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<mods:title id="A0E8E0D3F77971E46F2CDF0BCAE25129">Disentangling the Pelomedusa complex using type specimens and historical DNA (Testudines: Pelomedusidae)</mods:title>
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<mods:namePart id="AA0B116BE268E9F38C8B2095D9D6BC49">Fritz, Uwe</mods:namePart>
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<mods:namePart id="7B0D5B56B159D67118475C1FFE10DB78">Petzold, Alice</mods:namePart>
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<mods:affiliation id="C3B1C7641391948932DDE30A56EEEBC6">Museum of Zoology, Senckenberg Dresden, A. B. Meyer Building, 01109 Dresden, Germany</mods:affiliation>
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<mods:namePart id="36E6A8B7F1952729DCB60A31D247D040">Campbell, Patrick</mods:namePart>
<mods:affiliation id="4CC1E1ED4F416C4128E47F0B33AAF7E1">Department of Zoology, Darwin Centre (DC 1), Natural History Museum, Cromwell Road, South Kensington, London SW 7 5 BD, England</mods:affiliation>
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<mods:namePart id="C79929F8F8B36E780976236D8CD89B79">Hofmeyr, Margaretha D.</mods:namePart>
<mods:affiliation id="36EF818DFC241A59A8A391EC56A29636">Chelonian Biodiversity and Conservation, Department of Biodiversity and Conservation Biology, University of the Western Cape, Private Bag X 17, Bellville 7535, South Africa</mods:affiliation>
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<mods:namePart id="845B5B6213535A38B3AD4A87169906DB">Branch, William R.</mods:namePart>
<mods:affiliation id="76551C8E20B0F127F41A964447F0A160">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 Corresponding author. E-mail: uwe. fritz @ senckenberg. de</mods:affiliation>
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<treatment id="03B68780CF55102A18C3EEA7FB2FF882" ID-DOI="http://doi.org/10.5281/zenodo.5082877" ID-GBIF-Taxon="183403826" ID-Zenodo-Dep="5082877" LSID="urn:lsid:plazi:treatment:03B68780CF55102A18C3EEA7FB2FF882" httpUri="http://treatment.plazi.org/id/03B68780CF55102A18C3EEA7FB2FF882" lastPageId="6" lastPageNumber="507" pageId="5" pageNumber="506">
<subSubSection id="C305651DCF55102918C3EEA7FC9DFA1B" box="[151,995,1438,1464]" pageId="5" pageNumber="506" type="nomenclature">
<paragraph id="8BA03696CF55102918C3EEA7FC9DFA1B" blockId="5.[151,995,1438,1464]" box="[151,995,1438,1464]" pageId="5" pageNumber="506">
<heading id="D0E881FACF55102918C3EEA7FC9DFA1B" bold="true" box="[151,995,1438,1464]" fontSize="11" level="1" pageId="5" pageNumber="506" reason="1">
<emphasis id="B96BEA84CF55102918C3EEA7FC9DFA1B" bold="true" box="[151,995,1438,1464]" pageId="5" pageNumber="506">
<taxonomicName id="4C1F4D15CF55102918C3EEA7FD5CFA1B" authority="Lacepede, 1788" authorityName="Lacepede" authorityYear="1788" box="[151,546,1438,1464]" class="Reptilia" family="Testudinidae" genus="Testudo" kingdom="Animalia" order="Testudines" pageId="5" pageNumber="506" phylum="Chordata" rank="species" species="subrufa">
<emphasis id="B96BEA84CF55102918C3EEA7FE24FA1B" bold="true" box="[151,346,1438,1464]" italics="true" pageId="5" pageNumber="506">Testudo subrufa</emphasis>
<bibRefCitation id="EF8E4B67CF5510291935EEA7FD5CFA1B" author="Lacepede" box="[353,546,1438,1464]" pageId="5" pageNumber="506" refId="ref12474" refString="Lacepede, [B. G.] de (1788) Histoire naturelle des quadrupedes ovipares et des serpens. Tome premier. Hotel de Thou, Paris, (17) + (1, errata) + 651 pp., 1 folded table, 41 plates. http: // dx. doi. org / 10.5962 / bhl. title. 5036" type="book" year="1788">Lacepède, 1788</bibRefCitation>
</taxonomicName>
=
<emphasis id="B96BEA84CF5510291A15EEA7FC7AFA1B" bold="true" box="[577,772,1438,1464]" italics="true" pageId="5" pageNumber="506">Testudo subrufa</emphasis>
<bibRefCitation id="EF8E4B67CF5510291B5FEEA7FC9DFA1B" author="Bonnaterre" box="[779,995,1438,1464]" pageId="5" pageNumber="506" refId="ref11147" refString="Bonnaterre, [P. J.] (1789) Tableau encyclopedique et methodique des trois regnes de la nature. Erpetologie. Panckoucke, Hotel de Thou, Paris, xxviii + 70 + (1) pp., 35 plates. http: // dx. doi. org / 10.5962 / bhl. title. 59326" type="book" year="1789">Bonnaterre, 1789</bibRefCitation>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C305651DCF55102918C3EEDAFC85F857" pageId="5" pageNumber="506" type="description">
<paragraph id="8BA03696CF55102918C3EEDAFB38F92F" blockId="5.[151,1437,1507,2036]" pageId="5" pageNumber="506">
Only when
<bibRefCitation id="EF8E4B67CF551029194CEEDAFEBEFA5F" author="Mertens, R." box="[280,448,1507,1532]" pageId="5" pageNumber="506" pagination="139 - 142" refId="ref12650" refString="Mertens, R. (1937) Bemerkungen uber die Rassen von Pelomedusa subrufa (La Cepede). Zoologischer Anzeiger, 117, 139 - 142." type="journal article" year="1937">Mertens (1937</bibRefCitation>
: p. 139) and
<bibRefCitation id="EF8E4B67CF5510291A1AEEDAFC72FA5F" author="Loveridge, A." box="[590,780,1507,1532]" pageId="5" pageNumber="506" pagination="467 - 524" refId="ref12616" refString="Loveridge, A. (1941) Revision of the African terrapin [sic] of the family Pelomedusidae. Bulletin of the Museum of Comparative Zoology, 88, 467 - 524." type="journal article" year="1941">Loveridge (1941</bibRefCitation>
: p. 470) reinstated the scientific name
<taxonomicName id="4C1F4D15CF5510291CE3EEDAFE22F983" authority="(Lacepede, 1788)" baseAuthorityName="Lacepede" baseAuthorityYear="1788" class="Reptilia" family="Pelomedusidae" genus="Pelomedusa" kingdom="Animalia" order="Testudines" pageId="5" pageNumber="506" phylum="Chordata" rank="species" species="subrufa">
<emphasis id="B96BEA84CF5510291CE3EEDAFAE5FA5F" box="[1207,1435,1507,1532]" italics="true" pageId="5" pageNumber="506">Pelomedusa subrufa</emphasis>
(
<bibRefCitation id="EF8E4B67CF55102918CAED3EFE2AF983" author="Lacepede" box="[158,340,1543,1568]" pageId="5" pageNumber="506" refId="ref12474" refString="Lacepede, [B. G.] de (1788) Histoire naturelle des quadrupedes ovipares et des serpens. Tome premier. Hotel de Thou, Paris, (17) + (1, errata) + 651 pp., 1 folded table, 41 plates. http: // dx. doi. org / 10.5962 / bhl. title. 5036" type="book" year="1788">Lacepède, 1788</bibRefCitation>
)
</taxonomicName>
for helmeted terrapins, this name combination found wide acceptance. Previously,
<taxonomicName id="4C1F4D15CF5510291D47ED3EFED0F9E7" authority="(Schoepff, 1792)" baseAuthorityName="Schoepff" baseAuthorityYear="1792" class="Reptilia" family="Pelomedusidae" genus="Pelomedusa" kingdom="Animalia" order="Testudines" pageId="5" pageNumber="506" phylum="Chordata" rank="species" species="galeata">
<emphasis id="B96BEA84CF5510291D47ED3EFF95F9E7" italics="true" pageId="5" pageNumber="506">Pelomedusa galeata</emphasis>
(
<bibRefCitation id="EF8E4B67CF55102918ACED12FEDBF9E7" author="Schoepff, I. D." box="[248,421,1579,1604]" pageId="5" pageNumber="506" refId="ref13074" refString="Schoepff, I. D. (1792) Historia Testudinum iconibus illustrata. Ioannes Iacobus Palm, Erlangen, XII + 136 pp., 31 plates. http: // dx. doi. org / 10.5962 / bhl. title. 5109" type="book" year="1792">Schoepff, 1792</bibRefCitation>
)
</taxonomicName>
was used by most authors (see below under
<taxonomicName id="4C1F4D15CF5510291BC8ED12FA7EF9E7" authority="Schoepff, 1792" authorityName="Schoepff" authorityYear="1792" box="[924,1280,1579,1604]" class="Reptilia" family="Testudinidae" genus="Testudo" kingdom="Animalia" order="Testudines" pageId="5" pageNumber="506" phylum="Chordata" rank="species" species="galeata">
<emphasis id="B96BEA84CF5510291BC8ED12FB32F9E7" box="[924,1100,1579,1604]" italics="true" pageId="5" pageNumber="506">Testudo galeata</emphasis>
<bibRefCitation id="EF8E4B67CF5510291C06ED12FA7EF9E7" author="Schoepff, I. D." box="[1106,1280,1579,1604]" pageId="5" pageNumber="506" refId="ref13074" refString="Schoepff, I. D. (1792) Historia Testudinum iconibus illustrata. Ioannes Iacobus Palm, Erlangen, XII + 136 pp., 31 plates. http: // dx. doi. org / 10.5962 / bhl. title. 5109" type="book" year="1792">Schoepff, 1792</bibRefCitation>
</taxonomicName>
), even though John Edward Gray in his influential works consistently applied the species name
<taxonomicName id="4C1F4D15CF5510291C6FED76FBEFF9CB" baseAuthorityName="Lacepede" baseAuthorityYear="1788" box="[1083,1169,1615,1640]" class="Reptilia" family="Pelomedusidae" genus="Pelomedusa" kingdom="Animalia" order="Testudines" pageId="5" pageNumber="506" phylum="Chordata" rank="species" species="subrufa">
<emphasis id="B96BEA84CF5510291C6FED76FBEFF9CB" box="[1083,1169,1615,1640]" italics="true" pageId="5" pageNumber="506">subrufa</emphasis>
</taxonomicName>
(as
<taxonomicName id="4C1F4D15CF5510291C97ED76FE2BF92F" authority=", Gray 1831: p. 39" authorityName="Gray" authorityPageNumber="39" authorityYear="1831" class="Testudines" family="Chelidae" genus="Hydraspis" higherTaxonomySource="CoL" kingdom="Animalia" order="Pleurodira" pageId="5" pageNumber="506" phylum="Chordata" rank="species" species="subrufa">
<emphasis id="B96BEA84CF5510291C97ED76FAEAF9CB" box="[1219,1428,1615,1640]" italics="true" pageId="5" pageNumber="506">Hydraspis subrufa</emphasis>
,
<bibRefCitation id="EF8E4B67CF55102918C3ED4AFE6CF92F" author="Gray, J. E." box="[151,274,1651,1676]" pageId="5" pageNumber="506" refId="ref12093" refString="Gray, J. E. (1831) Synopsis Reptilium; or Short Descriptions of the Species of Reptiles. Part I. - Cataphracta. Tortoises, Crocodiles, and Enaliosaurians. Treuttel, Wurtz, and Co., London, viii + 85 pp." type="book" year="1831">Gray 1831</bibRefCitation>
: p. 39
</taxonomicName>
or
<taxonomicName id="4C1F4D15CF551029192EED4AFD21F92F" baseAuthorityName="Lacepede" baseAuthorityYear="1788" box="[378,607,1651,1676]" class="Reptilia" family="Pelomedusidae" genus="Pelomedusa" kingdom="Animalia" order="Testudines" pageId="5" pageNumber="506" phylum="Chordata" rank="species" species="subrufa">
<emphasis id="B96BEA84CF551029192EED4AFD21F92F" box="[378,607,1651,1676]" italics="true" pageId="5" pageNumber="506">Pelomedusa subrufa</emphasis>
</taxonomicName>
, e.g.,
<bibRefCitation id="EF8E4B67CF5510291AF0ED4AFC61F92F" author="Gray, J. E." box="[676,799,1651,1676]" pageId="5" pageNumber="506" refId="ref12142" refString="Gray, J. E. (1856 [&quot; 1855 &quot;]) Catalogue of Shield Reptiles in the Collection of the British Museum. Part I. Testudinata (Tortoises). British Museum, London, (1) + 79 + (2) pp., 42 plates. http: // dx. doi. org / 10.5962 / bhl. title. 5491" type="book" year="1856">Gray 1856</bibRefCitation>
: p. 53; 1863: pp. 99100).
</paragraph>
<paragraph id="8BA03696CF5510291893EDAEFDE3F8C7" blockId="5.[151,1437,1507,2036]" pageId="5" pageNumber="506">
The name
<taxonomicName id="4C1F4D15CF551029191BEDAEFD75F913" authorityName="Bonnaterre" authorityYear="1789" box="[335,523,1687,1712]" class="Reptilia" family="Testudinidae" genus="Testudo" kingdom="Animalia" order="Testudines" pageId="5" pageNumber="506" phylum="Chordata" rank="species" species="subrufa">
<emphasis id="B96BEA84CF551029191BEDAEFD75F913" box="[335,523,1687,1712]" italics="true" pageId="5" pageNumber="506">Testudo subrufa</emphasis>
</taxonomicName>
was originally coined by
<bibRefCitation id="EF8E4B67CF5510291B0EEDAEFB5EF913" author="Lacepede" box="[858,1056,1687,1712]" pageId="5" pageNumber="506" refId="ref12474" refString="Lacepede, [B. G.] de (1788) Histoire naturelle des quadrupedes ovipares et des serpens. Tome premier. Hotel de Thou, Paris, (17) + (1, errata) + 651 pp., 1 folded table, 41 plates. http: // dx. doi. org / 10.5962 / bhl. title. 5036" type="book" year="1788">Lacepède (1788)</bibRefCitation>
in his “
<emphasis id="B96BEA84CF5510291CC7EDA1FEE1F971" italics="true" pageId="5" pageNumber="506">Histoire Naturelle des Quadrupèdes Ovipares</emphasis>
”, where a specimen from the Royal Cabinet Paris was described under the name “
<emphasis id="B96BEA84CF5510291D2AED85FE78F95B" italics="true" pageId="5" pageNumber="506">La Roussâtre</emphasis>
” (p. 173). In the “
<emphasis id="B96BEA84CF5510291989EDE6FC8AF954" box="[477,1012,1759,1784]" italics="true" pageId="5" pageNumber="506">Synopsis methodica Quadrupedum oviparorum</emphasis>
” of the same work, a folded table in which binominals were applied to the individual species, Lacepède named this terrapin
<taxonomicName id="4C1F4D15CF5510291C32EC3AFA66F8BF" authorityName="Bonnaterre" authorityYear="1789" box="[1126,1304,1795,1820]" class="Reptilia" family="Testudinidae" genus="Testudo" kingdom="Animalia" order="Testudines" pageId="5" pageNumber="506" phylum="Chordata" rank="species" species="subrufa">
<emphasis id="B96BEA84CF5510291C32EC3AFA66F8BF" box="[1126,1304,1795,1820]" italics="true" pageId="5" pageNumber="506">Testudo subrufa</emphasis>
</taxonomicName>
. According to
<bibRefCitation id="EF8E4B67CF55102918E3EC1EFE0DF8E3" author="Lacepede" box="[183,371,1831,1856]" pageId="5" pageNumber="506" refId="ref12474" refString="Lacepede, [B. G.] de (1788) Histoire naturelle des quadrupedes ovipares et des serpens. Tome premier. Hotel de Thou, Paris, (17) + (1, errata) + 651 pp., 1 folded table, 41 plates. http: // dx. doi. org / 10.5962 / bhl. title. 5036" type="book" year="1788">Lacepède (1788</bibRefCitation>
: p. 173) the terrapin on which he based his description was obtained from Pierre Sonnerat (17451814) and came allegedly from “
<collectingCountry id="F3087606CF5510291A02EC72FDF0F8C7" box="[598,654,1867,1892]" name="India" pageId="5" pageNumber="506">Inde</collectingCountry>
”.
</paragraph>
<paragraph id="8BA03696CF5510291893EC56FC85F857" blockId="5.[151,1437,1507,2036]" pageId="5" pageNumber="506">
This specimen, which has to be identified with the
<typeStatus id="54A48834CF5510291B55EC56FC1DF82B" box="[769,867,1903,1928]" pageId="5" pageNumber="506" type="holotype">holotype</typeStatus>
of
<taxonomicName id="4C1F4D15CF5510291BDCEC56FB88F82B" authority="Lacepede, 1788" authorityName="Lacepede" authorityYear="1788" box="[904,1270,1903,1928]" class="Reptilia" family="Testudinidae" genus="Testudo" kingdom="Animalia" order="Testudines" pageId="5" pageNumber="506" phylum="Chordata" rank="species" species="subrufa">
<emphasis id="B96BEA84CF5510291BDCEC56FB45F82B" box="[904,1083,1903,1928]" italics="true" pageId="5" pageNumber="506">Testudo subrufa</emphasis>
<bibRefCitation id="EF8E4B67CF5510291C15EC56FB88F82B" author="Lacepede" box="[1089,1270,1903,1928]" pageId="5" pageNumber="506" refId="ref12474" refString="Lacepede, [B. G.] de (1788) Histoire naturelle des quadrupedes ovipares et des serpens. Tome premier. Hotel de Thou, Paris, (17) + (1, errata) + 651 pp., 1 folded table, 41 plates. http: // dx. doi. org / 10.5962 / bhl. title. 5036" type="book" year="1788">Lacepède, 1788</bibRefCitation>
</taxonomicName>
, is still present in the collection of the Muséum National dHistoire naturelle (
<bibRefCitation id="EF8E4B67CF5510291B25ECAAFC90F80F" author="Bour, R." box="[881,1006,1939,1964]" pageId="5" pageNumber="506" pagination="531 - 539" refId="ref11305" refString="Bour, R. (1982) Pelomedusa subrufa (Lacepede, 1788), Pelusios subniger (Lacepede, 1788) (Reptilia, Chelonii) et le sejour de Philibert Commerson a Madagascar. Bulletin du Museum National d'Histoire naturelle, 4 e Serie, Section A, 4, 531 - 539." type="journal article" year="1982">Bour 1982</bibRefCitation>
) and was examined by us; tissue for genetic examination was extracted. The type (MNHN 7970) is a shell of an adult terrapin (straight carapacial length
<quantity id="4CE79B73CF55102918C3ECE2FE7FF857" box="[151,257,2011,2036]" metricMagnitude="-1" metricUnit="m" metricValue="1.367" pageId="5" pageNumber="506" unit="cm" value="13.67">13.67 cm</quantity>
) with most epidermal scutes missing. Its plastron is broken (
<figureCitation id="13242A13CF5510291BFCECE2FC93F857" box="[936,1005,2011,2036]" captionStart="FIGURE 2" captionStartId="6.[151,250,954,976]" captionTargetBox="[212,1387,200,929]" captionTargetId="figure-447@6.[179,1408,185,941]" captionTargetPageId="6" captionText="FIGURE 2. Dorsal and ventral view of the holotype of Testudo subrufa Bonnaterre, 1789 (MNHN 7970, straight carapacial length 13.67 cm). Scale bar, 3 cm. Photos: M.D. Hofmeyr." figureDoi="http://doi.org/10.5281/zenodo.4915068" httpUri="https://zenodo.org/record/4915068/files/figure.png" pageId="5" pageNumber="506">Fig. 2</figureCitation>
).
</paragraph>
</subSubSection>
<caption id="DF60661ECF56102A18C3E883FD99FC4C" ID-DOI="http://doi.org/10.5281/zenodo.4915068" ID-Zenodo-Dep="4915068" httpUri="https://zenodo.org/record/4915068/files/figure.png" pageId="6" pageNumber="507" startId="6.[151,250,954,976]" targetBox="[212,1387,200,929]" targetPageId="6">
<paragraph id="8BA03696CF56102A18C3E883FD99FC4C" blockId="6.[151,1435,954,1007]" pageId="6" pageNumber="507">
<emphasis id="B96BEA84CF56102A18C3E883FE6AFC6C" bold="true" box="[151,276,954,976]" pageId="6" pageNumber="507">FIGURE 2.</emphasis>
Dorsal and ventral view of the holotype of
<taxonomicName id="4C1F4D15CF56102A1A8AE883FB45FC73" authority="Bonnaterre, 1789" authorityName="Bonnaterre" authorityYear="1789" box="[734,1083,954,976]" class="Reptilia" family="Testudinidae" genus="Testudo" kingdom="Animalia" order="Testudines" pageId="6" pageNumber="507" phylum="Chordata" rank="species" species="subrufa">
<emphasis id="B96BEA84CF56102A1A8AE883FCFCFC73" box="[734,898,954,976]" italics="true" pageId="6" pageNumber="507">Testudo subrufa</emphasis>
<bibRefCitation id="EF8E4B67CF56102A1BDDE882FB45FC73" author="Bonnaterre" box="[905,1083,954,976]" pageId="6" pageNumber="507" refId="ref11147" refString="Bonnaterre, [P. J.] (1789) Tableau encyclopedique et methodique des trois regnes de la nature. Erpetologie. Panckoucke, Hotel de Thou, Paris, xxviii + 70 + (1) pp., 35 plates. http: // dx. doi. org / 10.5962 / bhl. title. 59326" type="book" year="1789">Bonnaterre, 1789</bibRefCitation>
</taxonomicName>
(MNHN 7970, straight carapacial length 13.67 cm). Scale bar, 3 cm. Photos: M.D. Hofmeyr.
</paragraph>
</caption>
<subSubSection id="C305651DCF56102A1893EF2CFA1FFB1D" pageId="6" pageNumber="507" type="reference_group">
<paragraph id="8BA03696CF56102A1893EF2CFA1FFB1D" blockId="6.[151,1437,1045,1826]" pageId="6" pageNumber="507">
Since the International Commission on Zoological Nomenclature (
<bibRefCitation id="EF8E4B67CF56102A1BE0EF2CFB45FB8D" author="ICZN &amp; International Commission on Zoological Nomenclature" box="[948,1083,1045,1070]" pageId="6" pageNumber="507" pagination="55" refId="ref12417" refString="ICZN [International Commission on Zoological Nomenclature] (2005) Opinion 2104. Bulletin of Zoological Nomenclature, 62, 55." type="journal article" year="2005">ICZN 2005</bibRefCitation>
: Opinion 2104, Case 3226) has ruled that
<bibRefCitation id="EF8E4B67CF56102A1945EF00FE9BFBF1" author="Lacepede" box="[273,485,1081,1106]" pageId="6" pageNumber="507" refId="ref12474" refString="Lacepede, [B. G.] de (1788) Histoire naturelle des quadrupedes ovipares et des serpens. Tome premier. Hotel de Thou, Paris, (17) + (1, errata) + 651 pp., 1 folded table, 41 plates. http: // dx. doi. org / 10.5962 / bhl. title. 5036" type="book" year="1788">Lacepèdes (1788)</bibRefCitation>
<emphasis id="B96BEA84CF56102A19A8EF03FB71FBF3" box="[508,1039,1081,1106]" italics="true" pageId="6" pageNumber="507">Histoire Naturelle des Quadrupèdes Ovipares</emphasis>
” is an unavailable non-binominal work,
<bibRefCitation id="EF8E4B67CF56102A18B6EF67FECAFBD5" author="Bonnaterre" box="[226,436,1117,1142]" pageId="6" pageNumber="507" refId="ref11147" refString="Bonnaterre, [P. J.] (1789) Tableau encyclopedique et methodique des trois regnes de la nature. Erpetologie. Panckoucke, Hotel de Thou, Paris, xxviii + 70 + (1) pp., 35 plates. http: // dx. doi. org / 10.5962 / bhl. title. 59326" type="book" year="1789">Bonnaterre (1789)</bibRefCitation>
became author of
<taxonomicName id="4C1F4D15CF56102A1AC1EF64FC32FBD5" authorityName="Bonnaterre" authorityYear="1789" box="[661,844,1117,1142]" class="Reptilia" family="Testudinidae" genus="Testudo" kingdom="Animalia" order="Testudines" pageId="6" pageNumber="507" phylum="Chordata" rank="species" species="subrufa">
<emphasis id="B96BEA84CF56102A1AC1EF64FC32FBD5" box="[661,844,1117,1142]" italics="true" pageId="6" pageNumber="507">Testudo subrufa</emphasis>
</taxonomicName>
(and of other names) by re-publishing Lacepèdes descriptions using consistently binominal scientific names. The
<typeStatus id="54A48834CF56102A1B3AEFBBFCE0FB39" box="[878,926,1154,1178]" pageId="6" pageNumber="507">type</typeStatus>
status of the terrapin described by
<bibRefCitation id="EF8E4B67CF56102A1D65EFB8FF9FFB1D" author="Lacepede" pageId="6" pageNumber="507" refId="ref12474" refString="Lacepede, [B. G.] de (1788) Histoire naturelle des quadrupedes ovipares et des serpens. Tome premier. Hotel de Thou, Paris, (17) + (1, errata) + 651 pp., 1 folded table, 41 plates. http: // dx. doi. org / 10.5962 / bhl. title. 5036" type="book" year="1788">Lacepède (1788)</bibRefCitation>
and re-described by
<bibRefCitation id="EF8E4B67CF56102A1998EF9FFDEBFB1D" author="Bonnaterre" box="[460,661,1189,1214]" pageId="6" pageNumber="507" refId="ref11147" refString="Bonnaterre, [P. J.] (1789) Tableau encyclopedique et methodique des trois regnes de la nature. Erpetologie. Panckoucke, Hotel de Thou, Paris, xxviii + 70 + (1) pp., 35 plates. http: // dx. doi. org / 10.5962 / bhl. title. 59326" type="book" year="1789">Bonnaterre (1789</bibRefCitation>
: p. 28) remains unaffected by this transfer of formal authorship.
</paragraph>
</subSubSection>
<subSubSection id="C305651DCF56102A1893EFF3FB2FF882" pageId="6" pageNumber="507" type="materials_examined">
<paragraph id="8BA03696CF56102A1893EFF3FBCCF985" blockId="6.[151,1437,1045,1826]" pageId="6" pageNumber="507">
It is obvious that the original type locality (“
<collectingCountry id="F3087606CF56102A1A81EFF0FC72FB41" box="[725,780,1225,1250]" name="India" pageId="6" pageNumber="507">Inde</collectingCountry>
”) of
<taxonomicName id="4C1F4D15CF56102A1B18EFF0FB7DFB41" authorityName="Bonnaterre" authorityYear="1789" box="[844,1027,1225,1250]" class="Reptilia" family="Testudinidae" genus="Testudo" kingdom="Animalia" order="Testudines" pageId="6" pageNumber="507" phylum="Chordata" rank="species" species="subrufa">
<emphasis id="B96BEA84CF56102A1B18EFF0FB7DFB41" box="[844,1027,1225,1250]" italics="true" pageId="6" pageNumber="507">Testudo subrufa</emphasis>
</taxonomicName>
is in error. In accordance with the information provided by
<bibRefCitation id="EF8E4B67CF56102A19E6EFD4FD0CFAA5" author="Lacepede" box="[434,626,1261,1286]" pageId="6" pageNumber="507" refId="ref12474" refString="Lacepede, [B. G.] de (1788) Histoire naturelle des quadrupedes ovipares et des serpens. Tome premier. Hotel de Thou, Paris, (17) + (1, errata) + 651 pp., 1 folded table, 41 plates. http: // dx. doi. org / 10.5962 / bhl. title. 5036" type="book" year="1788">Lacepède (1788)</bibRefCitation>
,
<bibRefCitation id="EF8E4B67CF56102A1A2BEFD7FC56FAA5" author="Mertens, R." box="[639,808,1261,1286]" pageId="6" pageNumber="507" pagination="139 - 142" refId="ref12650" refString="Mertens, R. (1937) Bemerkungen uber die Rassen von Pelomedusa subrufa (La Cepede). Zoologischer Anzeiger, 117, 139 - 142." type="journal article" year="1937">Mertens (1937</bibRefCitation>
: p. 139) and
<bibRefCitation id="EF8E4B67CF56102A1BE9EFD4FB02FAA5" author="Loveridge, A." box="[957,1148,1261,1286]" pageId="6" pageNumber="507" pagination="467 - 524" refId="ref12616" refString="Loveridge, A. (1941) Revision of the African terrapin [sic] of the family Pelomedusidae. Bulletin of the Museum of Comparative Zoology, 88, 467 - 524." type="journal article" year="1941">Loveridge (1941</bibRefCitation>
: p. 470) believed that the type specimen of
<taxonomicName id="4C1F4D15CF56102A1935EE28FD6BFA89" authorityName="Bonnaterre" authorityYear="1789" box="[353,533,1297,1322]" class="Reptilia" family="Testudinidae" genus="Testudo" kingdom="Animalia" order="Testudines" pageId="6" pageNumber="507" phylum="Chordata" rank="species" species="subrufa">
<emphasis id="B96BEA84CF56102A1935EE28FD6BFA89" box="[353,533,1297,1322]" italics="true" pageId="6" pageNumber="507">Testudo subrufa</emphasis>
</taxonomicName>
originates from Sonnerat, who is known to have collected at the Cape of Good Hope. Therefore, they identified the type locality with the Cape. However, based on an extensive discussion of historical details,
<bibRefCitation id="EF8E4B67CF56102A190AEE63FE97FAD1" author="Bour, R." box="[350,489,1369,1394]" pageId="6" pageNumber="507" pagination="531 - 539" refId="ref11305" refString="Bour, R. (1982) Pelomedusa subrufa (Lacepede, 1788), Pelusios subniger (Lacepede, 1788) (Reptilia, Chelonii) et le sejour de Philibert Commerson a Madagascar. Bulletin du Museum National d'Histoire naturelle, 4 e Serie, Section A, 4, 531 - 539." type="journal article" year="1982">Bour (1982)</bibRefCitation>
concluded that the
<typeStatus id="54A48834CF56102A1A93EE60FC57FAD1" box="[711,809,1369,1394]" pageId="6" pageNumber="507" type="holotype">holotype</typeStatus>
of
<taxonomicName id="4C1F4D15CF56102A1B04EE60FB7DFAD1" authorityName="Bonnaterre" authorityYear="1789" box="[848,1027,1369,1394]" class="Reptilia" family="Testudinidae" genus="Testudo" kingdom="Animalia" order="Testudines" pageId="6" pageNumber="507" phylum="Chordata" rank="species" species="subrufa">
<emphasis id="B96BEA84CF56102A1B04EE60FB7DFAD1" box="[848,1027,1369,1394]" italics="true" pageId="6" pageNumber="507">Testudo subrufa</emphasis>
</taxonomicName>
was most probably not collected by Sonnerat, but by Philibert Commerson (17271773), who stayed on
<collectingCountry id="F3087606CF56102A1B91EE44FB32FA35" box="[965,1100,1405,1430]" name="Madagascar" pageId="6" pageNumber="507">Madagascar</collectingCountry>
in 1770. Some drawings of Commersons Malagasy specimens are still present in the Central Library of the Paris Museum.
<bibRefCitation id="EF8E4B67CF56102A1D5AEE9BFAE2FA19" author="Bour, R." box="[1294,1436,1441,1466]" pageId="6" pageNumber="507" pagination="531 - 539" refId="ref11305" refString="Bour, R. (1982) Pelomedusa subrufa (Lacepede, 1788), Pelusios subniger (Lacepede, 1788) (Reptilia, Chelonii) et le sejour de Philibert Commerson a Madagascar. Bulletin du Museum National d'Histoire naturelle, 4 e Serie, Section A, 4, 531 - 539." type="journal article" year="1982">Bour (1982)</bibRefCitation>
identified one of these drawings with the
<typeStatus id="54A48834CF56102A1ACBEEFCFC7FFA7D" box="[671,769,1477,1502]" pageId="6" pageNumber="507" type="holotype">holotype</typeStatus>
and designated “Taolañaro (Fort-Dauphin), République Malagasy [
<collectingCountry id="F3087606CF56102A1947EED0FEDCF9A1" box="[275,418,1513,1538]" name="Madagascar" pageId="6" pageNumber="507">Madagascar</collectingCountry>
]” (p. 535) as type locality of
<taxonomicName id="4C1F4D15CF56102A1AA1EED0FCD7F9A1" authorityName="Bonnaterre" authorityYear="1789" box="[757,937,1513,1538]" class="Reptilia" family="Testudinidae" genus="Testudo" kingdom="Animalia" order="Testudines" pageId="6" pageNumber="507" phylum="Chordata" rank="species" species="subrufa">
<emphasis id="B96BEA84CF56102A1AA1EED0FCD7F9A1" box="[757,937,1513,1538]" italics="true" pageId="6" pageNumber="507">Testudo subrufa</emphasis>
</taxonomicName>
. However, probably due to oversight,
<bibRefCitation id="EF8E4B67CF56102A1D30EED3FFA2F985" author="Bour, R." pageId="6" pageNumber="507" pagination="54 - 80" refId="ref11361" refString="Bour, R. (1985) Les tortues terrestres et d'eau douce de Madagascar et des iles voisines. Madagascar Recherches Scientifique, 18, 54 - 80." type="journal article" year="1985">Bour (1985</bibRefCitation>
: p. 56) endorsed later the Cape of Good Hope as type locality of
<taxonomicName id="4C1F4D15CF56102A1BE0ED34FB18F985" authorityName="Bonnaterre" authorityYear="1789" box="[948,1126,1549,1574]" class="Reptilia" family="Testudinidae" genus="Testudo" kingdom="Animalia" order="Testudines" pageId="6" pageNumber="507" phylum="Chordata" rank="species" species="subrufa">
<emphasis id="B96BEA84CF56102A1BE0ED34FB18F985" box="[948,1126,1549,1574]" italics="true" pageId="6" pageNumber="507">Testudo subrufa</emphasis>
</taxonomicName>
again.
</paragraph>
<paragraph id="8BA03696CF56102A1893ED08FB2FF882" blockId="6.[151,1437,1045,1826]" pageId="6" pageNumber="507">
All of our efforts to generate mtDNA sequences from the
<typeStatus id="54A48834CF56102A1BDCED08FC94F9E9" box="[904,1002,1585,1610]" pageId="6" pageNumber="507" type="holotype">holotype</typeStatus>
resulted only in contaminant fungal sequences. Therefore, in the absence of other evidence, we accept the type locality Taolañaro (
<collectingCountry id="F3087606CF56102A1CB6ED6CFA11F9CD" box="[1250,1391,1621,1646]" name="Madagascar" pageId="6" pageNumber="507">Madagascar</collectingCountry>
), as suggested by
<bibRefCitation id="EF8E4B67CF56102A1962ED43FEB9F931" author="Bour, R." box="[310,455,1657,1682]" pageId="6" pageNumber="507" pagination="531 - 539" refId="ref11305" refString="Bour, R. (1982) Pelomedusa subrufa (Lacepede, 1788), Pelusios subniger (Lacepede, 1788) (Reptilia, Chelonii) et le sejour de Philibert Commerson a Madagascar. Bulletin du Museum National d'Histoire naturelle, 4 e Serie, Section A, 4, 531 - 539." type="journal article" year="1982">Bour (1982)</bibRefCitation>
. Consequently, the name
<taxonomicName id="4C1F4D15CF56102A1AA7ED40FB07F931" authority="Bonnaterre, 1789" authorityName="Bonnaterre" authorityYear="1789" box="[755,1145,1657,1682]" class="Reptilia" family="Testudinidae" genus="Testudo" kingdom="Animalia" order="Testudines" pageId="6" pageNumber="507" phylum="Chordata" rank="species" species="subrufa">
<emphasis id="B96BEA84CF56102A1AA7ED40FCD7F931" box="[755,937,1657,1682]" italics="true" pageId="6" pageNumber="507">Testudo subrufa</emphasis>
<bibRefCitation id="EF8E4B67CF56102A1BE7ED43FB07F931" author="Bonnaterre" box="[947,1145,1657,1682]" pageId="6" pageNumber="507" refId="ref11147" refString="Bonnaterre, [P. J.] (1789) Tableau encyclopedique et methodique des trois regnes de la nature. Erpetologie. Panckoucke, Hotel de Thou, Paris, xxviii + 70 + (1) pp., 35 plates. http: // dx. doi. org / 10.5962 / bhl. title. 59326" type="book" year="1789">Bonnaterre, 1789</bibRefCitation>
</taxonomicName>
refers to lineage VIII of
<bibRefCitation id="EF8E4B67CF56102A18C3EDA4FE9EF915" 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="[151,480,1693,1718]" pageId="6" pageNumber="507" pagination="428 - 440" refId="ref13781" 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="B96BEA84CF56102A1906EDA7FEF2F915" box="[338,396,1693,1718]" italics="true" pageId="6" pageNumber="507">et al.</emphasis>
(2010)
</bibRefCitation>
, which is known to occur in
<collectingCountry id="F3087606CF56102A1B73EDA4FCCEF915" box="[807,944,1693,1718]" name="Madagascar" pageId="6" pageNumber="507">Madagascar</collectingCountry>
,
<collectingCountry id="F3087606CF56102A1BE8EDA7FB53F916" box="[956,1069,1694,1717]" name="Botswana" pageId="6" pageNumber="507">Botswana</collectingCountry>
,
<collectingCountry id="F3087606CF56102A1C6DEDA4FF9AF979" name="Democratic Republic of the Congo" pageId="6" pageNumber="507">the Democratic Republic of the Congo</collectingCountry>
,
<collectingCountry id="F3087606CF56102A18A6EDF8FE39F979" box="[242,327,1729,1754]" name="Malawi" pageId="6" pageNumber="507">Malawi</collectingCountry>
and
<collectingCountry id="F3087606CF56102A19D0EDF8FE9BF979" box="[388,485,1729,1754]" name="Namibia" pageId="6" pageNumber="507">Namibia</collectingCountry>
(
<figureCitation id="13242A13CF56102A19A3EDF8FD45F979" box="[503,571,1729,1754]" captionStart="FIGURE 1" captionStartId="5.[151,250,1100,1122]" captionTargetBox="[157,1429,201,1075]" captionTargetId="figure-427@5.[151,1436,195,1080]" captionTargetPageId="5" captionText="FIGURE 1. Maximum Likelihood tree for Pelomedusa, based on 1848 bp of mitochondrial DNA (12S, cyt b, ND4+tRNAs), rooted with Pelusios sinuatus. The tree topology completely matches the one inferred by Bayesian analyses. Numbers along nodes indicate bootstrap support and Bayesian posterior probabilities greater than 50 or 0.95, respectively (not shown for some terminal clades with short branches). Sample codes are voucher numbers; see Table S1 for explanation. Name-bearing types (red) and paralectotypes or topotypic samples (blue) highlighted. On the right, mitochondrial lineages of Vargas-Ramírez et al. (2010) indicated. No topotypic material of Pelomedusa galeata orangensis Hewitt, 1935 could be sequenced. However, the lost holotype is thought to come from the vicinity of Kimberley (South Africa) and all South African samples belong to lineage IX, so it can be concluded that Pelomedusa galeata orangensis represents the same lineage. Inset: Helmeted terrapin from the Addo Elephant National Park, South Africa. Photo: M. Gregorič." figureDoi="http://doi.org/10.5281/zenodo.4915066" httpUri="https://zenodo.org/record/4915066/files/figure.png" pageId="6" pageNumber="507">Fig. 1</figureCitation>
). The Malagasy populations are thought to be introduced from continental Africa (
<bibRefCitation id="EF8E4B67CF56102A18B9EDDCFD59F95D" 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="[237,551,1765,1790]" pageId="6" pageNumber="507" pagination="428 - 440" refId="ref13781" 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="B96BEA84CF56102A19FFEDDFFE9AF95D" box="[427,484,1765,1790]" italics="true" pageId="6" pageNumber="507">et al.</emphasis>
2010
</bibRefCitation>
;
<bibRefCitation id="EF8E4B67CF56102A1A67EDDFFD84F95D" author="Wong, R. A. &amp; Fong, J. J. &amp; Papenfuss, T. J." box="[563,762,1765,1790]" pageId="6" pageNumber="507" pagination="575 - 585" refId="ref13970" refString="Wong, R. A., Fong, J. J. &amp; Papenfuss, T. J. (2010) Phylogeography of the African helmeted terrapin, Pelomedusa subrufa: genetic structure, dispersal, and human introduction. Proceedings of the California Academy of Sciences, 61, 575 - 585." type="journal article" year="2010">
Wong
<emphasis id="B96BEA84CF56102A1A28EDDFFDC8F95D" box="[636,694,1765,1790]" italics="true" pageId="6" pageNumber="507">et al.</emphasis>
2010
</bibRefCitation>
). If lineage VIII is deemed to be taxonomically distinct, the name
<taxonomicName id="4C1F4D15CF56102A188EEC30FDE2F881" authority="(Bonnaterre, 1789)" baseAuthorityName="Bonnaterre" baseAuthorityYear="1789" box="[218,668,1801,1826]" class="Reptilia" family="Pelomedusidae" genus="Pelomedusa" kingdom="Animalia" order="Testudines" pageId="6" pageNumber="507" phylum="Chordata" rank="species" species="subrufa">
<emphasis id="B96BEA84CF56102A188EEC30FEBEF881" box="[218,448,1801,1826]" italics="true" pageId="6" pageNumber="507">Pelomedusa subrufa</emphasis>
(
<bibRefCitation id="EF8E4B67CF56102A199AEC30FDEAF881" author="Bonnaterre" box="[462,660,1801,1826]" pageId="6" pageNumber="507" refId="ref11147" refString="Bonnaterre, [P. J.] (1789) Tableau encyclopedique et methodique des trois regnes de la nature. Erpetologie. Panckoucke, Hotel de Thou, Paris, xxviii + 70 + (1) pp., 35 plates. http: // dx. doi. org / 10.5962 / bhl. title. 59326" type="book" year="1789">Bonnaterre, 1789</bibRefCitation>
)
</taxonomicName>
will have to be restricted to this taxon.
</paragraph>
</subSubSection>
</treatment>
</document>