Add updates up until 2025-01-23 17:02:47

This commit is contained in:
ggserver 2025-01-23 17:08:51 +00:00
parent 25789522ec
commit 6b89830877

View file

@ -0,0 +1,689 @@
<document id="06B83F9C82B224F1FC178E704CC252A4" ID-DOI="10.3897/vz.75.e138687" ID-ZooBank="10CFDEFC-8E1B-448E-9BFC-7A6C8B2AB776" ID-publisher-id="138687" URI-arpha="2C83DC53-3E6F-58EA-8773-0F1DEFB7D76A" URI-zoobank="http://zoobank.org/10CFDEFC-8E1B-448E-9BFC-7A6C8B2AB776" XM.bibliography_approvedBy="admin" XM.materialsCitations_approvedBy="admin" XM.taxonomicNames_approvedBy="admin" XM.treatments_approvedBy="admin" article-type="research-article" checkinTime="1737645806352" checkinUser="pensoft" docAuthor="Dufresnes, Christophe, Lukanov, Simeon, Gippner, Sven, Ambu, Johanna, Strachinis, Ilias, Arsovski, Dragan, Monod-Broca, Benjamin, Cayuela, Hugo, Lymberakis, Petros, Canestrelli, Daniele, Cogălniceanu, Dan, Poyarkov, Nikolay A., Litvinchuk, Spartak N., Suchan, Tomasz, Denoël, Mathieu &amp; Jablonski, Daniel" docDate="2025" docId="7B3AD50B77B55EE9A5EC0F768D307567" docLanguage="en" docName="Vertebrate Zoology 75: 1-30" docOrigin="Vertebrate Zoology 75" docSource="https://vertebrate-zoology.arphahub.com/article/138687/download/xml/" docStyle="DocumentStyle:PensoftTaxPub.0000.journal_article.generic" docStyleName="PensoftTaxPub.0000.journal_article.generic" docTitle="Bombina variegata subsp. rhodopensis Lukanov, Denoel, Jablonski &amp; Dufresnes 2025, subsp. nov." docType="treatment" docVersion="3" dtd-version="3.0" lastPageNumber="30" masterDocId="2C83DC533E6F58EA87730F1DEFB7D76A" masterDocTitle="Historical biogeography and systematics of yellow-bellied toads (Bombina variegata), with the description of a new subspecies from the Balkans" masterLastPageNumber="30" masterPageNumber="1" pageNumber="1" updateTime="1737651765380" updateUser="admin">
<mods:mods id="6198245E0FC6E1F0B9CD6C2C7CDD1F0A" xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo id="30DF4C714C5705661536A7CC469FBD41">
<mods:title id="C4B96229795392FCB9C2438A308C7700">Historical biogeography and systematics of yellow-bellied toads (Bombina variegata), with the description of a new subspecies from the Balkans</mods:title>
</mods:titleInfo>
<mods:name id="D717715F4A8A57E4425DB7B638F96CC0" type="personal">
<mods:role id="1FF3C63FC888D6044F01CC7F58ABF37B">
<mods:roleTerm id="98BDE04CE4F15BFD271D20A5DD0D077E">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="0D271903AB1F4C7B0280FCD2373F36D4">Dufresnes, Christophe</mods:namePart>
<mods:nameIdentifier id="7CAE93E61DA135CB1F3A3E1178A379F6" type="ORCID">0000-0002-8497-8908</mods:nameIdentifier>
<mods:affiliation id="286321348385BFED672A29384C04EA7F">Institut de Systématique, Evolution, Biodiversité (ISYEB), Muséum national dHistoire naturelle, CNRS, Sorbonne Université, EPHE-PSL, Université des Antilles, 55 rue Buffon, CP 51, 75005 Paris, France</mods:affiliation>
</mods:name>
<mods:name id="48CFF7A5A285449E4C1BA8485885A91B" type="personal">
<mods:role id="50058CE0257D1A18ADBCCA0FA1EB8CF3">
<mods:roleTerm id="4C2D5F9038D2367B10C85B9CA7F16842">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="7724394538A0D71C2B281A43BCBFC6E1">Lukanov, Simeon</mods:namePart>
<mods:affiliation id="3FF1498326DC4437CCF14C4848AD0CEB">Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, Tzar Osvoboditel 1, 1000 Sofia, Bulgaria</mods:affiliation>
</mods:name>
<mods:name id="233D7EB181C69EB97A78857077ECC4CA" type="personal">
<mods:role id="7804ACD9FD4DE31F9FF1FC0A1CC3BD58">
<mods:roleTerm id="EE24654A9450F1A6233B1A33676E51DA">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="DC8DDD9F60C6793C3CE1E66B72DFD6FB">Gippner, Sven</mods:namePart>
<mods:affiliation id="A537E7CFAC5717FEED6F75B08BDCB122">Zoological Institute, Technical University of Braunschweig, Mendelssohnstr. 4, 38106 Braunschweig, Germany</mods:affiliation>
</mods:name>
<mods:name id="651CC824B4DBE125189ED8C4864D09B4" type="personal">
<mods:role id="DED7A953B2C2BDB94BF5DD4D6AA17224">
<mods:roleTerm id="0EAF8A11C97BCA667AE58AF143A704E9">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="1F28C2E0AC38EA4386867BA1DB50BDB2">Ambu, Johanna</mods:namePart>
<mods:affiliation id="8569ADE7EBBA437D3B660DB7C318D370">Laboratory for Amphibian Systematics and Evolutionary Research, College of Biology &amp; the Environment, Nanjing Forestry University, Nanjing, China</mods:affiliation>
</mods:name>
<mods:name id="E50A8E1713FF401C8C2AF5941B25D81C" type="personal">
<mods:role id="FB188DF8FAFBFA6833DFE3E3294D4344">
<mods:roleTerm id="386591C4E3B8B197973CCCB7F79ED5BF">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="E37AA3D3638300CDB040DB19E007F622">Strachinis, Ilias</mods:namePart>
<mods:nameIdentifier id="92AFF55D490617A30BE443CE2148BED7" type="ORCID">0000-0001-9944-4522</mods:nameIdentifier>
<mods:affiliation id="8F4CC9221F6F20AC65C9F9595750F7E1">Department of Genetics, Development and Molecular Biology, School of Biology, Faculty of Natural Sciences, Aristotle University of Thessaloniki, 54636, Thessaloniki, Greece</mods:affiliation>
</mods:name>
<mods:name id="3AB720814A1157B33A21BA8FF79E9FB4" type="personal">
<mods:role id="DF10AF001F0C0EB1BA371437C056F7B4">
<mods:roleTerm id="6CB76B62D9F505AEEC53B5925F1C423B">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="86A710BBB992070E63EAE6C672E98135">Arsovski, Dragan</mods:namePart>
<mods:affiliation id="E7A805281E0DAEFBEB45077C726B8DBC">Macedonian Ecological Society, Arhimedova 5, 1000 Skopje, North Macedonia</mods:affiliation>
</mods:name>
<mods:name id="9AA767160748A1F2AC19BB3DEE9ED241" type="personal">
<mods:role id="B8DC7547E52A3B9FE0C2D0531FB7CA1E">
<mods:roleTerm id="607120F3460AEC91DC05FD43A8922AB3">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="5ED95CC491A5127041A948120327435C">Monod-Broca, Benjamin</mods:namePart>
<mods:nameIdentifier id="9A13A679A92C13BF9E2E49FF16D7B2E8" type="ORCID">0000-0002-9133-5548</mods:nameIdentifier>
<mods:affiliation id="9C7B5C79F760ED58288689660F780A5B">Université Claude Bernard Lyon 1, LEHNA UMR 5023, CNRS, ENTPE, F- 69622, Villeurbanne, France</mods:affiliation>
</mods:name>
<mods:name id="C6EBABA935CF2D205C91D2D3AA805622" type="personal">
<mods:role id="95554540EEA18AB049B2D1AAD5BCD00D">
<mods:roleTerm id="120A976E8C8DEBEAE4E427C8A9572DAF">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="0C5EF71C9CD8A8B069F3EB0D7D4317F0">Cayuela, Hugo</mods:namePart>
<mods:affiliation id="E7640416C18E4E41DBE1BEB6182E59F9">Université Claude Bernard Lyon 1, LEHNA UMR 5023, CNRS, ENTPE, F- 69622, Villeurbanne, France</mods:affiliation>
</mods:name>
<mods:name id="DDDD9453B441F7ED009FB5F2BC9D3ED7" type="personal">
<mods:role id="68B89E4AA404901FC885CC1D26DB6656">
<mods:roleTerm id="0D97ED762B76F1F2EE4A068464CA1313">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="3B4AE7741607F0D5C06042ADCED794BA">Lymberakis, Petros</mods:namePart>
<mods:nameIdentifier id="02434B6A2DD8C0E870F14B774F89FAE7" type="ORCID">0000-0002-5067-9600</mods:nameIdentifier>
<mods:affiliation id="6753A09A55D3181FFDE60E8C957FAE35">Université Claude Bernard Lyon 1, CNRS, VetAgro Sup, UMR 5558 Biométrie et Biologie Evolutive, Bat. G Mendel 43 bd du 11 Novembre 1918, 69622 Villeurbanne Cedex, France</mods:affiliation>
</mods:name>
<mods:name id="C748FC5645E5A554BEAEB308C1CBE653" type="personal">
<mods:role id="423FE2AE597227CDE9BEE145A2FC3548">
<mods:roleTerm id="6DB34892027086EBEFEAC328934E8931">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="9CAA294E9DC1C6FD601D23749ECA1712">Canestrelli, Daniele</mods:namePart>
<mods:affiliation id="E57DB48197B99D73221799E2AC539AAB">Natural History Museum of Crete, School of Sciences and Engineering, University of Crete, Heraklion, Greece</mods:affiliation>
</mods:name>
<mods:name id="AD25D50209E0B3E93B21E12102E73846" type="personal">
<mods:role id="E80BB4623D6E5EC2FBF24AEBAA881700">
<mods:roleTerm id="70B9E9E1C802A9526B30C5CFD6A77B02">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="CCFA37194A8F1ECAFC400F71BE00C0A0">Cogălniceanu, Dan</mods:namePart>
<mods:affiliation id="7A360FB3349FA46E265B7D4502291A6D">Department of Ecological and Biological Sciences, University of Tuscia, Largo dellUniversità snc, 01100 Viterbo, Italy</mods:affiliation>
</mods:name>
<mods:name id="622AACE350330EB83C143D1B0F665D99" type="personal">
<mods:role id="693C18844512D87CF7C70DFFAB707939">
<mods:roleTerm id="200E6F0AB434882F8D9C393540426E5B">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="821392AC0696DCE0B81802604E21BFE8">Poyarkov, Nikolay A.</mods:namePart>
<mods:nameIdentifier id="5343AD7E8CBDD18C206ADD1CD66C5ACF" type="ORCID">0000-0002-7576-2283</mods:nameIdentifier>
<mods:affiliation id="1F3E2B4A16A0E2D6B7E075917EE8E7F0">University Ovidius Constanţa, Faculty of Natural Sciences and Agricultural Sciences, A. Universităţii 1, 900470, Constanţa, Romania</mods:affiliation>
</mods:name>
<mods:name id="14664772BDC0C84B144800D5A39F52C1" type="personal">
<mods:role id="F5D6E63035AEFFBEA51359158B7A10AE">
<mods:roleTerm id="273133E1994A1E70A38E20DF7A325E9F">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="65AC704D5568D8F3F47F881068423B33">Litvinchuk, Spartak N.</mods:namePart>
<mods:affiliation id="19DD5F0D83BE1816CDFA5430BFF35D8E">Department of Vertebrate Zoology, Lomonosov Moscow State University, Leninskiye Gory, Moscow 119234, Russia &amp; Institute of Cytology, Russian Academy of Sciences, Saint Petersburg, Russia</mods:affiliation>
</mods:name>
<mods:name id="4E26117FC3323E3ED8467D469CAEC9DB" type="personal">
<mods:role id="3DC521EF5CF8F600AD110770B028EDCC">
<mods:roleTerm id="20E902DF50197FD70C61C0C81F2060C3">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="524FB89DAF4A7A4EDAF932605DB37C70">Suchan, Tomasz</mods:namePart>
<mods:nameIdentifier id="2B442E83972D8D01021E625C5188040A" type="ORCID">0000-0002-0811-8754</mods:nameIdentifier>
<mods:affiliation id="FB1A5CA3E56A343C94E9FC7BC659F975">Dagestan State University, Makhachkala, Russia</mods:affiliation>
</mods:name>
<mods:name id="4F5E9DAEC336283133AD15938F9890EE" type="personal">
<mods:role id="5A6BE85308367FF6820905C78830ED1F">
<mods:roleTerm id="A6251A7A74D3BDD1A8A9C494FFA8EEB2">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="68E72802C56A678447E5C8DC0A2CEBB2">Denoël, Mathieu</mods:namePart>
<mods:nameIdentifier id="C30BC9F1A9F9AA94F4A8AEA197349467" type="ORCID">0000-0002-3586-8323</mods:nameIdentifier>
<mods:affiliation id="E3873E3EC02BD1C9555ECB1C28D7A47A">W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz, 46, 31 512 Kraków, Poland</mods:affiliation>
</mods:name>
<mods:name id="927CC39EC7DCFE5C38CE67468404407B" type="personal">
<mods:role id="E8D3BFFC6C86AC918D1D9023B825092D">
<mods:roleTerm id="0C1EEB90283308B5F1654B466AF4FF24">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="9E6C6DAFF6C8FEBAB891856D6B744A9C">Jablonski, Daniel</mods:namePart>
<mods:nameIdentifier id="31D293D0CAE93BEA987A40332D65DC9B" type="ORCID">0000-0002-5394-0114</mods:nameIdentifier>
<mods:affiliation id="77C3AE5EF7BA0F6EEC685B24F4A8A7BB">Laboratory of Ecology and Conservation of Amphibians (LECA), FOCUS, University of Liège, Belgium</mods:affiliation>
</mods:name>
<mods:typeOfResource id="147EAA500D2F33A26678D3C0264FE032">text</mods:typeOfResource>
<mods:relatedItem id="F72F31A7F6CA0F6D47917F326B734BFB" type="host">
<mods:titleInfo id="D794784486EAEC1BB9D30FC90FF42D32">
<mods:title id="022F5DB8FB0BF173B5C7AF7B4DBBB9FA">Vertebrate Zoology</mods:title>
</mods:titleInfo>
<mods:part id="EC6237FD6F9576F9EEA99803033A6697">
<mods:date id="22006E3FAC275EAC9E93D852AA257F81">2025</mods:date>
<mods:detail id="FE2606282B78BAA0AF6EE67241AFB29D" type="pubDate">
<mods:number id="49726948C42BF43FA9DFB41C45C82FE4">2025-01-23</mods:number>
</mods:detail>
<mods:detail id="47E2F391E24B092A07A1BB82BB23FB23" type="volume">
<mods:number id="F8FC4F13D33836E90305E12DCDFEE1A4">75</mods:number>
</mods:detail>
<mods:extent id="4CF83C4F403C9BF1C17A16DE7FF547DA" unit="page">
<mods:start id="DA1D29E1961EE9B2C36562D6675FAC71">1</mods:start>
<mods:end id="2D88E964C25740A9D7A54DF990F84F52">30</mods:end>
</mods:extent>
</mods:part>
</mods:relatedItem>
<mods:classification id="2AFCE2AC60D3B4B16D81C2A72AD9E47F">journal article</mods:classification>
<mods:identifier id="0F9361DB2F53C50E03F9154ABE039BD8" type="DOI">10.3897/vz.75.e138687</mods:identifier>
<mods:identifier id="934BFFDC5DD848E6A19546E561F7102B" type="ZooBank">10CFDEFC-8E1B-448E-9BFC-7A6C8B2AB776</mods:identifier>
</mods:mods>
<treatment id="7B3AD50B77B55EE9A5EC0F768D307567" ID-arpha="7B3AD50B-77B5-5EE9-A5EC-0F768D307567" ID-zoobank="https://zoobank.org/E61B7D1D-FF8D-48A4-93C9-DE43EAC3B2B4" LSID="urn:lsid:plazi:treatment:7B3AD50B77B55EE9A5EC0F768D307567" LSID-ZBK="urn:lsid:zoobank.org:act:E61B7D1D-FF8D-48A4-93C9-DE43EAC3B2B4" httpUri="http://treatment.plazi.org/id/7B3AD50B77B55EE9A5EC0F768D307567">
<subSubSection id="078CBC933EAFEA3035ECE85472981FDB" type="nomenclature">
<paragraph id="432D84F69C428F4C8961C19641AA771E">
<taxonomicName id="30166C6278139250A62F0EBBE0990FF1" authority="Lukanov, Denoel, Jablonski &amp; Dufresnes" authorityName="Lukanov, Denoel, Jablonski &amp; Dufresnes" authorityYear="2025" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="subSpecies" species="variegata" status="subsp. nov." subSpecies="rhodopensis">
<emphasis id="5F7D23627EF927BC50B7AC018D2335A0" italics="true">Bombina variegata rhodopensis</emphasis>
Lukanov, Denoël, Jablonski &amp; Dufresnes
</taxonomicName>
<taxonomicNameLabel id="F57D83D27C43C12E2FD004D891B389CC" rank="subSpecies">subsp. nov.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection id="SECID0EFVAK" type="Identity">
<paragraph id="8A93FF89C8F53C242032CC1D2B413848">
<heading id="CF6E866F132DE85FBDC4D89291F965F0" reason="title">Identity.</heading>
</paragraph>
<paragraph id="CA399E81346AE380B6DF0B60224CDFBA">
Previously identified as a divergent mitochondrial (cyt
<emphasis id="0D19A62053741314E4C9D03557A56BD0" italics="true">b</emphasis>
) lineage, attributed to the subspecies
<taxonomicName id="F8EE045096F25A08E41C75F3A6A04E35" baseAuthorityName="Mertens and Muller" baseAuthorityYear="1940" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="subSpecies" species="variegata" subSpecies="scabra">
<emphasis id="1019B8CCFA03C05DE46984B2A2F69D03" italics="true">Bombina variegata scabra</emphasis>
</taxonomicName>
by
<bibRefCitation id="C4B50AD7AB1C79980191F51DB13DF570" DOI="10.1111/j.1365-294X.2007.03309.x" author="Hofman S &amp; Spolsky C &amp; Uzzell T &amp; Cogălniceanu D &amp; Babik W &amp; Szymura JM" etAl="et al." firstAuthor="Hofman" journalOrPublisher="Molecular Ecology" pagination="2301-2316" refId="B79" refString="Hofman S, Spolsky C, Uzzell T, Cogălniceanu D, Babik W, Szymura JM (2007) Phylogeography of the fire-bellied toads Bombina: Independent Pleistocene histories inferred from mitochondrial genomes. Molecular Ecology 16: 23012316. https://doi.org/10.1111/j.1365-294X.2007.03309.x" title="Phylogeography of the fire-bellied toads Bombina: Independent Pleistocene histories inferred from mitochondrial genomes." volume="16" year="2007">Hofman et al. (2007)</bibRefCitation>
and
<bibRefCitation id="63284ABEBB72ED4615F13F2ED45F63D4" DOI="10.1111/j.1365-294X.2011.05175.x" author="Fijarczyk A &amp; Nadachowska K &amp; Hofman S &amp; Litvinchuk SN &amp; Babik W &amp; Stuglik M &amp; Gollmann G &amp; Choleva L &amp; Cogălniceanu D &amp; Vukov T &amp; Džukić G &amp; Szymura JM" etAl="et al." firstAuthor="Fijarczyk" journalOrPublisher="Molecular Ecology" pagination="3381-3398" refId="B58" refString="Fijarczyk A, Nadachowska K, Hofman S, Litvinchuk SN, Babik W, Stuglik M, Gollmann G, Choleva L, Cogălniceanu D, Vukov T, Džukić G, Szymura JM (2011) Nuclear and mitochondrial phylogeography of the European fire-bellied toads Bombina bombina and Bombina variegata supports their independent histories. Molecular Ecology 20: 33813398. https://doi.org/10.1111/j.1365-294X.2011.05175.x" title="Nuclear and mitochondrial phylogeography of the European fire-bellied toads Bombina bombina and Bombina variegata supports their independent histories." volume="20" year="2011">Fijarczyk et al. (2011)</bibRefCitation>
and labeled as the “ Rhodopean ”, “ R ” or “ RD ”, in reference to its distribution in the Rhodope Mountains of southeastern
<collectingCountry id="FD363EBE42B02953FA79AA3F740E2BF3" name="Bulgaria">Bulgaria</collectingCountry>
and northeastern
<collectingCountry id="AF3DABFD4910986BACEE816B898F3292" name="Greece">Greece</collectingCountry>
. This new taxon corresponds to the Rhodope lineage (
<abbrev id="ABBRID0EEWAK" xlink_title="mitochondrial DNA">mtDNA</abbrev>
R) that is purple-coded in our figures. Given its level of divergence (see Discussion), we describe this lineage as a new subspecies,
<taxonomicName id="23FFA2C87E936D808D49C6A86139AB04" authorityName="Lukanov, Denoel, Jablonski &amp; Dufresnes" authorityYear="2025" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="subSpecies" species="variegata" status="subsp. nov." subSpecies="rhodopensis">
<emphasis id="6A6580988AA439061B4B21F6FC98D09C" italics="true">Bombina variegata rhodopensis</emphasis>
</taxonomicName>
<emphasis id="660496E0EEFCBC06D091AAFECCB41487" bold="true">
<taxonomicNameLabel id="305F15256D157AF5844E1450666C95F5" rank="subSpecies">subsp. nov.</taxonomicNameLabel>
</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="SECID0EYWAK" type="materials_examined">
<paragraph id="53FCE9CEB41C1AE315F37C0B596243AF">
<heading id="BA40A2870C2DD53D05DA5256175E7655" reason="title">
<typeStatus id="2A4BB25B7D0B9F582A249FEBCF80F074" type="holotype">Holotype</typeStatus>
.
</heading>
</paragraph>
<paragraph id="4DDCACF4AED0A41D96631D3ED36C98BF">
<materialsCitation id="8AC8EF10B8D15EB58B2102E278CB29CA" collectingDate="2024-04-23" collectionCode="IBER-BAS" collectorName="SL &amp; MD &amp; DJ &amp; CD" country="Bulgaria" county="The" elevation="563" latitude="41.7039" location="water fountain at the northeastern exit of Kostino" longLatPrecision="6" longitude="25.3028" municipality="Kardzhali Municipality" specimenCode="BG-IBER-VER-000010561" specimenCount="1" specimenCount-male="1" stateProvince="Kardzhali Province" typeStatus="holotype">
<specimenCode id="121FA395C801EE6FFF94A5631B61C35C">BG-IBER-VER-000010561</specimenCode>
,
<specimenCount id="D0E861E5A1712A3390AD94F5277FFA61" count="1" type="male">adult male</specimenCount>
collected on
<date id="E1B8AE806202819B07518BAC2AD1F3FB" value="2024-04-23">
<collectingDate id="CF5F08FE04A9EB82EB0A761DC5E99DAA" value="2024-04-23">23 April 2024</collectingDate>
</date>
by
<collectorName id="BDEE423F38C691E2B7B5B39D5DFEAFD0">SL</collectorName>
,
<collectorName id="6656DC64720BA8DA6E7F7ACF1E90015E">MD</collectorName>
,
<collectorName id="AAD343E3593481B5A1A22317CAB74C83">DJ</collectorName>
and
<collectorName id="0E0A0C6E7A6958E6DB65557D1DA8AFD8">CD</collectorName>
in a
<location id="39FCD9BB119955DACEE8A2F96EDA53FE" LSID="urn:lsid:plazi:treatment:7B3AD50B77B55EE9A5EC0F768D307567:39FCD9BB119955DACEE8A2F96EDA53FE" country="Bulgaria" county="The" latitude="41.7039" longLatPrecision="6" longitude="25.3028" municipality="Kardzhali Municipality" name="water fountain at the northeastern exit of Kostino" stateProvince="Kardzhali Province">water fountain at the northeastern exit of Kostino</location>
,
<collectingMunicipality id="84DB8EB90A8E63FD9369FB3F7F74EECC">Kardzhali Municipality</collectingMunicipality>
,
<collectingRegion id="4DF6BA626DBD4ABA358C304CA09B6828" country="Bulgaria" name="Kurdzhali">Kardzhali Province</collectingRegion>
,
<collectingCountry id="10B94E203B12F52659218C3D92A03A22" name="Bulgaria">Bulgaria</collectingCountry>
(
<named-content content-type="dwc:verbatimCoordinates" id="NCID0EBXAK" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[25.302800,41.703900]}">
<geoCoordinate id="8A80691F6DFC6B3E9B422FB518C2BB2F" degrees="41.7039" direction="north" orientation="latitude" precision="5" value="41.7039">41.7039 ° N</geoCoordinate>
,
<geoCoordinate id="4A1E9BDC69F81D855CC10E3752E05A45" degrees="25.3028" direction="east" orientation="longitude" precision="5" value="25.3028">25.3028 ° E</geoCoordinate>
</named-content>
; elevation:
<elevation id="1BBD9D3FD89568DA96BC5DF85AE06B37" metricMagnitude="2" metricUnit="m" metricValue="5.63" unit="m" value="563.0">
<quantity id="9FBC2383BF8F9DD16E8DF3711D2EB24F" metricMagnitude="2" metricUnit="m" metricValue="5.63" unit="m" value="563.0">563 m</quantity>
a. s. l.
</elevation>
), and curated at the
<named-content id="E94BE9F5896130B1F21597BC228EEF93" content-type="institution" xlink_href="http://grbio.org/institution/bulgarian-academy-science" xlink_type="simple">Institute of Biodiversity and Ecosystem Research of the Bulgarian Academy of Sciences</named-content>
(
<named-content id="AFB164C4DECF9AD7BB0AEFDBFCD3031F" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/bulgarian-academy-science" xlink_title="Institute of Biodiversity and Ecosystem Research of the Bulgarian Academy of Sciences">
<emphasis id="73A8A10D23660BF4306C231B08A04A95" bold="true">
<collectionCode id="01463E577D1D4C1FE621E2E1220B3642">IBER-BAS</collectionCode>
</emphasis>
</named-content>
), Sofia, Bulgaria
</materialsCitation>
. Measurements in mm: L. 42.7, F. 16.3, T. 17.3,
<abbrev id="ABBRID0EMXAK" xlink_title="head width, between the posterior edges of jaw articulations">Lt. c</abbrev>
. 12.8,
<abbrev id="ABBRID0EQXAK" xlink_title="inter-orbital distance, as the minimum distance between orbits">Sp. p</abbrev>
. 4.8,
<abbrev id="ABBRID0EUXAK" xlink_title="maximum length of the upper eyelid">L. o</abbrev>
. 4.4,
<abbrev id="ABBRID0EYXAK" xlink_title="inter-nostril distance">Sp. n</abbrev>
. 2.4. The holotype and the type locality are depicted on Fig.
<figureCitation id="3CFF3782DC697BC206DE01F88CCC88D0" captionStart="Figure 10" captionStartId="F10" captionText="Figure 10. The holotype and type locality of Bombina variegata rhodopensis subsp. nov. Credit: CD (top and middle) and DJ (bottom)." figureDoi="10.3897/vz.75.e138687.figure10" httpUri="https://binary.pensoft.net/fig/1230623">10</figureCitation>
.
</paragraph>
<caption id="93F331BC54D78CD861092B709826120A" ID-DOI="10.3897/vz.75.e138687.figure10" ID-arpha="4A9B5AC9-00F4-53F5-A7DA-CE869DDE7E1C" httpUri="https://binary.pensoft.net/fig/1230623" startId="F10">
<paragraph id="11553D8F5C305BDB53FEC6A74CAEA100">
<label id="3D8473BA0924B01A71780FCECEAF2888">Figure 10.</label>
</paragraph>
<paragraph id="885EC2EEB5EC5952DABFEAE9AC43C068">
The holotype and type locality of
<taxonomicName id="5B1260250E02EB1F96CF50802D3584DD" authorityName="Lukanov, Denoel, Jablonski &amp; Dufresnes" authorityYear="2025" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="subSpecies" species="variegata" status="subsp. nov." subSpecies="rhodopensis">
<emphasis id="5BFA3AE5A287C62AF24F36AEF6CF73DD" italics="true">Bombina variegata rhodopensis</emphasis>
</taxonomicName>
<emphasis id="54BC5B7B78A4ACB01C9AA85AE41B1D1E" bold="true">
<taxonomicNameLabel id="6A2649488B7D987DD1815D8CA506C180" rank="subSpecies">subsp. nov.</taxonomicNameLabel>
</emphasis>
Credit: CD (top and middle) and DJ (bottom).
</paragraph>
</caption>
<paragraph id="10371C0736C480608D82DC911F529A7F">
<heading id="DF2ED7E5061C250EC1F46CB253453A34" reason="title">
<typeStatus id="EA8C4EE91A3158AA331AD3DEF4D1569C" type="paratype">Paratypes</typeStatus>
.
</heading>
</paragraph>
<paragraph id="2799893F0198C90A368C776713011876">
<materialsCitation id="EB9BD08719F236C299DC4A65BA51BA5B" collectingDate="2024-04-23" collectionCode="NMNHS-BAS" collectorName="SL &amp; MD &amp; DJ &amp; CD" country="Bulgaria" county="The" elevation="563" latitude="41.7039" location="water fountain at the northeastern exit of Kostino" longLatPrecision="6" longitude="25.3028" municipality="Kardzhali Municipality" specimenCode="BG-NMNHS-HER-000000000552" specimenCount="1" specimenCount-male="1" stateProvince="Kardzhali Province" typeStatus="paratype">
<specimenCode id="942EA35213E69B3387C392DFABA25BC6">BG-NMNHS-HER-000000000552</specimenCode>
(L.
<quantity id="88E44B1AB184A214645AD2BA608718CA" metricMagnitude="-2" metricUnit="m" metricValue="4.16" unit="mm" value="41.6">41.6 mm</quantity>
),
<specimenCount id="0840D25DF9B18B4BA66E36B898E7839B" count="1" type="male">adult male</specimenCount>
collected on
<date id="80FEC6C1F7225D4C8A70A60BA846E31C" value="2024-04-23">
<collectingDate id="0335AD341E8C842B9EA469C731CE4CB4" value="2024-04-23">23 April 2024</collectingDate>
</date>
by
<collectorName id="222395EAC1F742FBE4C18DF0C7F90167">SL</collectorName>
,
<collectorName id="01B7C65DF5C43A7FDCF66803F6DB9FB5">MD</collectorName>
,
<collectorName id="6BDFE4746A82A3F7C580612F038517D9">DJ</collectorName>
and
<collectorName id="69428A3C5AF54BD9273AEE2C0456159B">CD</collectorName>
at the type locality and curated at the
<named-content id="B4477FFD7B2347A60D4648B64759E344" content-type="institution" xlink_href="http://grbio.org/institution/bulgarian-academy-science" xlink_type="simple">National Museum of Natural History of the Bulgarian Academy of Sciences</named-content>
(
<named-content id="718083673F9C02026A6271C85F6B6529" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/bulgarian-academy-science" xlink_title="National Museum of Natural History of the Bulgarian Academy of Sciences">
<emphasis id="67BEF62E11E3AAA9A2B65FE682465296" bold="true">
<collectionCode id="006E93BA0FD4C21D7B07C495E42B0BDB">NMNHS-BAS</collectionCode>
</emphasis>
</named-content>
), Sofia, Bulgaria
</materialsCitation>
;
<materialsCitation id="FF8C1EA1A3C930820862370D9208D44B" collectingDate="2024-04-24" collectionCode="MNHN" collectorName="SL &amp; MD &amp; DJ &amp; CD" country="Bulgaria" elevation="233" latitude="41.6108" location="Lisitsite" longLatPrecision="6" longitude="25.4543" municipality="Kardzhali Municipality" specimenCode="MNHN-RA-2024.0001" specimenCount="1" specimenCount-female="1" stateProvince="Kardzhali Province" typeStatus="paratype">
<specimenCode id="772802E1E47CF63F6C4B410207531642">
<named-content id="30EF9A58FC4CF14FB62B6F1834AD6573" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/museum-national-dhistoire-naturelle-2" xlink_title="Muséum National dHistoire Naturelle Paris">MNHN</named-content>
-RA-2024.0001
</specimenCode>
(L. 46.0 mm),
<specimenCount id="9C1568691FF99994C76CD8B7C53518D3" count="1" type="female">adult female</specimenCount>
collected on
<date id="5ED44026FD87EF1D7360DF7EB713F7E6" value="2024-04-24">
<collectingDate id="8B1950B9758FEF48057FB6323C56DA69" value="2024-04-24">24 April 2024</collectingDate>
</date>
by
<collectorName id="191EC1585D8F25E7F489DF6A4967C387">SL</collectorName>
,
<collectorName id="F392F441878706FC940E3050CE5FF5EC">MD</collectorName>
,
<collectorName id="B5F25EF069BE495C55725ED990719AE7">DJ</collectorName>
and
<collectorName id="A04FFF06492BEB1E3F0F326A5B86F843">CD</collectorName>
in
<location id="7877F83FC12B60F1B2339ECA853E5C26" LSID="urn:lsid:plazi:treatment:7B3AD50B77B55EE9A5EC0F768D307567:7877F83FC12B60F1B2339ECA853E5C26" country="Bulgaria" latitude="41.6108" longLatPrecision="6" longitude="25.4543" municipality="Kardzhali Municipality" name="Lisitsite" stateProvince="Kardzhali Province">Lisitsite</location>
,
<collectingMunicipality id="E3AC399A620D0B15A5563699F5771A31">Kardzhali Municipality</collectingMunicipality>
,
<collectingRegion id="BB6228D99DFC2F716B308230756C74BF" country="Bulgaria" name="Kurdzhali">Kardzhali Province</collectingRegion>
,
<collectingCountry id="3E635B33852A518562CA03AF661FF834" name="Bulgaria">Bulgaria</collectingCountry>
(
<named-content content-type="dwc:verbatimCoordinates" id="NCID0ESZAK" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[25.454300,41.610800]}">
<geoCoordinate id="3D13CA256267D93EA4187211E0B59616" degrees="41.6108" direction="north" orientation="latitude" precision="5" value="41.6108">41.6108 ° N</geoCoordinate>
,
<geoCoordinate id="B56A345CF49275034F7B4D1478AE6C94" degrees="25.4543" direction="east" orientation="longitude" precision="5" value="25.4543">25.4543 ° E</geoCoordinate>
</named-content>
; elevation:
<quantity id="D0527DFC9C9C8C3245DB782D26975B53" metricMagnitude="2" metricUnit="m" metricValue="2.33" unit="m" value="233.0">
<elevation id="EF09B4400193770BFF854DED92C219EE" metricMagnitude="2" metricUnit="m" metricValue="2.33" unit="m" value="233.0">233 m</elevation>
</quantity>
a. s. l.), and curated at
<named-content id="E4CEED7E26919620356F513618FCEBFE" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/museum-national-dhistoire-naturelle-2" xlink_title="Muséum National dHistoire Naturelle Paris">
<collectionCode id="84FA9874374B73D42D0669E20BFB52DF">MNHN</collectionCode>
</named-content>
</materialsCitation>
.
<materialsCitation id="299843ED8D252ED7919F10DFF18B4E61" collectingDate="2024-04-24" collectionCode="NHMW" collectorName="SL &amp; MD &amp; DJ &amp; CD" country="Bulgaria" elevation="753" latitude="41.8565" location="Panichkovo" longLatPrecision="6" longitude="25.1517" municipality="Chernoochene Municipality" specimenCode="NHMW 41962" specimenCount="1" specimenCount-male="1" stateProvince="Kardzhali Province" typeStatus="paratype">
<specimenCode id="1FC0363D6BA49275B9214123573FAC2C">
<named-content id="CFE398B7F6D2D9CC2711D9DBCB2F1DDA" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/naturhistorisches-museum-wien-0" xlink_title="Naturhistorisches Museum Wien">NHMW</named-content>
41962
</specimenCode>
(L.
<quantity id="DC39D67F762CA9A44E3930C27D521E90" metricMagnitude="-2" metricUnit="m" metricValue="3.97" unit="mm" value="39.7">39.7 mm</quantity>
),
<specimenCount id="07C10A008B4BF558D403B6E42E569CFD" count="1" type="male">adult male</specimenCount>
collected on
<date id="235F8BC344DFE6F2C8612094B9CB5E74" value="2024-04-24">
<collectingDate id="FAF89288243B1BCA5D891D4B475D113F" value="2024-04-24">24 April 2024</collectingDate>
</date>
by
<collectorName id="59444DA4AB917C2D2EE9C59A46698CE3">SL</collectorName>
,
<collectorName id="D4B05821D060BB8B80E835E4090E2793">MD</collectorName>
,
<collectorName id="96D25FF0E79346B2E4F8E601CC07B490">DJ</collectorName>
and
<collectorName id="74D864A988F283C19D5DF0F4BF738354">CD</collectorName>
in
<location id="69CB017CA69A65A68519F4B8309BD4C0" LSID="urn:lsid:plazi:treatment:7B3AD50B77B55EE9A5EC0F768D307567:69CB017CA69A65A68519F4B8309BD4C0" country="Bulgaria" latitude="41.8565" longLatPrecision="6" longitude="25.1517" municipality="Chernoochene Municipality" name="Panichkovo" stateProvince="Kardzhali Province">Panichkovo</location>
,
<collectingMunicipality id="89567146EFBC232EFE4C009E21E6F23F">Chernoochene Municipality</collectingMunicipality>
,
<collectingRegion id="7ADA05AACD76668568EC167BFD447C5E" country="Bulgaria" name="Kurdzhali">Kardzhali Province</collectingRegion>
,
<collectingCountry id="354A434820A0580799620356F72B02BD" name="Bulgaria">Bulgaria</collectingCountry>
(
<named-content content-type="dwc:verbatimCoordinates" id="NCID0EE1AK" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[25.151700,41.856500]}">
<geoCoordinate id="068BB85880FEC697568846066BDDEFD4" degrees="41.8565" direction="north" orientation="latitude" precision="5" value="41.8565">41.8565 ° N</geoCoordinate>
,
<geoCoordinate id="C578D67713D5E12534A7837565A2AD75" degrees="25.1517" direction="east" orientation="longitude" precision="5" value="25.1517">25.1517 ° E</geoCoordinate>
</named-content>
; elevation:
<quantity id="275153CD6EF73BC350C05E26239FE2CC" metricMagnitude="2" metricUnit="m" metricValue="7.529999999999999" unit="m" value="753.0">
<elevation id="8D3C52E3C37EBA131B59038C2AF57294" metricMagnitude="2" metricUnit="m" metricValue="7.529999999999999" unit="m" value="753.0">753 m</elevation>
</quantity>
a. s. l.) and curated at
<named-content id="C4DCD94CD2D028E82692B551BB4FFF04" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/naturhistorisches-museum-wien-0" xlink_title="Naturhistorisches Museum Wien">
<collectionCode id="24E73BE03D612A6322C8ECD954C144A9">NHMW</collectionCode>
</named-content>
</materialsCitation>
. These three specimens are depicted in Fig.
<figureCitation id="29469CAB6DBCC2149010FAE80BC3C55F" captionStart="Figure 11" captionStartId="F11" captionText="Figure 11. The paratypes of Bombina variegata rhodopensis subsp. nov. Credit: CD." figureDoi="10.3897/vz.75.e138687.figure11" httpUri="https://binary.pensoft.net/fig/1230624">11</figureCitation>
.
</paragraph>
<caption id="6EBE276CC900FFE5E52004C2D09D90A7" ID-DOI="10.3897/vz.75.e138687.figure11" ID-arpha="0E5CA17B-8E48-5237-8780-8CF8B4AF141F" httpUri="https://binary.pensoft.net/fig/1230624" startId="F11">
<paragraph id="EB9D8EB1FA6482D396AA4BBC0363D28E">
<label id="272FCFD75443AD46307D74CE55950AC5">Figure 11.</label>
</paragraph>
<paragraph id="186F19E1F98E78F7C7381ACC25D8D417">
The paratypes of
<taxonomicName id="CA158529D602D6D6F2C60A53009AC051" authorityName="Lukanov, Denoel, Jablonski &amp; Dufresnes" authorityYear="2025" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="subSpecies" species="variegata" status="subsp. nov." subSpecies="rhodopensis">
<emphasis id="054A655FCF4955DDDB4A3A48E295BA42" italics="true">Bombina variegata rhodopensis</emphasis>
</taxonomicName>
<emphasis id="4EF1A9F555CF555F637E26B82EEA55EB" bold="true">
<taxonomicNameLabel id="2188B68FBD13112DC8DDD25BF5086D98" rank="subSpecies">subsp. nov.</taxonomicNameLabel>
</emphasis>
Credit: CD.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="SECID0EQ2AK" type="diagnosis">
<paragraph id="AECFABCE2056CF011C70B68C84926D66">
<heading id="AD6A64ECE74CB426F959BA1502AC6391" reason="title">Diagnosis.</heading>
</paragraph>
<paragraph id="E327E35E12EBEACFE05C23DF7A6CF5DB">
General characteristics similar to those of the yellow-bellied toad
<taxonomicName id="462D55038842972C70EE0D7DA8A0A256" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="species" species="variegata">
<emphasis id="0570782C50A506F2CE5396666C543C63" italics="true">B. variegata</emphasis>
</taxonomicName>
. It is the sister taxon of the Balkan subspecies
<taxonomicName id="AF9468D2A00F1078B248074FBD6C6CD8" baseAuthorityName="Mertens and Muller" baseAuthorityYear="1940" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="variety" species="variegata" variety="scabra">
<emphasis id="B0EE9832133074796A6E62CF5E1D70AC" italics="true">B. v. scabra</emphasis>
</taxonomicName>
, from which it is distinguished by substantial mitochondrial, nuclear (especially phylogenomic) and morphological divergence. Specifically,
<taxonomicName id="EC3116A0ED888F41389BB2578A36B6E9" authorityName="Lukanov, Denoel, Jablonski &amp; Dufresnes" authorityYear="2025" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="subSpecies" species="variegata" subSpecies="rhodopensis">
<emphasis id="C1E30B4D6470C485D951EF0C1B1811DE" italics="true">B. v. rhodopensis</emphasis>
</taxonomicName>
<emphasis id="CD781DA7FC71BE912DDAC60FE519AB28" bold="true">subsp. nov.</emphasis>
differs from
<taxonomicName id="E068774605D91EB78022A37607BE6CD2" baseAuthorityName="Mertens and Muller" baseAuthorityYear="1940" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="variety" species="variegata" variety="scabra">
<emphasis id="3429F110B3EEF91C13C14D462DCA1095" italics="true">B. v. scabra</emphasis>
</taxonomicName>
by 1.3 % of sequence divergence at
<abbrev id="ABBRID0EQ4AK" xlink_title="16S ribosomal DNA">16 S</abbrev>
, 1.6 % of sequence divergence at
<abbrev id="ABBRID0EU4AK" xlink_title="cytochrome c oxidase subunit I">cox 1</abbrev>
, and 2.4 % of sequence divergence at cyt
<emphasis id="0DA5752F6DAE4F57EDB26334CEDBBD85" italics="true">b</emphasis>
(Table
<tableCitation id="70E3540288A30077198BF8649173D85E" captionStart="Table 1" captionStartId="T1" captionText="Table 1. Pairwise distances between mitochondrial lineages at the DNA barcoding genes 16 S, cox 1 and cyt b. See Fig. 1 for the distribution of the B. variegata lineages." httpUri="http://table.plazi.org/id/FB06175F484E9F93053CB28FDA3B5B1C" tableUuid="FB06175F484E9F93053CB28FDA3B5B1C">1</tableCitation>
). Based on the mitochondrial time tree, the two taxa initiated their divergence during the Early Pleistocene, either around 2.1 or 1.4 Mya, depending on the calibration (Fig.
<figureCitation id="DE21FA11706F852B44CC368090FCA491" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Bayesian phylogeny obtained with BEAST based on full mitogenomes (17,240 kb). Nodes are annotated with dating estimates and their 95 % HPD (node bars) obtained from two calibrations (dark grey: calibration I; light grey: calibration II). Sequence labels indicate accession numbers, geographic origins, taxa and mitochondrial lineages. Asterisks denote full branch support (posterior probabilities of 1.0)." figureDoi="10.3897/vz.75.e138687.figure2" httpUri="https://binary.pensoft.net/fig/1230615">2</figureCitation>
). On the cyt
<emphasis id="44A181569186ECE07B81AABC71265192" italics="true">b</emphasis>
gene, the following nucleotides distinguish
<taxonomicName id="C0DFC8B2EF018955D81AF0565E85A846" authorityName="Lukanov, Denoel, Jablonski &amp; Dufresnes" authorityYear="2025" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="subSpecies" species="variegata" subSpecies="rhodopensis">
<emphasis id="E829EC5D020690224069DB8B4E374407" italics="true">B. v. rhodopensis</emphasis>
</taxonomicName>
<emphasis id="81511C6C985C28DA3C7BA7C8C7C3D1A8" bold="true">subsp. nov.</emphasis>
from any other
<taxonomicName id="F1C241E78504D47BFDF4CF803BFDF536" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="species" species="variegata">
<emphasis id="1FBD7161A50CCF33B2162994EED3F47E" italics="true">B. variegata</emphasis>
</taxonomicName>
subspecies: a “ G ” in the site 469, a “ G ” in the site 726 and an “ A ” in the site 801 (positions relative to the full gene sequence). The new subspecies also features a unique combination of alleles at the nuclear genes ncx 1, rag- 1, rag- 2 and rho (Figs
<figureCitation id="6DDF635B75D1E6691DA9D4FEF4E922ED" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Cloudogram of the nuclear species trees obtained with * BEAST based on four gene fragments (3715 bp) and distinguishing samples representative of the different B. variegata mitochondrial lineages. Node labels indicate Bayesian posterior probabilities; sequence labels indicate taxa and mitochondrial lineages; the thick and thin lines show the average tree root and all the sampled trees, respectively." figureDoi="10.3897/vz.75.e138687.figure3" httpUri="https://binary.pensoft.net/fig/1230616">3</figureCitation>
, S 3), as well as at 4759 RAD loci (Fig.
<figureCitation id="4D4BD12F25B523AA2CCDF4E84B9E6F7A" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Maximum-likelihood phylogeny obtained with IQ-TREE of 4759 concatenated RAD loci (553,608 bp). Colors distinguish the different B. variegata mitochondrial lineages. Sequence labels indicate sample names, geographic origins, taxa and mitochondrial lineages. Asterisks denote robust branch support (bootstrap &gt; 95). The tree is rooted by B. bombina." figureDoi="10.3897/vz.75.e138687.figure4" httpUri="https://binary.pensoft.net/fig/1230617">4</figureCitation>
). Externally,
<taxonomicName id="3A187A0AD13A3B9ED4215DAD78EAC2C4" authorityName="Lukanov, Denoel, Jablonski &amp; Dufresnes" authorityYear="2025" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="subSpecies" species="variegata" subSpecies="rhodopensis">
<emphasis id="2961A11C4FC78FD5A9C2515DABE92CF4" italics="true">B. v. rhodopensis</emphasis>
</taxonomicName>
<emphasis id="DA6E4AD684BCE3E98E04401904152019" bold="true">subsp. nov.</emphasis>
differs from
<taxonomicName id="62C35200B8F94935031C4BC23EC6EDB0" baseAuthorityName="Mertens and Muller" baseAuthorityYear="1940" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="variety" species="variegata" variety="scabra">
<emphasis id="4C0F065F0DA21C0984AE4610B97BA7E2" italics="true">B. v. scabra</emphasis>
</taxonomicName>
by having on average a shorter femur, tibia, and first toe, as well as bigger eyes, noting that these rely on a small sample of
<taxonomicName id="2285D17187D2063622E431D830E0CED9" authorityName="Lukanov, Denoel, Jablonski &amp; Dufresnes" authorityYear="2025" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="subSpecies" species="variegata" subSpecies="rhodopensis">
<emphasis id="6CE47C08073EA5F471977A9E10FEE2BB" italics="true">B. v. rhodopensis</emphasis>
</taxonomicName>
<emphasis id="5D50D19227C15E9BBD1A313CA6BDD104" bold="true">subsp. nov.</emphasis>
specimens (Table
<tableCitation id="74A8607893FAE31440DC2C29795833A0" captionStart="Table 2" captionStartId="T2" captionText="Table 2. Average and standard deviation of the morphological characters measured in 58 specimens representing the Rhodope lineage and the south northwestern Balkan populations of B. v. scabra, distinguishing females and males (in mm). Values for the extended body length dataset (178 specimens) are given on the last line. For character abbreviations, see Methods (section Morphometric analyses in Bombina variegata scabra)." httpUri="http://table.plazi.org/id/3B840D2ACEBED5138148BAFAED33AFB4" tableUuid="3B840D2ACEBED5138148BAFAED33AFB4">2</tableCitation>
). Body sizes are on average similar (
<quantity id="17F153B06EFFF244DCFA2E7E1D59279A" metricMagnitude="-2" metricUnit="m" metricValue="4.36" unit="mm" value="43.6">43.6 mm</quantity>
in
<taxonomicName id="31C094FC7162AA43837E70FD9F455BEC" authorityName="Lukanov, Denoel, Jablonski &amp; Dufresnes" authorityYear="2025" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="subSpecies" species="variegata" subSpecies="rhodopensis">
<emphasis id="29CBCB41EDC6BD8FC53362EF75FB322D" italics="true">B. v. rhodopensis</emphasis>
</taxonomicName>
<emphasis id="F87E115772CC1BD53E8C021974A49B01" bold="true">subsp. nov.</emphasis>
vs.
<quantity id="79C07D9E13F23D941289A4F4BBB9BA8A" metricMagnitude="-2" metricUnit="m" metricValue="4.33" unit="mm" value="43.3">43.3 mm</quantity>
in
<taxonomicName id="58C077E6A3BB1361EC9EB1FB610DD74E" baseAuthorityName="Mertens and Muller" baseAuthorityYear="1940" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="variety" species="variegata" variety="scabra">
<emphasis id="21B1BEE870D56F1DF0E4F368262A555F" italics="true">B. v. scabra</emphasis>
</taxonomicName>
), with higher variation between the sexes in
<taxonomicName id="045E910752BB2478399DA3D3622B7E49" authorityName="Lukanov, Denoel, Jablonski &amp; Dufresnes" authorityYear="2025" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="subSpecies" species="variegata" subSpecies="rhodopensis">
<emphasis id="DF3F3F8CE18A28CDB48112D20167C63E" italics="true">B. v. rhodopensis</emphasis>
</taxonomicName>
<emphasis id="B0DD3FD431BAA6820AC16AF242DA2F6B" bold="true">subsp. nov.</emphasis>
(Fig.
<figureCitation id="3985BAA40F648982B9809569E9A34F08" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Morphological comparison of specimens attributed to the two lineages of B. v. scabra. A PCA on 11 characters corrected by body length distinguishing the males (triangles) and females (circles) of each lineage; B boxplots showing variation in body length (L.) among sexes and lineages. For the list of characters, see Methods (section Morphometric analyses in Bombina variegata scabra)." figureDoi="10.3897/vz.75.e138687.figure5" httpUri="https://binary.pensoft.net/fig/1230618">5</figureCitation>
). From the examined specimens, the sexes appear morphologically dimorphic (Table
<tableCitation id="5D02744B634F7EAB4D4EADCB133A3B94" captionStart="Table 2" captionStartId="T2" captionText="Table 2. Average and standard deviation of the morphological characters measured in 58 specimens representing the Rhodope lineage and the south northwestern Balkan populations of B. v. scabra, distinguishing females and males (in mm). Values for the extended body length dataset (178 specimens) are given on the last line. For character abbreviations, see Methods (section Morphometric analyses in Bombina variegata scabra)." httpUri="http://table.plazi.org/id/3B840D2ACEBED5138148BAFAED33AFB4" tableUuid="3B840D2ACEBED5138148BAFAED33AFB4">2</tableCitation>
, Fig.
<figureCitation id="A2E9DE8ADCE6179E16703EF29318E763" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Morphological comparison of specimens attributed to the two lineages of B. v. scabra. A PCA on 11 characters corrected by body length distinguishing the males (triangles) and females (circles) of each lineage; B boxplots showing variation in body length (L.) among sexes and lineages. For the list of characters, see Methods (section Morphometric analyses in Bombina variegata scabra)." figureDoi="10.3897/vz.75.e138687.figure5" httpUri="https://binary.pensoft.net/fig/1230618">5</figureCitation>
), as also seen in
<taxonomicName id="03BE92209602DA710115F1A481B902E0" baseAuthorityName="Mertens and Muller" baseAuthorityYear="1940" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="variety" species="variegata" variety="scabra">
<emphasis id="21238531FF620E896E4808AD4B121254" italics="true">B. v. scabra</emphasis>
</taxonomicName>
(
<bibRefCitation id="49B75DD04726C6C02F2CBC860F88322E" author="Radojicic" etAl="et al." firstAuthor="Radojicic" journalOrPublisher="Folia Zoologica" pagination="129-140" refId="B138" refString="Radojičić J, Cvetković D, Tomović L, Džukić G, Kalezic ML (2002) Sexual dimorphism in fire-belied toads Bombina spp. from the central Balkans. Folia Zoologica 51: 129140." title="Sexual dimorphism in fire-belied toads Bombina spp. from the central Balkans." volume="51" year="2002">Radojičić et al. 2002</bibRefCitation>
; Fig.
<figureCitation id="6F7CD1FC1DBF392C3DF7C95E9E47A3F0" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Morphological comparison of specimens attributed to the two lineages of B. v. scabra. A PCA on 11 characters corrected by body length distinguishing the males (triangles) and females (circles) of each lineage; B boxplots showing variation in body length (L.) among sexes and lineages. For the list of characters, see Methods (section Morphometric analyses in Bombina variegata scabra)." figureDoi="10.3897/vz.75.e138687.figure5" httpUri="https://binary.pensoft.net/fig/1230618">5</figureCitation>
). Compared to
<taxonomicName id="BF81702A271DE715E39F1C0F695BCB9E" baseAuthorityName="Mertens and Muller" baseAuthorityYear="1940" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="variety" species="variegata" variety="scabra">
<emphasis id="EA274374918C7FF0EEBFFDD30EA4F666" italics="true">B. v. scabra</emphasis>
</taxonomicName>
,
<taxonomicName id="A555AFE0D35EEAED164FB236EFB98583" authorityName="Lukanov, Denoel, Jablonski &amp; Dufresnes" authorityYear="2025" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="subSpecies" species="variegata" subSpecies="rhodopensis">
<emphasis id="76F7F453119728D857BFB25ACF6B7B87" italics="true">B. v. rhodopensis</emphasis>
</taxonomicName>
<emphasis id="C87D9BAED2F6F11B4576839051E034EF" bold="true">subsp. nov.</emphasis>
features a higher average proportion of yellow vs. dark coloration on the ventral side, although with wide overlap (Fig.
<figureCitation id="52086909DE638E695DDF017EA60AE576" captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Proportion of color / dark coloration on the ventral side of specimens previously attributed to B. v. scabra. Bracket indicates significant difference (Kruskal-Wallis test) at P &lt; 0.05. The body area analyzed is emphasized in the two colored (top) and dark (bottom) examples." figureDoi="10.3897/vz.75.e138687.figure7" httpUri="https://binary.pensoft.net/fig/1230620">7</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="SECID0EZEBK" type="etymology">
<paragraph id="F2426994084E124284E8BF5DA1CEEE88">
<heading id="DBD34E83FA77F9DF44533BC36AE7BECB" reason="title">Etymology.</heading>
</paragraph>
<paragraph id="3E9B09B68E806D4CF0BAC905D8AD7D7B">
The name
<emphasis id="1244D921B00A10F3498224C13DE5E9DF" italics="true">rhodopensis</emphasis>
is a Latin toponymic adjective given in reference to the Rhodope Mountains in the southeastern part of the Balkan Peninsula (
<collectingCountry id="9D8707DA637EF7ACF87DBED21A2A2CC6" name="Bulgaria">Bulgaria</collectingCountry>
and
<collectingCountry id="DEF795027342DB60D131F4C6A3447A05" name="Greece">Greece</collectingCountry>
) where the new taxon is mostly distributed. It spotlights a rare case of Rhodope endemism in vertebrates Rhodope endemics are so far known only from plants and invertebrates.
</paragraph>
</subSubSection>
<subSubSection id="SECID0EJFBK" type="Vernacular names">
<paragraph id="E49C49DC5E1117E5A54371C9DED33F9E">
<heading id="AEEBA203B45D74E1990B607E4F9CEAEE" reason="title">Vernacular names.</heading>
</paragraph>
<paragraph id="A9AA42D2FC016E69ED8569A4359C59F3">Rhodope yellow-bellied toad (English), Родопска жълтокоремна бумка (Bulgarian), Κιτρινομπομπίνα της Ροδόπης (Greek), Rodop Sarılı Kurbağa (Turkish), Sonneur à ventre jaune des Rhodopes (French), Kunka žltobruchá rodopská (Slovak).</paragraph>
</subSubSection>
<subSubSection id="SECID0EOFBK" type="distribution">
<paragraph id="188ED3434F5BECE59E260173CC4B6601">
<heading id="EC7A9400E1F084E839CB23C44A28F653" reason="title">Distribution.</heading>
</paragraph>
<paragraph id="0E9481E1C65CD0ED8EB5B716EA6D9924">
<taxonomicName id="78D474D7BADB20E2B617F20A1708CF9E" authorityName="Lukanov, Denoel, Jablonski &amp; Dufresnes" authorityYear="2025" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="subSpecies" species="variegata" subSpecies="rhodopensis">
<emphasis id="75D619767FA719B6DF1F8AE471E3BF9D" italics="true">Bombina v. rhodopensis</emphasis>
</taxonomicName>
<emphasis id="D536B048BDF9E8252F5155791C4050C6" bold="true">subsp. nov.</emphasis>
is essentially restricted to the Rhodope Mountains and their foothills (from sea level up to
<quantity id="36C28D71D509BD19D190AE7943508B4A" metricMagnitude="3" metricUnit="m" metricValue="1.6" unit="m" value="1600.0">1600 m</quantity>
a. s. l.) in southeastern
<collectingCountry id="D9EDADD1820434F707B133CBB0B58D1C" name="Bulgaria">Bulgaria</collectingCountry>
, northeastern
<collectingCountry id="098769A674A28C5FF02C343947485F33" name="Greece">Greece</collectingCountry>
, and the adjacent part of Turkish Thrace (
<bibRefCitation id="4DE9419054617715CC3C869DB669C1C7" refId="B67" refString="Global Biodiversity Information Facility (2024) https://www.gbif.org [accessed on 28 September 2024]." url="https://www.gbif.org" year="2024">Global Biodiversity Information Facility 2024</bibRefCitation>
). In
<collectingCountry id="42AE64FE7137A86590CFFA309517ABB6" name="Greece">Greece</collectingCountry>
, it was documented eastward up to the National Forest Park of Dadia Lefkimi Soufli close to Evros (Maritsa) River which makes the border with
<collectingCountry id="E3C889B27BE77F30B42CF9272B8D5251" name="Turkey">Turkey</collectingCountry>
(
<bibRefCitation id="FC2888826DCAAE19171F8D6EA6D1E6B1" author="Petrov" editor="Beron P &amp; Popov A" firstAuthor="Petrov" journalOrPublisher="2. Biodiversity of Eastern Rhodopes (Bulgaria and Greece) Pensoft and National Museum of Natural History, Sofia" pagination="863-879" refId="B131" refString="Petrov BP (2004) The herpetofauna (Amphibia and Reptilia) of the Eastern Rhodopes (Bulgaria and Greece). In: Beron P, Popov A (Eds) Biodiversity of Bulgaria. 2. Biodiversity of Eastern Rhodopes (Bulgaria and Greece) Pensoft and National Museum of Natural History, Sofia, 863879." title="The herpetofauna (Amphibia and Reptilia) of the Eastern Rhodopes (Bulgaria and Greece)." volumeTitle="Biodiversity of Bulgaria." year="2004">Petrov 2004</bibRefCitation>
; Valakos et al. 2008; Kret and Poirazidis 2015;
<bibRefCitation id="696613111CC1A938D3A7132E3BC4D649" author="Pafilis and Maragkou" firstAuthor="Pafilis" journalOrPublisher="Broken Hill, Nicosia" refId="B128" refString="Pafilis P, Maragkou P (2020) Ατλας Αμφιβίων &amp; Ερπετών της Ελλάδας [Atlas of the Amphibians and Reptiles of Greece]. Broken Hill, Nicosia, 231 pp." title="Ατλας Αμφιβίων &amp; Ερπετών της Ελλάδας [Atlas of the Amphibians and Reptiles of Greece]." year="2020">Pafilis and Maragkou 2020</bibRefCitation>
;
<bibRefCitation id="F4C10EB8BB68A6C3B82D93B6526EA189" accessDate="[accessed 28 September 2024]" author="Strachinis" firstAuthor="Strachinis" journalOrPublisher="Herpetofauna of Greece" refId="B161" refString="Strachinis I (2024) Herpetofauna of Greece. http://www.herpetofauna.gr [accessed 28 September 2024]." url="http://www.herpetofauna.gr" year="2024">Strachinis 2024</bibRefCitation>
). In
<collectingCountry id="41EEA603AC29D0A95DA7291FD61AD384" name="Bulgaria">Bulgaria</collectingCountry>
, population isolates exist east of the Rhodopes, namely in Sakar Mountain and perhaps Strandzha Mountain near the Black Sea (
<bibRefCitation id="7446E9D1283C187BB28092376B8BBF01" author="Boev" etAl="et al." firstAuthor="Boev" refId="B11" refString="Boev Z, Georgiev B, Raychev R, Georgieva U (2008) The wetland zone of the Malkotarnovska River near the town of Malko Tamovo (Burgas District, SE Bulgaria) An example of rich biodiversity (vertebrate fauna) under intensive anthropogenic impact. In: 7 th International Symposium “ Ecology Sustainable Development ”. Proceedings of the symposium, Vratsa 2325 October 2008. Union of Scientists in Bulgaria, Vratsa, 6672." year="2008">Boev et al. 2008</bibRefCitation>
;
<bibRefCitation id="FAAB00E53ECBCEC6F139CE4179AF351B" author="Stojanov" etAl="et al." firstAuthor="Stojanov" journalOrPublisher="Chimaira, Frankfurt am Main" refId="B160" refString="Stojanov A, Tzankov N, Naumov B (2011) Die Amphibien und Reptilien Bulgariens. Chimaira, Frankfurt am Main, 582 pp." title="Die Amphibien und Reptilien Bulgariens." year="2011">Stojanov et al. 2011</bibRefCitation>
). In
<collectingCountry id="BC4DB919D3F9084F6966722388AF9141" name="Turkey">Turkey</collectingCountry>
, it is very rare and restricted to Karacahasan Mountain (Enez District), close to the Evros River (
<bibRefCitation id="CC2464D73171D83BCECD308A8427A2A9" author="Karis" etAl="et al." firstAuthor="Karis" journalOrPublisher="Herpetological Bulletin" pagination="25-27" refId="B90" refString="Kariş M, Veith M, Göçmen B, Oğuz MA, Şener D, Kurt B (2017) Genetic confirmation of the occurrence and notes on the ecology of the yellow-bellied toad, Bombina variegata (L., 1758) (Amphibia: Bombinatoridae) in the European part of Turkey. Herpetological Bulletin 139: 2527." title="Genetic confirmation of the occurrence and notes on the ecology of the yellow-bellied toad, Bombina variegata (L., 1758) (Amphibia: Bombinatoridae) in the European part of Turkey." volume="139" year="2017">Kariş et al. 2017</bibRefCitation>
). The eastern and northern margins of
<taxonomicName id="2DF7BFAC09C15073B2BB3EE0568A2636" authorityName="Lukanov, Denoel, Jablonski &amp; Dufresnes" authorityYear="2025" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="subSpecies" species="variegata" subSpecies="rhodopensis">
<emphasis id="2EF3AB1E69FCE19776E381A5B48AE1A9" italics="true">B. v. rhodopensis</emphasis>
</taxonomicName>
<emphasis id="24DC9DFAB0DC275DD996DF0B393CCF87" bold="true">subsp. nov.</emphasis>
correspond to the shifts from forest hills towards open lowland habitats colonized by the fire-bellied toad (
<taxonomicName id="B998A8ED6A7183FDA3FA03FC0C04007C" baseAuthorityName="Oken" baseAuthorityYear="1816" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="species" species="bombina">
<emphasis id="928F431FDD6078B9DB20FFED138EEB0E" italics="true">B. bombina</emphasis>
</taxonomicName>
), where they probably form hybrid zones. The southern boundary of the range follows the coastal foothills of the Rhodopes (Valakos et al. 2008). According to the
<abbrev id="ABBRID0E4HBK" xlink_title="mitochondrial DNA">mtDNA</abbrev>
barcoding, the transition with
<taxonomicName id="33BCDF829F1C737CE9FCFA051EAEB20B" baseAuthorityName="Mertens and Muller" baseAuthorityYear="1940" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="variety" species="variegata" variety="scabra">
<emphasis id="571074E10AE018A4A706B5606C2160A8" italics="true">B. v. scabra</emphasis>
</taxonomicName>
in the west might follow the Nestos (Mesta) River valley, which separates the Rhodopes from the Rila massif in the north, and from the Pirin massif in the west; the
<abbrev id="ABBRID0EQIBK" xlink_title="mitochondrial DNA">mtDNA</abbrev>
of both subspecies were reported in the middle part of the Nestos River in
<collectingCountry id="DE82009E6D654DDA4310BDA6A993D25D" name="Greece">Greece</collectingCountry>
(Platanias-Pteleas). Nevertheless, many sampling gaps remain in the Pirin, western Rhodopes and southern Rila, so the exact subspecies boundaries shall be fine-tuned by multilocus genotyping.
</paragraph>
</subSubSection>
<subSubSection id="SECID0EUIBK" type="natural history">
<paragraph id="55A9FE878A0A9EE8E4E3602F2F687377">
<heading id="582D4AE0854C71368479A66ED211984E" reason="title">Natural history.</heading>
</paragraph>
<paragraph id="7BA14FF4642AD56FAD0A78DE161AF49F">
The new subspecies inhabits similar habitats as
<taxonomicName id="DD8C76CFC9C0E20E15EF45CD62B2A401" baseAuthorityName="Mertens and Muller" baseAuthorityYear="1940" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="variety" species="variegata" variety="scabra">
<emphasis id="5D552A5F62460C9D318C53D5A600FB55" italics="true">B. v. scabra</emphasis>
</taxonomicName>
, being found in various aquatic sites such as mountain brooks, rivers, ponds, natural and artificial lakes or water-filled ruts and puddles (
<bibRefCitation id="035447DCC42BD9891D764768D165A087" author="Petrov" editor="Beron P &amp; Popov A" firstAuthor="Petrov" journalOrPublisher="2. Biodiversity of Eastern Rhodopes (Bulgaria and Greece) Pensoft and National Museum of Natural History, Sofia" pagination="863-879" refId="B131" refString="Petrov BP (2004) The herpetofauna (Amphibia and Reptilia) of the Eastern Rhodopes (Bulgaria and Greece). In: Beron P, Popov A (Eds) Biodiversity of Bulgaria. 2. Biodiversity of Eastern Rhodopes (Bulgaria and Greece) Pensoft and National Museum of Natural History, Sofia, 863879." title="The herpetofauna (Amphibia and Reptilia) of the Eastern Rhodopes (Bulgaria and Greece)." volumeTitle="Biodiversity of Bulgaria." year="2004">Petrov 2004</bibRefCitation>
). The type series was found in drinking throughs, a valuable habitat for this subspecies as it is for
<taxonomicName id="F62718F1B6E0CC921A4BC17393F5744C" baseAuthorityName="Mertens and Muller" baseAuthorityYear="1940" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="variety" species="variegata" variety="scabra">
<emphasis id="C8881108BA415A853AD4BB3B7819A55E" italics="true">B. v. scabra</emphasis>
</taxonomicName>
in northwestern
<collectingCountry id="B5AF076551AEC86BA78AEC0D8471B7B9" name="Greece">Greece</collectingCountry>
(
<bibRefCitation id="6C7BB673BFC11072F77A0FB3E8765A85" author="Denoel" firstAuthor="Denoel" journalOrPublisher="Herpetozoa" pagination="49-64" refId="B33" refString="Denoël M (2004) Distribution and characteristics of aquatic habitats of newts and yellow-bellied toads in the district of Ioannina (Epirus, Greece). Herpetozoa 17: 4964." title="Distribution and characteristics of aquatic habitats of newts and yellow-bellied toads in the district of Ioannina (Epirus, Greece)." volume="17" year="2004">Denoël 2004</bibRefCitation>
). The breeding season starts in March and lasts until late July. Reports of several thousand eggs laid at the bottom of a slow-flowing stream in Eastern Rhodopes suggested possibly up to 200 per female (
<bibRefCitation id="31059023FE68464823DD13ED02C81438" author="Stojanov" etAl="et al." firstAuthor="Stojanov" journalOrPublisher="Chimaira, Frankfurt am Main" refId="B160" refString="Stojanov A, Tzankov N, Naumov B (2011) Die Amphibien und Reptilien Bulgariens. Chimaira, Frankfurt am Main, 582 pp." title="Die Amphibien und Reptilien Bulgariens." year="2011">Stojanov et al. 2011</bibRefCitation>
). Toads are most active during the day and at dusk, and hibernation occurs on land. Diet studies indicate that Bulgarian populations are mostly insectivorous, preferring water beetles and winged insects, with arachnids and snails having a minor share (
<bibRefCitation id="04D6B13A5D00532715F549884CBA6317" author="Donev" firstAuthor="Donev" refId="B35" refString="Donev A (1984) Studies on the diet of the yellow-bellied toad (Bombina variegata L.). Scientific Works of PU “ P. Hilendarski ” 22 (Biology): 115120." year="1984">Donev 1984</bibRefCitation>
).
<taxonomicName id="18FD78915572ABDD45B6B5743FDB2877" authorityName="Lukanov, Denoel, Jablonski &amp; Dufresnes" authorityYear="2025" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="subSpecies" species="variegata" subSpecies="rhodopensis">
<emphasis id="23A42919B189448D6407944CB9E5591F" italics="true">Bombina v. rhodopensis</emphasis>
</taxonomicName>
<emphasis id="379B958EEE4D76A058AE2E745591C8CE" bold="true">subsp. nov.</emphasis>
displays the anti-predator defense posture (“ Unkenreflex ”), well-known in
<taxonomicName id="954498685E498482F027BF64ED36513C" authorityName="Oken" authorityYear="1816" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="genus">
<emphasis id="BE0764F760C069CBAB74D4F7DB3BA5F2" italics="true">Bombina</emphasis>
</taxonomicName>
toads (Baijger 1980), and which we could observe in
<collectingCountry id="82C18E47CD4A1026BE79EF757F900F0B" name="Bulgaria">Bulgaria</collectingCountry>
(see also
<bibRefCitation id="C8507700A599FA1808F4E8F0B086FEC6" author="Karis" etAl="et al." firstAuthor="Karis" journalOrPublisher="Herpetological Bulletin" pagination="25-27" refId="B90" refString="Kariş M, Veith M, Göçmen B, Oğuz MA, Şener D, Kurt B (2017) Genetic confirmation of the occurrence and notes on the ecology of the yellow-bellied toad, Bombina variegata (L., 1758) (Amphibia: Bombinatoridae) in the European part of Turkey. Herpetological Bulletin 139: 2527." title="Genetic confirmation of the occurrence and notes on the ecology of the yellow-bellied toad, Bombina variegata (L., 1758) (Amphibia: Bombinatoridae) in the European part of Turkey." volume="139" year="2017">Kariş et al. 2017</bibRefCitation>
for
<collectingCountry id="1124FE39F3C1C0529D25008B30ECFB6B" name="Turkey">Turkey</collectingCountry>
).
</paragraph>
</subSubSection>
<subSubSection id="SECID0EELBK" type="Conservation">
<paragraph id="A154E9C973080996718951FE3ADE3CB2">
<heading id="78A9217C2E7635C1D9068A253AE51A92" reason="title">Conservation.</heading>
</paragraph>
<paragraph id="6C04B8B5F20E8D776AAC078757187FEE">
<taxonomicName id="C01392FF4E5A2761C09ADCDE602BF334" authorityName="Lukanov, Denoel, Jablonski &amp; Dufresnes" authorityYear="2025" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="subSpecies" species="variegata" subSpecies="rhodopensis">
<emphasis id="742D0C676BBFB1B711615EF52DB033E4" italics="true">Bombina v. rhodopensis</emphasis>
</taxonomicName>
<emphasis id="C846B62305F255AB9B3AEE8C61D600EF" bold="true">subsp. nov.</emphasis>
was reported widespread over the eastern Rhodopes of
<collectingCountry id="7715F954972A1E59DD2006B305765402" name="Greece">Greece</collectingCountry>
and
<collectingCountry id="2BAC1334CE391E0297FD61CDA45FD658" name="Bulgaria">Bulgaria</collectingCountry>
(
<bibRefCitation id="064965CC4A86CFBA888020AF2A23E149" author="Petrov" editor="Beron P &amp; Popov A" firstAuthor="Petrov" journalOrPublisher="2. Biodiversity of Eastern Rhodopes (Bulgaria and Greece) Pensoft and National Museum of Natural History, Sofia" pagination="863-879" refId="B131" refString="Petrov BP (2004) The herpetofauna (Amphibia and Reptilia) of the Eastern Rhodopes (Bulgaria and Greece). In: Beron P, Popov A (Eds) Biodiversity of Bulgaria. 2. Biodiversity of Eastern Rhodopes (Bulgaria and Greece) Pensoft and National Museum of Natural History, Sofia, 863879." title="The herpetofauna (Amphibia and Reptilia) of the Eastern Rhodopes (Bulgaria and Greece)." volumeTitle="Biodiversity of Bulgaria." year="2004">Petrov 2004</bibRefCitation>
), but due to its much smaller distribution than that of
<taxonomicName id="53ADB340FFC07C53226A99178ED69D04" baseAuthorityName="Mertens and Muller" baseAuthorityYear="1940" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="variety" species="variegata" variety="scabra">
<emphasis id="CFFAC2D6A472517CB7EFAC6D2E770D9E" italics="true">B. v. scabra</emphasis>
</taxonomicName>
, it may be more vulnerable. In
<collectingCountry id="C18BC733E8619802DACEDFEE4EF926CB" name="Greece">Greece</collectingCountry>
, it is abundant above
<quantity id="E9C43805297B40B61258D558A7B16A13" metricMagnitude="2" metricUnit="m" metricValue="2.0" unit="m" value="200.0">200 m</quantity>
(
<bibRefCitation id="E97E55D7F778D24C4F0105FA1A3316C1" accessDate="[accessed 28 September 2024]" author="Strachinis" firstAuthor="Strachinis" journalOrPublisher="Herpetofauna of Greece" refId="B161" refString="Strachinis I (2024) Herpetofauna of Greece. http://www.herpetofauna.gr [accessed 28 September 2024]." url="http://www.herpetofauna.gr" year="2024">Strachinis 2024</bibRefCitation>
). In
<collectingCountry id="5E3A3EB1DBE7ECA9F8E23E243F5CCE9F" name="Turkey">Turkey</collectingCountry>
, it subsists only in a few isolated localities with potentially small population sizes (e. g.,
<bibRefCitation id="F8E51834C653267D3C12251895BA3510" author="Karis" etAl="et al." firstAuthor="Karis" journalOrPublisher="Herpetological Bulletin" pagination="25-27" refId="B90" refString="Kariş M, Veith M, Göçmen B, Oğuz MA, Şener D, Kurt B (2017) Genetic confirmation of the occurrence and notes on the ecology of the yellow-bellied toad, Bombina variegata (L., 1758) (Amphibia: Bombinatoridae) in the European part of Turkey. Herpetological Bulletin 139: 2527." title="Genetic confirmation of the occurrence and notes on the ecology of the yellow-bellied toad, Bombina variegata (L., 1758) (Amphibia: Bombinatoridae) in the European part of Turkey." volume="139" year="2017">Kariş et al. 2017</bibRefCitation>
), and any potential pressure (habitat change, collection) might threaten its persistence in the country. Informed conservation management would benefit from assessing the continuity of the distribution of
<taxonomicName id="B371F25A364573274C3E64A2AC3E8B81" authorityName="Lukanov, Denoel, Jablonski &amp; Dufresnes" authorityYear="2025" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="subSpecies" species="variegata" subSpecies="rhodopensis">
<emphasis id="A8E6B90A5571E968D2A4BC823B528016" italics="true">B. v. rhodopensis</emphasis>
</taxonomicName>
<emphasis id="51F08EB6C96D3F3A2D8E81877A6B6D4D" bold="true">subsp. nov.</emphasis>
across its range, in respect to known occurrence data (e. g., Valakos et al. 2008;
<bibRefCitation id="D4276BD9A251276A89981EBA28311090" refId="B67" refString="Global Biodiversity Information Facility (2024) https://www.gbif.org [accessed on 28 September 2024]." url="https://www.gbif.org" year="2024">Global Biodiversity Information Facility 2024</bibRefCitation>
) and from quantifying habitat loss and threats. As
<taxonomicName id="1B61439E0013D974DCA4DF0419ACE200" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="species" species="variegata">
<emphasis id="3F843D687C23CCD8997819A9794DADFD" italics="true">B. variegata</emphasis>
</taxonomicName>
is listed in the European Union Habitats Directive Annex 2 (= “ Natura 2000 ”), dedicated protected areas could be declared for the conservation of
<taxonomicName id="A457A89BBB77073B384B28DC7529BB37" authorityName="Lukanov, Denoel, Jablonski &amp; Dufresnes" authorityYear="2025" class="Amphibia" family="Bombinatoridae" genus="Bombina" kingdom="Animalia" order="Anura" phylum="Chordata" rank="subSpecies" species="variegata" subSpecies="rhodopensis">
<emphasis id="7A8149D1E78328AA220DC3AC21B0A839" italics="true">B. v. rhodopensis</emphasis>
</taxonomicName>
<emphasis id="1D9580BB3F72659D964C1EAAD9F4CF7E" bold="true">subsp. nov.</emphasis>
</paragraph>
</subSubSection>
</treatment>
</document>