treatments-xml/data/CE/95/59/CE9559CB92277D26D0C32C0A88986CDD.xml
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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.859.33634" ID-GBIF-Dataset="1ebfdc7c-50af-4535-bee7-2fb58e3f2b81" ID-PMC="PMC6616056" ID-Pensoft-Pub="1313-2970-859-131" ID-PubMed="31327926" ModsDocAuthor="" ModsDocDate="2019" ModsDocID="1313-2970-859-131" ModsDocOrigin="ZooKeys 859" ModsDocTitle="Call a spade a spade: taxonomy and distribution of Pelobates, with description of a new Balkan endemic" checkinTime="1562137562851" checkinUser="pensoft" docAuthor="Dufresnes, Christophe, Strachinis, Ilias, Tzoras, Elias, Litvinchuk, Spartak N. &amp; Denoel, Mathieu" docDate="2019" docId="CE9559CB92277D26D0C32C0A88986CDD" docLanguage="en" docName="ZooKeys 859: 131-158" docOrigin="ZooKeys 859" docSource="http://dx.doi.org/10.3897/zookeys.859.33634" docTitle="Pelobates balcanicus Karaman 1928" docType="treatment" docVersion="5" lastPageNumber="144" masterDocId="FFE13C3F7A123710D3023035D907FF8A" masterDocTitle="Call a spade a spade: taxonomy and distribution of Pelobates, with description of a new Balkan endemic" masterLastPageNumber="158" masterPageNumber="131" pageNumber="143" updateTime="1668167462434" updateUser="ExternalLinkService">
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<mods:titleInfo>
<mods:title>Call a spade a spade: taxonomy and distribution of Pelobates, with description of a new Balkan endemic</mods:title>
</mods:titleInfo>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Dufresnes, Christophe</mods:namePart>
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<mods:namePart>Strachinis, Ilias</mods:namePart>
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<mods:namePart>Tzoras, Elias</mods:namePart>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Litvinchuk, Spartak N.</mods:namePart>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Denoel, Mathieu</mods:namePart>
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<mods:date>2019</mods:date>
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<mods:number>859</mods:number>
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<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.859.33634</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-859-131</mods:identifier>
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<treatment ID-GBIF-Taxon="158498309" LSID="urn:lsid:plazi:treatment:CE9559CB92277D26D0C32C0A88986CDD" httpUri="http://treatment.plazi.org/id/CE9559CB92277D26D0C32C0A88986CDD" lastPageId="13" lastPageNumber="144" pageId="12" pageNumber="143">
<subSubSection pageId="12" pageNumber="143" type="nomenclature">
<paragraph pageId="12" pageNumber="143">
<taxonomicName authority="Karaman, 1928" authorityName="Karaman" authorityYear="1928" class="Amphibia" family="Pelobatidae" genus="Pelobates" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pelobates balcanicus" order="Anura" pageId="12" pageNumber="143" phylum="Chordata" rank="species" species="balcanicus">Pelobates balcanicus Karaman, 1928</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection pageId="12" pageNumber="143" type="diagnosis">
<paragraph pageId="12" pageNumber="143">Diagnosis.</paragraph>
<paragraph pageId="12" pageNumber="143">
Resembling
<taxonomicName lsidName="P. syriacus" pageId="12" pageNumber="143" rank="species" species="syriacus">P. syriacus</taxonomicName>
with which it was previously considered a synonym (
<bibRefCitation author="Frost, D" journalOrPublisher="Annalen des Naturhistorischen Museums in Wien" pageId="20" pageNumber="151" url="http://research.amnh.org/herpetology/amphibia/index.html" year="2019">Frost 2019</bibRefCitation>
). Large toad with whitish metatarsal spades and a flat skull. Sexes of similar size (Fig. 2). Various motifs with gray, yellow or greenish colors, but rarely brown (unlike the sympatric
<taxonomicName lsidName="P. fuscus" pageId="12" pageNumber="143" rank="species" species="fuscus">P. fuscus</taxonomicName>
, P.
<normalizedToken originalValue="Székely">Szekely</normalizedToken>
pers. comm.); frequently specked with orange dots, sometimes heavily (perhaps more than in
<taxonomicName lsidName="P. syriacus" pageId="12" pageNumber="143" rank="species" species="syriacus">P. syriacus</taxonomicName>
) (Fig. 4). Based on 25 biometric characters,
<bibRefCitation author="Ugurtas, IH" journalOrPublisher="Israel Journal of Zoology" pageId="24" pageNumber="155" pagination="13 - 32" title="Morphological differentiation of eastern spadefoot toad (Pelobatessyriacus) populations." volume="48" year="2002">
<normalizedToken originalValue="Uğurtas">Ugurtas</normalizedToken>
et al. (2002)
</bibRefCitation>
showed that the
<taxonomicName lsidName="P. balcanicus" pageId="12" pageNumber="143" rank="species" species="balcanicus">P. balcanicus</taxonomicName>
populations from the Balkans are morphologically very variable and differentiated from Asia Minor (
<taxonomicName lsidName="P. syriacus" pageId="12" pageNumber="143" rank="species" species="syriacus">P. syriacus</taxonomicName>
); yet
<taxonomicName lsidName="P. syriacus" pageId="12" pageNumber="143" rank="species" species="syriacus">P. syriacus</taxonomicName>
populations from European Turkey (Edirne, genetically confirmed by
<bibRefCitation author="Dufresnes, C" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="20" pageNumber="151" url="https://doi.org/10.1111/mec.15133" year="2019 b">Dufresnes et al. 2019b</bibRefCitation>
) and southeastern Bulgaria (Primorsko) grouped with
<taxonomicName lsidName="P. balcanicus" pageId="12" pageNumber="143" rank="species" species="balcanicus">P. balcanicus</taxonomicName>
(
<bibRefCitation author="Ugurtas, IH" journalOrPublisher="Israel Journal of Zoology" pageId="24" pageNumber="155" pagination="13 - 32" title="Morphological differentiation of eastern spadefoot toad (Pelobatessyriacus) populations." volume="48" year="2002">
<normalizedToken originalValue="Uğurtas">Ugurtas</normalizedToken>
et al. 2002
</bibRefCitation>
).
<bibRefCitation pageId="12" pageNumber="143">
<normalizedToken originalValue="Roček">Rocek</normalizedToken>
(1981)
</bibRefCitation>
only found one cranial difference: the processus posterior parasphenoidei is present in
<taxonomicName lsidName="P. syriacus" pageId="12" pageNumber="143" rank="species" species="syriacus">P. syriacus</taxonomicName>
but not developed in
<taxonomicName lsidName="P. balcanicus" pageId="12" pageNumber="143" rank="species" species="balcanicus">P. balcanicus</taxonomicName>
. Morphometric assessments associated to genetic data are needed. Based on populations of
<taxonomicName lsidName="P. balcanicus" pageId="12" pageNumber="143" rank="subSpecies" species="balcanicus" subSpecies="balcanicus">P. balcanicus balcanicus</taxonomicName>
, average SVL = 67 mm (48-100 mm) for females (n = 16 populations) and 68 mm (46-94 mm) for males (n = 15 populations) (Suppl. material 1, Table S1; Fig. 2). The karyotype (
<taxonomicName lsidName="P. b." pageId="12" pageNumber="143" rank="subSpecies" species="b." subSpecies="balcanicus">P. b. balcanicus</taxonomicName>
) consists of six large and seven small pairs of two-armed chromosomes. NORs (secondary constrictions) are in the short arm of pair 7 (
<bibRefCitation pageId="12" pageNumber="143">Belcheva et al. 1977</bibRefCitation>
). The nuclear DNA content (calculated from flow cytometry) averages 7.9 pg (
<bibRefCitation author="Litvinchuk, SN" journalOrPublisher="Organisms Diversity and Evolution," pageId="21" pageNumber="152" pagination="433 - 451" title="Phylogeographic patterns of genetic diversity in the common spadefoot toad, Pelobatesfuscus (Anura: Pelobatidae), reveals evolutionary history, postglacial range expansion and secondary contact." url="https://doi.org/10.1007/s13127-013-0127-5" volume="13" year="2013">Litvinchuk et al. 2013</bibRefCitation>
; data from
<taxonomicName lsidName="P. b." pageId="12" pageNumber="143" rank="subSpecies" species="b." subSpecies="balcanicus">P. b. balcanicus</taxonomicName>
). As shown in Table 1,
<taxonomicName lsidName="P. balcanicus" pageId="12" pageNumber="143" rank="species" species="balcanicus">P. balcanicus</taxonomicName>
differs from
<taxonomicName lsidName="P. syriacus" pageId="12" pageNumber="143" rank="species" species="syriacus">P. syriacus</taxonomicName>
by ~7.4% at mtDNA and ~0.31% at nuclear DNA (
<bibRefCitation author="Dufresnes, C" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="20" pageNumber="151" url="https://doi.org/10.1111/mec.15133" year="2019 b">Dufresnes et al. 2019b</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="12" pageNumber="143" type="taxonomy">
<paragraph pageId="12" pageNumber="143">Taxonomy.</paragraph>
<paragraph pageId="12" pageNumber="143">
Originally described as a subspecies of the Eastern spadefoot,
<taxonomicName class="Amphibia" family="Pelobatidae" genus="Pelobates" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pelobates syriacus" order="Anura" pageId="12" pageNumber="143" phylum="Chordata" rank="subSpecies" species="syriacus" subSpecies="balcanicus">Pelobates syriacus balcanicus</taxonomicName>
Karaman, 1928; type locality: Dojran Lake, North Macedonia; type(s): most likely include the skeleton described by
<bibRefCitation author="Karaman, S" journalOrPublisher="Bulletin de la Societe Scientifique de Skoplje, Section des Sciences Naturelles" pageId="21" pageNumber="152" pagination="129 - 143" title="III Prilog herpetologii Jugoslavije - Contribution a l'Herpetologie de la Jugoslavija." volume="4" year="1928">Karaman (1928)</bibRefCitation>
, deposited at MMNH (Skopje, North Macedonia), but destroyed in an earthquake in 1963 (V. Sidorovska pers. comm.); the MMNH currently hosts one specimen from the type locality, MMNH-A-699 (collected in 2001). This taxon represents a distinct species from
<taxonomicName lsidName="P. syriacus" pageId="12" pageNumber="143" rank="species" species="syriacus">P. syriacus</taxonomicName>
, due its old divergence (&gt;6 My) and the absence of contemporary introgression at their area of contact in European Turkey, consistent with advanced reproductive isolation (
<bibRefCitation author="Dufresnes, C" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="20" pageNumber="151" url="https://doi.org/10.1111/mec.15133" year="2019 b">Dufresnes et al. 2019b</bibRefCitation>
). Therefore, we herein remove
<taxonomicName lsidName="P. balcanicus" pageId="12" pageNumber="143" rank="species" species="balcanicus">P. balcanicus</taxonomicName>
from its previous synonymy with
<taxonomicName lsidName="P. syriacus" pageId="12" pageNumber="143" rank="species" species="syriacus">P. syriacus</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection lastPageId="13" lastPageNumber="144" pageId="12" pageNumber="143" type="distribution">
<paragraph pageId="12" pageNumber="143">Distribution.</paragraph>
<paragraph lastPageId="13" lastPageNumber="144" pageId="12" pageNumber="143">
<taxonomicName class="Amphibia" family="Pelobatidae" genus="Pelobates" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pelobates balcanicus" order="Anura" pageId="12" pageNumber="143" phylum="Chordata" rank="species" species="balcanicus">Pelobates balcanicus</taxonomicName>
is restricted to the Balkan Peninsula, 0-920 m a.s.l. (
<bibRefCitation author="Dzukic, G" journalOrPublisher="Zeitschrift fur Feldherpetologie" pageId="20" pageNumber="151" pagination="61 - 78" title="Contemporary chorology of the spadefoot toads (Pelobates spp.) in the Balkan Peninsula." volume="15" year="2008">
<normalizedToken originalValue="Džukić">Dzukic</normalizedToken>
et al. 2008
</bibRefCitation>
) (Fig. 1). In the north, it is present in northern Serbia and northwestern Romania. It follows the Danube River from Serbia to the Black Sea in Romania (
<bibRefCitation author="Szekely, P" journalOrPublisher="Herpetology Notes" pageId="23" pageNumber="154" pagination="481 - 484" title="Range extension for the Eastern spadefoot toad Pelobatessyriacus (Boettger, 1889) (Anura: Pelobatidae)." volume="6" year="2013">
<normalizedToken originalValue="Székely">Szekely</normalizedToken>
et al. 2013
</bibRefCitation>
;
<bibRefCitation author="Ţeran, L" journalOrPublisher="North-Western Journal of Zoology" pageId="23" pageNumber="154" pagination="179 - 182" title="New data on the distribution of Eastern spadefoot toad Pelobatessyriacus Boettger, 1889 (Anura: Pelobatidae) in the Pannonian Plain." volume="13" year="2017">
<normalizedToken originalValue="Ţeran">Ţeran</normalizedToken>
et al. 2017
</bibRefCitation>
). There are yet some possible gaps along the Danube (e.g. around the Iron Gate:
<bibRefCitation author="Vukov, T" journalOrPublisher="Bulletin of the Natural History Museum" pageId="24" pageNumber="155" pagination="90 - 112" title="Amphibians in Serbia: distribution and diversity patterns." url="https://doi.org/10.5937/bnhmb1306090V" volume="6" year="2013">Vukov et al. 2013</bibRefCitation>
;
<bibRefCitation author="Ţeran, L" journalOrPublisher="North-Western Journal of Zoology" pageId="23" pageNumber="154" pagination="179 - 182" title="New data on the distribution of Eastern spadefoot toad Pelobatessyriacus Boettger, 1889 (Anura: Pelobatidae) in the Pannonian Plain." volume="13" year="2017">
<normalizedToken originalValue="Ţeran">Ţeran</normalizedToken>
et al. 2017
</bibRefCitation>
). In the north-west, the Great Morova River in Serbia marks its western margin (
<bibRefCitation author="Dzukic, G" journalOrPublisher="Zeitschrift fur Feldherpetologie" pageId="20" pageNumber="151" pagination="61 - 78" title="Contemporary chorology of the spadefoot toads (Pelobates spp.) in the Balkan Peninsula." volume="15" year="2008">
<normalizedToken originalValue="Džukić">Dzukic</normalizedToken>
et al. 2008
</bibRefCitation>
). Northern ranges are currently disconnected from the southern populations (
<bibRefCitation author="Vukov, T" journalOrPublisher="Bulletin of the Natural History Museum" pageId="24" pageNumber="155" pagination="90 - 112" title="Amphibians in Serbia: distribution and diversity patterns." url="https://doi.org/10.5937/bnhmb1306090V" volume="6" year="2013">Vukov et al. 2013</bibRefCitation>
) of North Macedonia, eastern Albania (a single location), south-west Bulgaria (Strimon River), and Greece (
<bibRefCitation author="Dzukic, G" journalOrPublisher="Zeitschrift fur Feldherpetologie" pageId="20" pageNumber="151" pagination="61 - 78" title="Contemporary chorology of the spadefoot toads (Pelobates spp.) in the Balkan Peninsula." volume="15" year="2008">
<normalizedToken originalValue="Džukić">Dzukic</normalizedToken>
et al. 2008
</bibRefCitation>
;
<bibRefCitation author="Mollov, I" journalOrPublisher="Scientific Researches of the Union of Scientists in Bulgaria-Plovdiv, Series B Natural Sciences and the Humanities" pageId="22" pageNumber="153" pagination="132 - 136" title="New data on the distribution of the Syrian Spadefoot (Pelobatessyriacusbalcanicus Karaman, 1928) in Bulgaria." volume="8" year="2006">Mollov et al. 2006</bibRefCitation>
;
<bibRefCitation author="Szabolcs, IM" journalOrPublisher="NorthWestern Journal of Zoology" pageId="23" pageNumber="154" pagination="175 - 176" title="First record of the eastern spadefoot toad (Pelobatessyriacus Boettger, 1889) in Albania." volume="13" year="2017">Szabolcs and Mizsei 2017</bibRefCitation>
). In the 1980s,
<bibRefCitation pageId="12" pageNumber="143">Sofianidou (2012)</bibRefCitation>
reported the species along the western coastline of the Adriatic Sea and the northern coastline of the Gulf of Corinth (Greece), but there is no recent observation in this region. Elsewhere
<pageBreakToken pageId="13" pageNumber="144" start="start">in</pageBreakToken>
Greece, it is present in Peloponnese (
<taxonomicName lsidName="P. balcanicus" pageId="13" pageNumber="144" rank="subSpecies" species="balcanicus" subSpecies="chloeae">P. balcanicus chloeae</taxonomicName>
ssp. nov., see below), in the eastern parts of the mainland, and along the Aegean Sea shores, from Sterea Ellas to the Evros River, until it reaches
<taxonomicName lsidName="P. syriacus" pageId="13" pageNumber="144" rank="species" species="syriacus">P. syriacus</taxonomicName>
in Thrace (
<bibRefCitation author="Dzukic, G" journalOrPublisher="Zeitschrift fur Feldherpetologie" pageId="20" pageNumber="151" pagination="61 - 78" title="Contemporary chorology of the spadefoot toads (Pelobates spp.) in the Balkan Peninsula." volume="15" year="2008">
<normalizedToken originalValue="Džukić">Dzukic</normalizedToken>
et al. 2008
</bibRefCitation>
; Sofianidou, 2012). The spadefoots known from the Maritsa (Evros) River in southern Bulgaria, and along the western coasts of the Black Sea, may correspond to
<taxonomicName lsidName="P. syriacus" pageId="13" pageNumber="144" rank="species" species="syriacus">P. syriacus</taxonomicName>
(Stojanov et al. 2011;
<bibRefCitation author="Dufresnes, C" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="20" pageNumber="151" url="https://doi.org/10.1111/mec.15133" year="2019 b">Dufresnes et al. 2019b</bibRefCitation>
). IUCN status: Not Evaluated; previously included in
<taxonomicName lsidName="P. syriacus" pageId="13" pageNumber="144" rank="species" species="syriacus">P. syriacus</taxonomicName>
assessment.
</paragraph>
</subSubSection>
<subSubSection pageId="13" pageNumber="144" type="diversity">
<paragraph pageId="13" pageNumber="144">Diversity.</paragraph>
<paragraph pageId="13" pageNumber="144">
Using mtDNA and genomic data,
<bibRefCitation author="Dufresnes, C" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="20" pageNumber="151" url="https://doi.org/10.1111/mec.15133" year="2019 b">Dufresnes et al. (2019b)</bibRefCitation>
evidenced a Pleistocene split (~2 My) for spadefoots from the Peloponnese (
<taxonomicName lsidName="P. balcanicus" pageId="13" pageNumber="144" rank="subSpecies" species="balcanicus" subSpecies="chloeae">P. balcanicus chloeae</taxonomicName>
ssp. nov.). In the rest of the range, at least three glacial lineages (&lt;1 My) were identified: a first one in the eastern ranges, from the Carpathians to the Black Sea and as south as Greek Thrace; a second one in western ranges from Serbia to northern Greece; and a third one on the coastal island of Evia (north-east of Peloponnese). The eastern and western lineages widely admix. Populations from central Greece are yet to be examined.
</paragraph>
</subSubSection>
</treatment>
</document>