{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T01:16:40Z","timestamp":1787015800183,"version":"build-2736575974"},"reference-count":101,"publisher":"Wiley","issue":"13","license":[{"start":{"date-parts":[[2023,4,7]],"date-time":"2023-04-07T00:00:00Z","timestamp":1680825600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/onlinelibrary.wiley.com\/termsAndConditions#vor"}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Molecular Ecology"],"published-print":{"date-parts":[[2023,7]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>\n                    Within the Balkan Peninsula, topographic and climatic agents have promoted biodiversity and shaped the speciation history of many ectotherms. Here, we targeted an iconic European reptile, the nose\u2010horned viper species\u2010complex (\n                    <jats:italic>Vipera ammodytes<\/jats:italic>\n                    ), and explored its spatial and temporal evolution. We (i) utilized genome\u2010wide single nucleotide polymorphisms to infer genetic structure and build a time\u2010calibrated species\u2010tree, and (ii) applied species distribution modelling with niche\u2010divergence tests among major phylogenomic clades. Geographically structured genetic diversity was found. Cycles of recurrent isolation and expansion during glacial\u2013interglacial periods led to allopatric speciation and to secondary contacts and formation of multiple hybrid zones throughout the Balkan Peninsula. Deep divergence is still detected among populations separated by old and imminent biogeographical barriers (Pindos Mountain Range, the Cyclades islands, etc.), but in most cases speciation is incomplete. At the other end of the speciation continuum, we recognize two well\u2010differentiated lineages, currently lacking any evidence of gene flow; one is distributed in the Northwestern Balkans and the other in the Southeastern Balkans, further expanding into Asia. Despite their split 5 million years ago, there is no evidence of ecological divergence, as speciation probably occurred in niche\u2010pockets of analogous environments. These two lineages probably represent different species, while\n                    <jats:italic>V.\u2009transcaucasiana<\/jats:italic>\n                    does not merit species status. By comparing the genomic phylogenies to an updated mitochondrial one, we propose an evolutionary scenario that resolves all mitonuclear conflicts, according to which the history of the\n                    <jats:italic>V.\u2009ammodytes<\/jats:italic>\n                    species\u2010complex was shaped by complex processes, including a major event of introgressive hybridization with asymmetric mitochondrial capture.\n                  <\/jats:p>","DOI":"10.1111\/mec.16944","type":"journal-article","created":{"date-parts":[[2023,3,31]],"date-time":"2023-03-31T10:40:11Z","timestamp":1680259211000},"page":"3624-3640","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":19,"title":["Genome\u2010wide single nucleotide polymorphisms reveal recurrent waves of speciation in niche\u2010pockets, in Europe's most venomous snake"],"prefix":"10.1111","volume":"32","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5008-6012","authenticated-orcid":false,"given":"Evanthia","family":"Thanou","sequence":"first","affiliation":[{"name":"Ecology Backshop  Loutraki Greece"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Daniel","family":"Jablonski","sequence":"additional","affiliation":[{"name":"Department of Zoology Comenius University in Bratislava  Bratislava Slovakia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Panagiotis","family":"Kornilios","sequence":"additional","affiliation":[{"name":"Department of Biology University of Patras  Patras Greece"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2023,4,7]]},"reference":[{"key":"e_1_2_9_2_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ympev.2016.07.029"},{"key":"e_1_2_9_3_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.marpetgeo.2005.10.004"},{"key":"e_1_2_9_4_1","doi-asserted-by":"publisher","DOI":"10.1111\/mec.15856"},{"key":"e_1_2_9_5_1","doi-asserted-by":"publisher","DOI":"10.1111\/evo.13296"},{"key":"e_1_2_9_6_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pcbi.1003537"},{"key":"e_1_2_9_7_1","doi-asserted-by":"publisher","DOI":"10.1111\/mec.15836"},{"key":"e_1_2_9_8_1","doi-asserted-by":"publisher","DOI":"10.21425\/F5FBG44158"},{"key":"e_1_2_9_9_1","first-page":"1917","article-title":"Inferring species trees directly from biallelic genetic markers: Bypassing gene trees in a full coalescent analysis","volume":"29","author":"Bryant D.","year":"2012","journal-title":"Molecular Phylogenetics and Evolution"},{"key":"e_1_2_9_10_1","first-page":"2107","article-title":"Mitochondrial DNA phylogeography of the polytypic North American rat snake Elaphe obsoleta: A critique of the subspecies concept","volume":"54","author":"Burbrink F. T.","year":"2000","journal-title":"Evolution"},{"key":"e_1_2_9_11_1","first-page":"319","article-title":"Note eco\u2010morfologiche su alcune species ofidiche egee, con particolare riferimento alle popolazioni delle cicladi centro\u2010orientali (Reptilia)","volume":"34","author":"Cattaneo A.","year":"2010","journal-title":"Naturalista Siciliano"},{"key":"e_1_2_9_12_1","doi-asserted-by":"publisher","DOI":"10.1038\/38229"},{"key":"e_1_2_9_13_1","doi-asserted-by":"publisher","DOI":"10.1186\/s12862-022-01994-y"},{"key":"e_1_2_9_14_1","doi-asserted-by":"publisher","DOI":"10.1093\/bioinformatics\/btu530"},{"key":"e_1_2_9_15_1","doi-asserted-by":"publisher","DOI":"10.2298\/ABS181020028C"},{"key":"e_1_2_9_16_1","first-page":"263","article-title":"Paleogeography, geodynamic processes and event stratigraphy during the late Cenozoic of the Aegean area","volume":"85","author":"Dermitzakis M. D.","year":"1990","journal-title":"Atti della Accademia Nazionale dei Lincei"},{"key":"e_1_2_9_17_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.tox.2021.152724"},{"key":"e_1_2_9_18_1","doi-asserted-by":"publisher","DOI":"10.1186\/s12862-018-1179-0"},{"key":"e_1_2_9_19_1","doi-asserted-by":"publisher","DOI":"10.1093\/bioinformatics\/btu121"},{"key":"e_1_2_9_20_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-294X.2005.02553.x"},{"key":"e_1_2_9_21_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.tig.2012.03.009"},{"key":"e_1_2_9_22_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1558-5646.1985.tb00420.x"},{"key":"e_1_2_9_23_1","doi-asserted-by":"publisher","DOI":"10.1093\/biomethods\/bpab017"},{"key":"e_1_2_9_24_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.quascirev.2012.03.002"},{"key":"e_1_2_9_25_1","first-page":"1005","article-title":"Fossil mammals resolve regional patterns of Eurasian climate change over 20 million years","volume":"4","author":"Fortelius M.","year":"2002","journal-title":"Evolutionary Ecology Research"},{"key":"e_1_2_9_26_1","doi-asserted-by":"publisher","DOI":"10.1163\/15685381-bja10007"},{"key":"e_1_2_9_27_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1502-3931.2008.00097.x"},{"key":"e_1_2_9_28_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ympev.2004.11.004"},{"key":"e_1_2_9_29_1","unstructured":"GBIF. (2021).Vipera ammodytes in GBIF secretariat: GBIF backbone taxonomy. GBIF occurrence download fromhttps:\/\/doi.org\/10.15468\/dl.58wq2v Retrieved March 1 2021 fromGBIF.org"},{"key":"e_1_2_9_30_1","volume-title":"Endoglyphs and other major venomous snakes of the world. A checklist","author":"Golay P.","year":"1993"},{"key":"e_1_2_9_31_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-1-4020-2854-0"},{"key":"e_1_2_9_32_1","doi-asserted-by":"publisher","DOI":"10.1093\/sysbio\/syq010"},{"key":"e_1_2_9_33_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ympev.2013.05.004"},{"key":"e_1_2_9_34_1","first-page":"81","volume-title":"Handbuch der Reptillien und Amphibien Europas, IIB Schlangen (Serpentes) III Viperidae","author":"Heckes U.","year":"2005"},{"key":"e_1_2_9_35_1","doi-asserted-by":"publisher","DOI":"10.1038\/35016000"},{"key":"e_1_2_9_36_1","doi-asserted-by":"publisher","DOI":"10.1002\/joc.1276"},{"key":"e_1_2_9_37_1","doi-asserted-by":"publisher","DOI":"10.1038\/267399a0"},{"key":"e_1_2_9_38_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.quascirev.2011.08.016"},{"key":"e_1_2_9_39_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.quascirev.2010.06.032"},{"key":"e_1_2_9_40_1","doi-asserted-by":"publisher","DOI":"10.1016\/B978-0-12-800049-6.00077-9"},{"key":"e_1_2_9_41_1","doi-asserted-by":"publisher","DOI":"10.1111\/geb.12716"},{"key":"e_1_2_9_42_1","doi-asserted-by":"publisher","DOI":"10.5194\/cp-10-607-2014"},{"key":"e_1_2_9_43_1","doi-asserted-by":"publisher","DOI":"10.1093\/biolinnean\/blaa224"},{"key":"e_1_2_9_44_1","doi-asserted-by":"publisher","DOI":"10.1186\/s12862-016-0669-1"},{"key":"e_1_2_9_45_1","unstructured":"Jarvis A. Reuter H. I. Nelson A. &Guevara E.(2008).Hole\u2010filled SRTM for the globe Version 4. Retrieved April 25 2021 fromhttp:\/\/www.cgiarcsi.org\/data\/srtm\u201090mdigital\u2010elevation\u2010database\u2010v4\u20101"},{"key":"e_1_2_9_46_1","doi-asserted-by":"publisher","DOI":"10.1093\/bioinformatics\/btx025"},{"key":"e_1_2_9_47_1","doi-asserted-by":"publisher","DOI":"10.1127\/0372-8854\/2009\/0053S1-0169"},{"key":"e_1_2_9_48_1","doi-asserted-by":"publisher","DOI":"10.1111\/zsc.12018"},{"key":"e_1_2_9_49_1","doi-asserted-by":"publisher","DOI":"10.1111\/1755-0998.12387"},{"key":"e_1_2_9_50_1","doi-asserted-by":"publisher","DOI":"10.1111\/zsc.12243"},{"key":"e_1_2_9_51_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ympev.2011.11.035"},{"key":"e_1_2_9_52_1","doi-asserted-by":"publisher","DOI":"10.1111\/jbi.13524"},{"key":"e_1_2_9_53_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.epsl.2009.12.020"},{"key":"e_1_2_9_54_1","doi-asserted-by":"publisher","DOI":"10.1111\/jbi.12057"},{"key":"e_1_2_9_55_1","doi-asserted-by":"publisher","DOI":"10.1017\/S0003598X00083733"},{"key":"e_1_2_9_56_1","doi-asserted-by":"publisher","DOI":"10.1093\/bioinformatics\/btm404"},{"key":"e_1_2_9_57_1","doi-asserted-by":"publisher","DOI":"10.1111\/geb.13075"},{"key":"e_1_2_9_58_1","volume-title":"True vipers: Natural history and toxinology of Old World vipers","author":"Mallow D.","year":"2003"},{"key":"e_1_2_9_59_1","doi-asserted-by":"publisher","DOI":"10.1093\/biolinnean\/blx038"},{"key":"e_1_2_9_60_1","doi-asserted-by":"publisher","DOI":"10.1111\/jzs.12534"},{"key":"e_1_2_9_61_1","doi-asserted-by":"publisher","DOI":"10.1111\/jbi.13861"},{"key":"e_1_2_9_62_1","doi-asserted-by":"publisher","DOI":"10.1111\/jzs.12115"},{"key":"e_1_2_9_63_1","volume-title":"Snake species of the world: A taxonomic and geographic reference","author":"McDiarmid R. W.","year":"1999"},{"key":"e_1_2_9_64_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.zool.2018.01.005"},{"key":"e_1_2_9_65_1","doi-asserted-by":"publisher","DOI":"10.1093\/molbev\/mst024"},{"key":"e_1_2_9_66_1","doi-asserted-by":"publisher","DOI":"10.1093\/molbev\/msu300"},{"key":"e_1_2_9_67_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-2699.2004.01222.x"},{"key":"e_1_2_9_68_1","doi-asserted-by":"publisher","DOI":"10.1002\/tax.602007"},{"key":"e_1_2_9_69_1","first-page":"103","article-title":"Vipers of Caucasus: Taxonomic considerations","volume":"8","author":"Nilson G.","year":"1999","journal-title":"Kaupia (Darmstadt)"},{"key":"e_1_2_9_70_1","doi-asserted-by":"publisher","DOI":"10.1534\/genetics.112.145037"},{"key":"e_1_2_9_71_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0037135"},{"key":"e_1_2_9_72_1","doi-asserted-by":"publisher","DOI":"10.1111\/mec.15832"},{"key":"e_1_2_9_73_1","unstructured":"Phillips S. J. Dud\u00edk M. &Schapire R. E.(2017).Maxent software for modeling species niches and distributions (Version 3.4.1). Retrieved June 17 2021 fromhttp:\/\/biodiversityinformatics.amnh.org\/open_source\/maxent\/"},{"key":"e_1_2_9_74_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pgen.1002967"},{"key":"e_1_2_9_75_1","doi-asserted-by":"publisher","DOI":"10.1144\/SP433.11"},{"key":"e_1_2_9_76_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-294X.2004.02396.x"},{"key":"e_1_2_9_77_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ympev.2018.03.021"},{"key":"e_1_2_9_78_1","unstructured":"R Development Core Team. (2020).R: A language and environment for statistical computing. R Foundation for Statistical Computing. Accessed September 02 2020. Retrieved fromhttp:\/\/www.R\u2010project.org"},{"key":"e_1_2_9_79_1","unstructured":"Rambaut A. &Drummond A. J.(2009).Tracer ver. 1.5. [Computer software]. Institute of Evolutionary Biology University of Edinburgh. Accessed March 24 2020. Retrieved fromhttp:\/\/tree.bio.ed.ac.uk\/software\/tracer"},{"key":"e_1_2_9_80_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature08365"},{"key":"e_1_2_9_81_1","doi-asserted-by":"publisher","DOI":"10.1163\/15685381-00002935"},{"key":"e_1_2_9_82_1","doi-asserted-by":"publisher","DOI":"10.11646\/zootaxa.3795.3.6"},{"key":"e_1_2_9_83_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ympev.2016.07.005"},{"key":"e_1_2_9_84_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-019-41290-2"},{"key":"e_1_2_9_85_1","first-page":"157","article-title":"Pleistocene glacial history of Mount Olympus, Greece: Neotectonic uplift, equilibrium line elevations, and implications for climatic change","volume":"409","author":"Smith G. W.","year":"2006","journal-title":"Geological Society of America Special Papers"},{"key":"e_1_2_9_86_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ympev.2019.05.026"},{"key":"e_1_2_9_87_1","doi-asserted-by":"publisher","DOI":"10.1093\/sysbio\/syy006"},{"key":"e_1_2_9_88_1","doi-asserted-by":"publisher","DOI":"10.1111\/zsc.12471"},{"key":"e_1_2_9_89_1","volume-title":"PAUP*. Phylogenetic analysis using parsimony (* and other methods). Version 4","author":"Swofford D. L.","year":"2003"},{"key":"e_1_2_9_90_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41559-018-0777-y"},{"key":"e_1_2_9_91_1","doi-asserted-by":"publisher","DOI":"10.1163\/15685381-00002947"},{"key":"e_1_2_9_92_1","doi-asserted-by":"publisher","DOI":"10.1093\/cz\/zoz018"},{"key":"e_1_2_9_93_1","doi-asserted-by":"publisher","DOI":"10.1093\/biolinnean\/blab074"},{"key":"e_1_2_9_94_1","first-page":"191","article-title":"Systematics of the nose\u2010horned viper (Vipera ammodytes, Linnaeus, 1758)","volume":"16","author":"Tomovi\u0107 L.","year":"2006","journal-title":"Herpetological Journal"},{"key":"e_1_2_9_95_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1095-8312.2010.01513.x"},{"key":"e_1_2_9_96_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ympev.2007.11.002"},{"key":"e_1_2_9_97_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1600-0587.2009.06142.x"},{"key":"e_1_2_9_98_1","doi-asserted-by":"publisher","DOI":"10.1186\/1742-9994-10-13"},{"key":"e_1_2_9_99_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ympev.2010.04.030"},{"key":"e_1_2_9_100_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ympev.2008.08.019"},{"key":"e_1_2_9_101_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ympev.2015.02.002"},{"key":"e_1_2_9_102_1","doi-asserted-by":"publisher","DOI":"10.1186\/s12862-016-0647-7"}],"container-title":["Molecular Ecology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1111\/mec.16944","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,9,28]],"date-time":"2023-09-28T05:14:49Z","timestamp":1695878089000},"score":1,"resource":{"primary":{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/10.1111\/mec.16944"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,4,7]]},"references-count":101,"journal-issue":{"issue":"13","published-print":{"date-parts":[[2023,7]]}},"alternative-id":["10.1111\/mec.16944"],"URL":"https:\/\/doi.org\/10.1111\/mec.16944","archive":["Portico"],"relation":{},"ISSN":["0962-1083","1365-294X"],"issn-type":[{"value":"0962-1083","type":"print"},{"value":"1365-294X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2023,4,7]]},"assertion":[{"value":"2022-07-26","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2023-03-27","order":1,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2023-04-07","order":2,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}]}}