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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="44D1003976A37083B5FAB001D3340736" docLanguage="en" docName="ZooKeys 859: 131-158" docOrigin="ZooKeys 859" docSource="http://dx.doi.org/10.3897/zookeys.859.33634" docTitle="Pelobates fuscus Laurenti 1768" docType="treatment" docVersion="5" lastPageNumber="138" 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="138" updateTime="1668167462434" updateUser="ExternalLinkService">
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<mods:title>Call a spade a spade: taxonomy and distribution of Pelobates, with description of a new Balkan endemic</mods:title>
</mods:titleInfo>
<mods:name type="personal">
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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>
</mods:name>
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<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Litvinchuk, Spartak N.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<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>
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<treatment ID-GBIF-Taxon="158498313" LSID="urn:lsid:plazi:treatment:44D1003976A37083B5FAB001D3340736" httpUri="http://treatment.plazi.org/id/44D1003976A37083B5FAB001D3340736" lastPageNumber="138" pageId="7" pageNumber="138">
<subSubSection pageId="7" pageNumber="138" type="nomenclature">
<paragraph pageId="7" pageNumber="138">
<taxonomicName authority="Laurenti, 1768" authorityName="Laurenti" authorityYear="1768" class="Amphibia" family="Pelobatidae" genus="Pelobates" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pelobates fuscus" order="Anura" pageId="7" pageNumber="138" phylum="Chordata" rank="species" species="fuscus">
<pageBreakToken pageId="7" pageNumber="138" start="start">Pelobates</pageBreakToken>
fuscus (Laurenti, 1768)
</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection pageId="7" pageNumber="138" type="diagnosis">
<paragraph pageId="7" pageNumber="138">Diagnosis.</paragraph>
<paragraph pageId="7" pageNumber="138">
Small spadefoot characterized by pale grayish metatarsal spades and a domed skull. The webbing of the hindfeet is well developed. Males are smaller than females (Fig. 2). The species can be found in a spectrum of gray, brown, or yellowish colors, but rarely greenish (P.
<normalizedToken originalValue="Székely">Szekely</normalizedToken>
pers. comm.), and features patterns such as stripes or blotches of varying sizes; variable presence of orange dots, from almost absent to very abundant (Fig. 3). In Eastern Europe, it differs from its sister species
<taxonomicName lsidName="P. vespertinus" pageId="7" pageNumber="138" rank="species" species="vespertinus">P. vespertinus</taxonomicName>
by most individuals having numerous dark rounded spots on a light dorsum (
<bibRefCitation author="Suriadna, NN" journalOrPublisher="Zbirnyk Prac' Zoologichnogo Muzeyu" pageId="23" pageNumber="154" pagination="67 - 87" title="Distribution, morphological variability and peculiarities of biology of spadefood toads (Amphibia, Anura, Pelobatidae) in the south of Ukraine." volume="47" year="2016">Suriadna et al. 2016</bibRefCitation>
) and lacking a dark stripe between the eyes (
<bibRefCitation pageId="7" pageNumber="138">Lada et al. 2005</bibRefCitation>
). Average SVL = 54 mm (range: 37-78 mm) for females (n = 21 populations) and 47 mm (36-65 mm) for males (n = 21 populations) (Suppl. material 1, Table S1; Fig. 2). The karyotype consists of seven large and six small pairs of two-armed chromosomes (
<bibRefCitation pageId="7" pageNumber="138">
<normalizedToken originalValue="Mészáros">Meszaros</normalizedToken>
1973
</bibRefCitation>
;
<bibRefCitation author="Schmid, M" journalOrPublisher="Chromosoma" pageId="23" pageNumber="154" pagination="271 - 284" title="Chromosome banding in Amphibia. XI. Constitutive heterochromatin, nucleolus organizers, 18 S + 28 S and 5 S ribosomal RNA genes in Ascaphidae, Pipidae, Discoglossidae and Pelobatidae." url="https://doi.org/10.1007/BF00294784" volume="95" year="1987">Schmid et al. 1987</bibRefCitation>
;
<bibRefCitation author="Manilo, VV" journalOrPublisher="Vestnik Zoologii" pageId="22" pageNumber="153" pagination="91 - 94" title="Comparative karyological research of the &quot; western &quot; form of the common spadefoot, Pelobatesfuscus (Amphibia, Pelobatidae), from Kyivska, Chernigivska and Zakarpatska Region of Ukraine." volume="38" year="2004">Manilo and Radchenko 2004</bibRefCitation>
;
<bibRefCitation author="Manilo, VV" journalOrPublisher="Zbirnyk Prac' Zoologichnogo Muzeyu" pageId="22" pageNumber="153" pagination="109 - 115" title="Comparative karyological research of two species of spadefoot toads Pelobatesfuscus and P. vespertinus (Pelobatidae, Anura, Amphibia) from territory of Ukraine." volume="44" year="2013">Manilo and Manuilova 2013</bibRefCitation>
;
<bibRefCitation author="Suriadna, NN" journalOrPublisher="Vestnik Zoologii" pageId="23" pageNumber="154" pagination="511 - 520" title="Comparative analysis of karyotypes of two cryptic species of pelobatid frogs (Amphibia, Anura) of Ukraine." url="https://doi.org/10.2478/vzoo-2014-0061" volume="48" year="2014">Suriadna 2014</bibRefCitation>
). Centromeric C-bands are obvious in pairs 2, 6, and 7-13 (
<bibRefCitation author="Schmid, M" journalOrPublisher="Chromosoma" pageId="23" pageNumber="154" pagination="271 - 284" title="Chromosome banding in Amphibia. XI. Constitutive heterochromatin, nucleolus organizers, 18 S + 28 S and 5 S ribosomal RNA genes in Ascaphidae, Pipidae, Discoglossidae and Pelobatidae." url="https://doi.org/10.1007/BF00294784" volume="95" year="1987">Schmid et al. 1987</bibRefCitation>
). NORs are in the short arm of pair 7 (
<bibRefCitation author="Schmid, M" journalOrPublisher="Chromosoma" pageId="23" pageNumber="154" pagination="83 - 103" title="Chromosome banding in Amphibia. IV. Differentiation of GC- and AT-Rich chromosome regions in Anura." url="https://doi.org/10.1007/BF00292043" volume="77" year="1980">Schmid 1980</bibRefCitation>
,
<bibRefCitation author="Schmid, M" journalOrPublisher="Chromosoma" pageId="23" pageNumber="154" pagination="327 - 344" title="Chromosome banding in Amphibia. VII. Analysis of the structure and variability of NORs in Anura." url="https://doi.org/10.1007/BF00327634" volume="87" year="1982">1982</bibRefCitation>
). The nuclear DNA content (calculated from flow cytometry) averages 8.7-9.0 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>
).
</paragraph>
</subSubSection>
<subSubSection pageId="7" pageNumber="138" type="taxonomy">
<paragraph pageId="7" pageNumber="138">Taxonomy.</paragraph>
<paragraph pageId="7" pageNumber="138">
Originally described as
<taxonomicName class="Amphibia" family="Bufonidae" genus="Bufo" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bufo fuscus" order="Anura" pageId="7" pageNumber="138" phylum="Chordata" rank="species" species="fuscus">Bufo fuscus</taxonomicName>
Laurenti, 1768; type locality: not specifically designated (&quot;in paludibus, rarissime hospitantur in continenti&quot;, in swamps, rarely on the land); type(s): the specimens depicted by
<bibRefCitation author="Roesel von Rosenhof, AJ" journalOrPublisher="Historia Naturalis Ranarum Nostratium in Qua Omnes earum Proprietates Praesertim quae ad Generationem Ipsarum Pertinent, Fusius Enarrantur / Die natuerliche Historie der Froesche hiesigen Landes, Albrecht von Haller, Nuernberg" pageId="23" pageNumber="154" title="." url="https://doi.org/10.5962/bhl.title.149946" year="1758">
<normalizedToken originalValue="Rösel">Roesel</normalizedToken>
von Rosenhof (1758
</bibRefCitation>
: pls XVII, XVIII), expressively cited by
<bibRefCitation author="Laurenti, JN" journalOrPublisher="Johann Thomas von Trattner, Vienna" pageId="21" pageNumber="152" title="Specimen medicum exhibens synopsin reptilium emendatam cum experimentis circa venena et antidota reptilium austriacorum." url="https://doi.org/10.5962/bhl.title.5108" year="1768">Laurenti (1768)</bibRefCitation>
; although controversial (see
<bibRefCitation pageId="7" pageNumber="138">
<normalizedToken originalValue="Nöllert">Noellert</normalizedToken>
et al. 2012
</bibRefCitation>
;
<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>
), the additional mention of pl. XV (p. 122), a drawing of a dissected
<taxonomicName class="Amphibia" family="Ranidae" genus="Pelophylax" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pelophylax" order="Anura" pageId="7" pageNumber="138" phylum="Chordata" rank="genus">Pelophylax</taxonomicName>
, could simply be an error.
<normalizedToken originalValue="Rösel">Roesel</normalizedToken>
depicted the amphibians of Germany, and
<bibRefCitation pageId="7" pageNumber="138">Shaw (1802)</bibRefCitation>
accordingly mentioned that
<normalizedToken originalValue="Rösel">Roesel</normalizedToken>
found his specimens in the neighborhood of
<normalizedToken originalValue="“Nurenberg”">&quot;Nurenberg&quot;</normalizedToken>
(
<normalizedToken originalValue="Nürnberg">Nuernberg</normalizedToken>
), Germany, which could then apply as the type locality. Seven junior synonyms.
<taxonomicName class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Rana alliacea" order="Anura" pageId="7" pageNumber="138" phylum="Chordata" rank="species" species="alliacea">Rana alliacea</taxonomicName>
Shaw, 1802; type locality: not specifically designated, but
<bibRefCitation pageId="7" pageNumber="138">Shaw (1802)</bibRefCitation>
refers to
<normalizedToken originalValue="Rösels">Roesel's</normalizedToken>
toads from
<normalizedToken originalValue="Nürnberg">Nuernberg</normalizedToken>
, Germany; type(s): the toad illustrated by the author (pl. 41), which may very well corresponds to the amplexed female on the top right of pl. XVII in
<bibRefCitation author="Roesel von Rosenhof, AJ" journalOrPublisher="Historia Naturalis Ranarum Nostratium in Qua Omnes earum Proprietates Praesertim quae ad Generationem Ipsarum Pertinent, Fusius Enarrantur / Die natuerliche Historie der Froesche hiesigen Landes, Albrecht von Haller, Nuernberg" pageId="23" pageNumber="154" title="." url="https://doi.org/10.5962/bhl.title.149946" year="1758">
<normalizedToken originalValue="Rösel">Roesel</normalizedToken>
von Rosenhof (1758)
</bibRefCitation>
, of identical posture and color patterns.
<taxonomicName class="Amphibia" family="Pelobatidae" genus="Bombinator" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Bombinator marmorata" order="Anura" pageId="7" pageNumber="138" phylum="Chordata" rank="species" species="marmorata">Bombinator marmorata</taxonomicName>
Sturm, 1828; type locality: near Penig, Germany; holotype: the frog illustrated by the author.
<taxonomicName class="Amphibia" family="Pelobatidae" genus="Cultripes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cultripes minor" order="Anura" pageId="7" pageNumber="138" phylum="Chordata" rank="species" species="minor">Cultripes minor</taxonomicName>
<normalizedToken originalValue="Müller">Mueller</normalizedToken>
, 1832; type locality:
<normalizedToken originalValue="“unbekannt”">&quot;unbekannt&quot;</normalizedToken>
(unknown); type(s): not mentioned.
<taxonomicName class="Amphibia" family="Pelobatidae" genus="Pelobates" higherTaxonomySource="CoL" infraspecific-rank="var." kingdom="Animalia" lsidName="Pelobates fuscus var. lividis" order="Anura" pageId="7" pageNumber="138" phylum="Chordata" rank="variety" species="fuscus" variety="lividis">Pelobates fuscus var. lividis</taxonomicName>
Koch, 1872: type locality: &quot;von den Wiesen in der
<normalizedToken originalValue="Nähe">Naehe</normalizedToken>
des
<normalizedToken originalValue="Röder-Wäldchens">Roeder-Waeldchens</normalizedToken>
bei Frankfurt&quot; (the meadows in the
<normalizedToken originalValue="Röder">Roeder</normalizedToken>
groove near Frankfurt), Germany; type(s): not mentioned;
<taxonomicName class="Amphibia" family="Pelobatidae" genus="Pelobates" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pelobates insubricus" order="Anura" pageId="7" pageNumber="138" phylum="Chordata" rank="species" species="insubricus">Pelobates insubricus</taxonomicName>
Cornalia, 1873; type locality: nearby Milano, Italy; type(s): not mentioned, most likely deposited at MSNM, but presumably lost since (
<bibRefCitation author="Blackburn, DC" journalOrPublisher="Herpetological Monographs" pageId="18" pageNumber="149" pagination="24 - 45" title="An annotated catalog of the type specimens of Amphibia in the collection of the Museo Civico di Storia Naturale, Milan, Italy." url="https://doi.org/10.1655/HERPETOLOGICA-D-13-00008" volume="28" year="2014">Blackburn and Scali 2014</bibRefCitation>
).
<taxonomicName class="Amphibia" family="Pelobatidae" genus="Pelobates" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pelobates latifrons" order="Anura" pageId="7" pageNumber="138" phylum="Chordata" rank="species" species="latifrons">Pelobates latifrons</taxonomicName>
<normalizedToken originalValue="Herón-Royer">Heron-Royer</normalizedToken>
, 1888; type locality: &quot;environ de Turin&quot; (nearby Torino), Italy; type(s): not mentioned.
<taxonomicName class="Amphibia" family="Pelobatidae" genus="Pelobates" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pelobates praefuscus" order="Anura" pageId="7" pageNumber="138" phylum="Chordata" rank="species" species="praefuscus">Pelobates praefuscus</taxonomicName>
Khosatzky, 1985; type locality: Etuliya, Moldova; holotype: ZISP 21N RNA M-1, a Pliocene fossil (according to
<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>
). The Italian populations, for long considered as a subspecies
<taxonomicName lsidName="P. f." pageId="7" pageNumber="138" rank="subSpecies" species="f." subSpecies="insubricus">P. f. insubricus</taxonomicName>
, have been a matter of debate until recently because they bear private mtDNA haplotypes (
<bibRefCitation author="Crottini, A" journalOrPublisher="Molecular Ecology" pageId="19" pageNumber="150" pagination="2734 - 2754" title="Fossorial but widespread: the phylogeography of the common spadefoot toad (Pelobatesfuscus), and the role of the Po Valley as a major source of genetic variability." url="https://doi.org/10.1111/j.1365-294X.2007.03274.x" volume="16" year="2007">Crottini et al. 2007</bibRefCitation>
).
<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>
synonymized this taxon with
<taxonomicName lsidName="P. fuscus" pageId="7" pageNumber="138" rank="species" species="fuscus">P. fuscus</taxonomicName>
, given the weak divergence of these haplotypes, together with the lack of differentiation of allozyme and genome content. As it stands,
<taxonomicName lsidName="P. fuscus" pageId="7" pageNumber="138" rank="species" species="fuscus">P. fuscus</taxonomicName>
should thus be considered a monotypic taxon.
</paragraph>
</subSubSection>
<subSubSection pageId="7" pageNumber="138" type="distribution">
<paragraph pageId="7" pageNumber="138">Distribution.</paragraph>
<paragraph pageId="7" pageNumber="138">
Widespread distribution in western, central and eastern Europe (0-810 m elevation a.s.l.), but absent from the northern European countries and most of southern Europe (
<bibRefCitation author="Sillero, N" journalOrPublisher="Amphibia-Reptilia" pageId="23" pageNumber="154" pagination="1 - 31" title="Updated distribution and biogeography of amphibians and reptiles of Europe." url="https://doi.org/10.1163/15685381-00002935" volume="35" year="2014">Sillero et al. 2014</bibRefCitation>
;
<normalizedToken originalValue="Nöllert">Noellert</normalizedToken>
at al. 2012) (Fig. 1). In the west, it reaches the eastern edge of the Netherlands (
<bibRefCitation author="Creemers, RCM" journalOrPublisher="Nationaal Natuurhistorisch Museum Naturalis, Leiden" pageId="19" pageNumber="150" title="De amfibieen en reptielen van Nederland." year="2009">Creemers and Van Delft 2009</bibRefCitation>
), the eastern part of Flanders in Belgium (
<bibRefCitation author="Bauwens, D" journalOrPublisher="De Wielewaal Natuurvereniging, Turnhout" pageId="18" pageNumber="149" title="Verspreiding van Amfibieen en Reptielen in Vlaanderen." year="1996">Bauwens and Claus 1996</bibRefCitation>
), the western parts of Nordrhein-Westfalens and the south-east of Rheinland-Pflaz in Germany (
<bibRefCitation pageId="7" pageNumber="138">Bitz et al. 1996</bibRefCitation>
;
<bibRefCitation pageId="7" pageNumber="138">Chmela and Kronshage 2011</bibRefCitation>
), the north-eastern side of France (particularly along the Rhine River,
<bibRefCitation pageId="7" pageNumber="138">Eggert and Vacher 2012</bibRefCitation>
). In the north, it extends to northern Netherlands (
<bibRefCitation author="Creemers, RCM" journalOrPublisher="Nationaal Natuurhistorisch Museum Naturalis, Leiden" pageId="19" pageNumber="150" title="De amfibieen en reptielen van Nederland." year="2009">Creemers and Van Delft 2009</bibRefCitation>
), the North Sea coastline of Germany (
<bibRefCitation pageId="7" pageNumber="138">
<normalizedToken originalValue="Nöllert">Noellert</normalizedToken>
and
<normalizedToken originalValue="Günther">Guenther</normalizedToken>
1996
</bibRefCitation>
) and Denmark, the south of Sweden, as well as the coastline of the Baltic Sea from Germany to Estonia, and eastward until it reaches
<taxonomicName lsidName="P. vespertinus" pageId="7" pageNumber="138" rank="species" species="vespertinus">P. vespertinus</taxonomicName>
in Russia (
<bibRefCitation author="Kuzmin, SL" journalOrPublisher="Pensoft Publisher, Sofia" pageId="21" pageNumber="152" title="The amphibians of the former Soviet Union." year="1999">Kuzmin 1999</bibRefCitation>
;
<bibRefCitation author="Nystroem, P" journalOrPublisher="Biological Conservation" pageId="23" pageNumber="154" pagination="399 - 411" title="A documented amphibian decline over 40 years: possible causes and implications for species recovery." url="https://doi.org/10.1016/j.biocon.2007.05.007" volume="138" year="2007">
<normalizedToken originalValue="Nyström">Nystroem</normalizedToken>
et al. 2007
</bibRefCitation>
;
<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>
;
<bibRefCitation author="Sillero, N" journalOrPublisher="Amphibia-Reptilia" pageId="23" pageNumber="154" pagination="1 - 31" title="Updated distribution and biogeography of amphibians and reptiles of Europe." url="https://doi.org/10.1163/15685381-00002935" volume="35" year="2014">Sillero et al. 2014</bibRefCitation>
). The contact zone with the latter is well delineated from the Kursk region in Russia to the Black Sea coast (
<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>
). From there, it is present westward along the Black Sea coast of Ukraine to north-eastern Bulgaria (
<bibRefCitation author="Kuzmin, SL" journalOrPublisher="Pensoft Publisher, Sofia" pageId="21" pageNumber="152" title="The amphibians of the former Soviet Union." year="1999">Kuzmin 1999</bibRefCitation>
;
<bibRefCitation author="Stojankov, A" journalOrPublisher="Chimaira, Frankfurth am Main" pageId="23" pageNumber="154" title="Die Amphibien und Reptilien Bulgariens." year="2011">Stojankov et al. 2011</bibRefCitation>
). The southern edges extend along the Danube at the borders of Romania and Bulgaria (
<bibRefCitation author="Stojankov, A" journalOrPublisher="Chimaira, Frankfurth am Main" pageId="23" pageNumber="154" title="Die Amphibien und Reptilien Bulgariens." year="2011">Stojankov et al. 2011</bibRefCitation>
) and across Serbia (
<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>
), eastern Croatia, northern Bosnia and Herzegovina, Slovenia (
<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="Curic, A" journalOrPublisher="North-Western Journal of Zoology" pageId="19" pageNumber="150" pagination="50 - 59" title="New data and distribution of common spadefoot toad Pelobatesfuscus (Laurenti, 1768) (Anura: Pelobatidae) in Western Balkans." volume="14" year="2018">
<normalizedToken originalValue="Curić">Curic</normalizedToken>
et al. 2018
</bibRefCitation>
), northern and eastern Austria around the Alps (
<bibRefCitation author="Cabela, A" journalOrPublisher="Umweltbundesamt, Wien" pageId="19" pageNumber="150" title="Atlas zur Verbreitung und Oekologie der Amphibien und Reptilien in Oesterreich." year="2001">Cabela et al. 2001</bibRefCitation>
), and southern Germany (
<normalizedToken originalValue="Nöllert">Noellert</normalizedToken>
and Gunther 1996). The species is also present in a large area of northern Italy, especially in the Po Valley (
<bibRefCitation pageId="7" pageNumber="138">Andreone 2006</bibRefCitation>
). Last, isolated populations persist in central France (Indre, Loiret, Indre-et-Loire:
<bibRefCitation pageId="7" pageNumber="138">Eggert and Vacher 2012</bibRefCitation>
) and western Bulgaria (around Sofia:
<bibRefCitation author="Stojankov, A" journalOrPublisher="Chimaira, Frankfurth am Main" pageId="23" pageNumber="154" title="Die Amphibien und Reptilien Bulgariens." year="2011">Stojankov et al. 2011</bibRefCitation>
). IUCN Status: Not Evaluated, considered Least Concern when grouped with
<taxonomicName lsidName="P. vespertinus" pageId="7" pageNumber="138" rank="species" species="vespertinus">P. vespertinus</taxonomicName>
(
<bibRefCitation author="Agasyan, A" pageId="18" pageNumber="149" url="https://doi.org/10.2305/IUCN.UK.2009.RLTS.T16498A5951455.en" year="2009 a">Agasyan et al. 2009a</bibRefCitation>
). Declines have been reported for more than a century in various parts of Europe, which have caused a regression of the distribution limits (
<bibRefCitation author="Dzukic, G" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="20" pageNumber="151" url="https://doi.org/10.3409/173491505783995699" year="2005">
<normalizedToken originalValue="Džukić">Dzukic</normalizedToken>
et al. 2005
</bibRefCitation>
;
<bibRefCitation author="Eggert, C" journalOrPublisher="Conservation Genetics" pageId="20" pageNumber="151" pagination="185 - 195" title="The declining spadefoot toad, Pelobatesfuscus (Pelobatidae): paleo and recent environmental changes as a major influence on current population structure and status." url="https://doi.org/10.1007/s10592-006-9124-y" volume="7" year="2006">Eggert et al. 2006</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="7" pageNumber="138" type="diversity">
<paragraph pageId="7" pageNumber="138">Diversity.</paragraph>
<paragraph pageId="7" pageNumber="138">
The phylogeographic work by
<bibRefCitation author="Crottini, A" journalOrPublisher="Molecular Ecology" pageId="19" pageNumber="150" pagination="2734 - 2754" title="Fossorial but widespread: the phylogeography of the common spadefoot toad (Pelobatesfuscus), and the role of the Po Valley as a major source of genetic variability." url="https://doi.org/10.1111/j.1365-294X.2007.03274.x" volume="16" year="2007">Crottini et al. (2007)</bibRefCitation>
and
<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>
characterized two refugial groups for this species (as the &quot;western lineage of
<taxonomicName lsidName="P. fuscus" pageId="7" pageNumber="138" rank="species" species="fuscus">P. fuscus</taxonomicName>
&quot;), based on shallow mtDNA divergence and allozyme differentiation: in the Balkans/northern Italy and on the western shores of the Black Sea coast. This seems supported by weak genomic differentiation among Central-European samples (
<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>
). The refugial areas bear nearly all the genetic diversity of the species, which was lost in the derived northern populations, following post-glacial colonizations (
<bibRefCitation author="Eggert, C" journalOrPublisher="Conservation Genetics" pageId="20" pageNumber="151" pagination="185 - 195" title="The declining spadefoot toad, Pelobatesfuscus (Pelobatidae): paleo and recent environmental changes as a major influence on current population structure and status." url="https://doi.org/10.1007/s10592-006-9124-y" volume="7" year="2006">Eggert et al. 2006</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>