="21" lastPageNumber="544" pageId="20" pageNumber="543">
This candidate species from
<collectingCountry id="3BBC7619F923FFD4FE05F8EDFD92F8D1" box="[486,603,1877,1902]" name="Cameroon" pageId="20" pageNumber="543">Cameroon</collectingCountry>
corresponds to mtDNA lineage I of
<bibRefCitation id="273A4B78F923FFD4FBEEF8EDFA96F8D1" author="Vargas-Ramirez, M. &amp; Vences, M. &amp; Branch, W. R. &amp; Daniels, S. R. &amp; Glaw, F. &amp; Hofmeyr, M. D. &amp; Kuchling, G. &amp; Maran, J. &amp; Papenfuss, T. J. &amp; Siroky, P. &amp; Vieites, D. R. &amp; Fritz, U." box="[1037,1375,1877,1902]" pageId="20" pageNumber="543" pagination="428 - 440" refId="ref18768" refString="Vargas-Ramirez, M., Vences, M., Branch, W. R., Daniels, S. R., Glaw, F., Hofmeyr, M. D., Kuchling, G., Maran, J., Papenfuss, T. J., Siroky, P., Vieites, D. R. &amp; Fritz, U. (2010) Deep genealogical lineages in the widely distributed African helmeted terrapin: evidence from mitochondrial and nuclear DNA (Testudines: Pelomedusidae: Pelomedusa subrufa). Molecular Phylogenetics and Evolution, 56, 428 - 440. http: // dx. doi. org / 10.1016 / j. ympev. 2010.03.019" type="journal article" year="2010">
Vargas-Ramírez
<emphasis id="71DFEA9BF923FFD4FB28F8EEFACEF8D1" box="[1227,1287,1877,1902]" italics="true" pageId="20" pageNumber="543">et al.</emphasis>
(2010)
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. It is genetically clearly distinct and differs in 12S and cyt
<emphasis id="71DFEA9BF923FFD4FCF4F8C1FCECF82D" box="[791,805,1913,1938]" italics="true" pageId="20" pageNumber="543">b</emphasis>
sequences by uncorrected
<emphasis id="71DFEA9BF923FFD4FB86F8C2FBBAF82E" box="[1125,1139,1914,1937]" italics="true" pageId="20" pageNumber="543">p</emphasis>
distances resembling the divergences among other
<taxonomicName id="84AB4D0AF923FFD4FE54F825FD50F809" box="[439,665,1949,1974]" class="Reptilia" family="Pelomedusidae" genus="Pelomedusa" kingdom="Animalia" order="Testudines" pageId="20" pageNumber="543" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="71DFEA9BF923FFD4FE54F825FD89F809" box="[439,576,1949,1974]" italics="true" pageId="20" pageNumber="543">Pelomedusa</emphasis>
species
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(
<tableCitation id="0E290332F923FFD4FD44F825FCCEF809" box="[679,775,1949,1974]" captionStart="TABLE 1" captionStartId="5.[151,239,259,282]" captionTargetPageId="5" captionText="TABLE 1. Average uncorrected p distances (percentages) for a 360-bp-long alignment of the 12S gene of Pelomedusa species using the pairwise deletion option of MEGA 6.06. Between-group divergences below diagonal; within-group divergences on the diagonal in boldface. Roman numerals denote the mitochondrial lineage followed by the suggested species name; n = number of sequences." httpUri="http://table.plazi.org/id/17D46601F932FFC5FF74FEBBFD8DFE39" pageId="20" pageNumber="543" tableUuid="17D46601F932FFC5FF74FEBBFD8DFE39">Tables 1</tableCitation>
and
<tableCitation id="0E290332F923FFD4FCDDF825FC87F809" box="[830,846,1949,1974]" captionStart="TABLE 3" captionStartId="7.[151,239,890,913]" captionTargetPageId="7" captionText="TABLE 3. Average uncorrected p distances (percentages) for a 674-bp-long alignment of the cyt b gene of Pelomedusa species using the pairwise deletion option of MEGA 6.06. Between-group divergences below diagonal; within-group divergences on the diagonal in boldface. Roman numerals denote the mitochondrial lineage followed by the suggested species name; n = number of sequences." pageId="20" pageNumber="543">3</tableCitation>
). Since there is just one genetically verified voucher specimen available (Zoologisches Forschungsmuseum Alexander Koenig, Bonn, ZFMK 15171, Mokolo, Margui- Wandala, Extreme North Province,
<collectingCountry id="3BBC7619F922FFD5FDD6FF2FFD65FF0F" box="[565,684,151,176]" name="Cameroon" pageId="21" pageNumber="544">Cameroon</collectingCountry>
, coll. Wolfgang Böhme,
<date id="37151049F922FFD5FC32FF2FFB07FF0F" box="[977,1230,151,176]" pageId="21" pageNumber="544" value="1974-02-14" valueMax="1974-02-17" valueMin="1974-02-14">1417 February 1974</date>
), we refrain from describing this species formally. ZFMK 15171 is a dark-coloured subadult terrapin of
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straight carapacial length, resembling in gross morphology
<taxonomicName id="84AB4D0AF922FFD5FD9CFF59FCF0FF47" authorityName="Petzold &amp; Vargas-Ramírez &amp; Kehlmaier &amp; Vamberger &amp; Branch &amp; Preez &amp; Hofmeyr &amp; Meyer &amp; Schleicher &amp; Široký &amp; Fritz" authorityYear="2014" box="[639,825,223,248]" class="Reptilia" family="Pelomedusidae" genus="Pelomedusa" kingdom="Animalia" order="Testudines" pageId="21" pageNumber="544" phylum="Chordata" rank="species" species="schweinfurthi">
<emphasis id="71DFEA9BF922FFD5FD9CFF59FCF0FF47" box="[639,825,223,248]" italics="true" pageId="21" pageNumber="544">P. schweinfurthi</emphasis>
</taxonomicName>
or dark-coloured
<taxonomicName id="84AB4D0AF922FFD5FBF4FF59FB6DFF47" authorityName="Petzold &amp; Vargas-Ramírez &amp; Kehlmaier &amp; Vamberger &amp; Branch &amp; Preez &amp; Hofmeyr &amp; Meyer &amp; Schleicher &amp; Široký &amp; Fritz" authorityYear="2014" box="[1047,1188,223,248]" class="Reptilia" family="Pelomedusidae" genus="Pelomedusa" kingdom="Animalia" order="Testudines" pageId="21" pageNumber="544" phylum="Chordata" rank="species" species="variabilis">
<emphasis id="71DFEA9BF922FFD5FBF4FF59FB6DFF47" box="[1047,1188,223,248]" italics="true" pageId="21" pageNumber="544">P. variabilis</emphasis>
</taxonomicName>
. However, unlike
<taxonomicName id="84AB4D0AF922FFD5FA64FF59FEE6FEA2" authorityName="Petzold &amp; Vargas-Ramírez &amp; Kehlmaier &amp; Vamberger &amp; Branch &amp; Preez &amp; Hofmeyr &amp; Meyer &amp; Schleicher &amp; Široký &amp; Fritz" authorityYear="2014" class="Reptilia" family="Pelomedusidae" genus="Pelomedusa" kingdom="Animalia" order="Testudines" pageId="21" pageNumber="544" phylum="Chordata" rank="species" species="schweinfurthi">
<emphasis id="71DFEA9BF922FFD5FA64FF59FEE6FEA2" italics="true" pageId="21" pageNumber="544">P. schweinfurthi</emphasis>
</taxonomicName>
, the pectoral scutes of ZFMK 15171 are triangular and do not reach the plastral midseam. Another terrapin from the same source region was only sampled, but not collected (Museum of Zoology, Senckenberg Dresden, Tissue Collection, MTD T 5183, Maroua, Extreme North Province,
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) and is genetically very similar. Our phylogenetic analyses of mtDNA sequences place candidate species A in the well-supported northern clade of
<taxonomicName id="84AB4D0AF922FFD5FF14FE2CFE49FE12" authorityName="Wagler" authorityYear="1830" box="[247,384,404,429]" class="Reptilia" family="Pelomedusidae" genus="Pelomedusa" kingdom="Animalia" order="Testudines" pageId="21" pageNumber="544" phylum="Chordata" rank="genus">
<emphasis id="71DFEA9BF922FFD5FF14FE2CFE49FE12" box="[247,384,404,429]" italics="true" pageId="21" pageNumber="544">Pelomedusa</emphasis>
</taxonomicName>
(
<figureCitation id="DB902A0CF922FFD5FE6CFE2CFE18FE12" box="[399,465,404,429]" captionStart="FIGURE 1" captionStartId="3.[151,250,999,1021]" captionTargetBox="[167,1426,219,969]" captionTargetId="figure-480@3.[151,1436,199,975]" captionTargetPageId="3" captionText="FIGURE 1. Maximum Likelihood tree for helmeted terrapins (Pelomedusa spp.) using 1848 bp of mitochondrial DNA (12S, cyt b, ND4+tRNAs), rooted with Pelusios sinuatus. Terminal clades collapsed to cartoons. The topology of the Bayesian 50% majority rule consensus tree was identical. Numbers along branches indicate bootstrap support and clade support under BI (posterior probabilities) greater than 50 or 0.95, respectively. Asterisks represent maximum support under both methods. Colours and symbol correspond to the map (Fig. 2). On the right, proposed species names indicated; in brackets, mtDNA lineages. For clade membership of individual samples, see Table S1. The questionable sample MTD T 5484 from Swellendam District (Western Cape), South Africa, is highlighted in red (see text). The shown terrapins are Pelomedusa neumanni (top; Kakamega, Kenya) and P. galeata (bottom; Port Elizabeth, South Africa—photos: H. Prokop and W.R. Branch)." figureDoi="http://doi.org/10.5281/zenodo.4915085" httpUri="https://zenodo.org/record/4915085/files/figure.png" pageId="21" pageNumber="544">Fig. 1</figureCitation>
). Therein, it is best understood as having a basal position.
</paragraph>
<paragraph id="43143689F922FFD5FF24FE0FFBDCFD4F" blockId="21.[151,1440,151,752]" pageId="21" pageNumber="544">
<taxonomicName id="84AB4D0AF922FFD5FF24FE0FFE99FE6F" authorityName="Wagler" authorityYear="1830" box="[199,336,439,464]" class="Reptilia" family="Pelomedusidae" genus="Pelomedusa" kingdom="Animalia" order="Testudines" pageId="21" pageNumber="544" phylum="Chordata" rank="genus">
<emphasis id="71DFEA9BF922FFD5FF24FE0FFE99FE6F" box="[199,336,439,464]" italics="true" pageId="21" pageNumber="544">Pelomedusa</emphasis>
</taxonomicName>
candidate species A differs from all other
<taxonomicName id="84AB4D0AF922FFD5FCA6FE0FFBE0FE6F" box="[837,1065,439,464]" class="Reptilia" family="Pelomedusidae" genus="Pelomedusa" kingdom="Animalia" order="Testudines" pageId="21" pageNumber="544" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="71DFEA9BF922FFD5FCA6FE0FFC07FE6F" box="[837,974,439,464]" italics="true" pageId="21" pageNumber="544">Pelomedusa</emphasis>
species
</taxonomicName>
by the presence of guanine (G) instead of cytosine (C) at position 212, by the presence of thymine (T) instead of cytosine (C) at position 222, by the presence of cytosine (C) instead of thymine (T) or adenine (A) at position 302, by the presence of cytosine (C) instead of adenine (A) or guanine (G) at position 325, by the presence of thymine (T) instead of adenine (A) or guanine (G) at position 343, and by the presence of adenine (A) instead of cytosine (C) or thymine (T) at position 345 of the 360-bp-long reference alignment of the 12S rRNA gene (Supporting Information). In addition,
<taxonomicName id="84AB4D0AF922FFD5FF74FD37FEE9FD17" authorityName="Wagler" authorityYear="1830" box="[151,288,655,680]" class="Reptilia" family="Pelomedusidae" genus="Pelomedusa" kingdom="Animalia" order="Testudines" pageId="21" pageNumber="544" phylum="Chordata" rank="genus">
<emphasis id="71DFEA9BF922FFD5FF74FD37FEE9FD17" box="[151,288,655,680]" italics="true" pageId="21" pageNumber="544">Pelomedusa</emphasis>
</taxonomicName>
candidate species A differs from all species except possibly
<taxonomicName id="84AB4D0AF922FFD5FC2CFD29FBF4FD17" baseAuthorityName="Ruppell" baseAuthorityYear="1835" box="[975,1085,655,680]" class="Reptilia" family="Pelomedusidae" genus="Pelomedusa" kingdom="Animalia" order="Testudines" pageId="21" pageNumber="544" phylum="Chordata" rank="species" species="gehafie">
<emphasis id="71DFEA9BF922FFD5FC2CFD29FBF4FD17" box="[975,1085,655,680]" italics="true" pageId="21" pageNumber="544">P. gehafie</emphasis>
</taxonomicName>
and
<taxonomicName id="84AB4D0AF922FFD5FB96FD29FB0FFD17" authorityName="Petzold &amp; Vargas-Ramírez &amp; Kehlmaier &amp; Vamberger &amp; Branch &amp; Preez &amp; Hofmeyr &amp; Meyer &amp; Schleicher &amp; Široký &amp; Fritz" authorityYear="2014" box="[1141,1222,655,680]" class="Reptilia" family="Pelomedusidae" genus="Pelomedusa" kingdom="Animalia" order="Testudines" pageId="21" pageNumber="544" phylum="Chordata" rank="species" species="kobe">
<emphasis id="71DFEA9BF922FFD5FB96FD29FB0FFD17" box="[1141,1222,655,680]" italics="true" pageId="21" pageNumber="544">P. kobe</emphasis>
</taxonomicName>
by the presence of thymine (T) instead of cytosine (C) at position 50 and by the presence of thymine (T) instead of adenine (A) or cytosine (C) at position 65 (character states in
<taxonomicName id="84AB4D0AF922FFD5FD43FD61FCC7FD4F" baseAuthorityName="Ruppell" baseAuthorityYear="1835" box="[672,782,727,752]" class="Reptilia" family="Pelomedusidae" genus="Pelomedusa" kingdom="Animalia" order="Testudines" pageId="21" pageNumber="544" phylum="Chordata" rank="species" species="gehafie">
<emphasis id="71DFEA9BF922FFD5FD43FD61FCC7FD4F" box="[672,782,727,752]" italics="true" pageId="21" pageNumber="544">P. gehafie</emphasis>
</taxonomicName>
and
<taxonomicName id="84AB4D0AF922FFD5FCA7FD61FC5CFD4F" authorityName="Petzold &amp; Vargas-Ramírez &amp; Kehlmaier &amp; Vamberger &amp; Branch &amp; Preez &amp; Hofmeyr &amp; Meyer &amp; Schleicher &amp; Široký &amp; Fritz" authorityYear="2014" box="[836,917,727,752]" class="Reptilia" family="Pelomedusidae" genus="Pelomedusa" kingdom="Animalia" order="Testudines" pageId="21" pageNumber="544" phylum="Chordata" rank="species" species="kobe">
<emphasis id="71DFEA9BF922FFD5FCA7FD61FC5CFD4F" box="[836,917,727,752]" italics="true" pageId="21" pageNumber="544">P. kobe</emphasis>
</taxonomicName>
unknown).
</paragraph>
</subSubSection>
</treatment>
</document>