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Vranka Pruhoploutvá, Rod Cottus, Haplotypy, Taxonomie, Fylogeneze, DNA Barcoding

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Vranka Pruhoploutvá, Rod Cottus, Haplotypy, Taxonomie, Fylogeneze, DNA Barcoding MASARYKOVA UNIVERZITA PŘÍRODOVĚDECKÁ FAKULTA ÚSTAV EXPERIMENTÁLNÍ BIOLOGIE Genetická diverzita vranky pruhoploutvé (Cottus poecilopus, Heckel, 1837) ve vodách Karpatského oblouku (hraniční oblast Baltského a Černého moře) Disertační práce Mgr. Eva Marešová Školitel: Mgr. Jan Mendel, Ph.D. Brno 2017 Bibliografický záznam Autor: Mgr. Eva Marešová Přírodovědecká fakulta, Masarykova univerzita Ústav experimentální biologie Název práce: Genetická diverzita vranky pruhoploutvé (Cottus poecilopus, Heckel, 1837) ve vodách Karpatského oblouku (hraniční oblast Baltského a Černého moře) Studijní program: Biologie Studijní obor: Obecná a molekulární genetika Vedoucí práce: Mgr. Jan Mendel, Ph.D. Ústav biologie obratlovců, AV ČR, v. v. i Akademický rok: 2016/2017 Počet stran: 105 Klíčová slova: genetická diverzita, mitochondriální geny, cytochrom b, kontrolní oblast, COI, jaderný intron S7 r-proteinu, hybridizace, mikrosatelity, sekvenace, vranka pruhoploutvá, rod Cottus, haplotypy, taxonomie, fylogeneze, DNA barcoding Bibliographic entry Author: Mgr. Eva Marešová Faculty of Science, Masaryk University Department of Experimental Biology Title of Thesis: Genetic diversity of the Alpine bullhead (Cottus poecilopus, Heckel, 1837) in the Carpathian arc (Baltic and Black Sea basins contact zone) Degree Programme: Biology Field of Study: General and Molecular Genetics Supervisor: Mgr. Jan Mendel, Ph.D. Institute of Vertebrate Biology, Czech Academy of Sciences Academic Year: 2016/2017 Number of Pages: 105 Keywords: genetic diversity, mitochondrial genes, cytochrome b, control region, COI, nuclear intron of the S7 ribosomal protein gene, hybridization, sequencing, Alpine bullhead, genus Cottus, haplotypes, taxonomy, phylogenetics, DNA barcoding ABSTRAKT Předkládaná dizertační práce vychází z malé lokální studie, která poprvé odhaluje vysokou vnitrodruhovou variabilitu vranky pruhoploutvé. Do té doby byl region střední Evropy považován za geneticky homogenní. Dizertace rozšiřuje molekulárně-genetické analýzy na celou hlavní oblast výskytu tohoto druhu a přináší nový pohled na taxonomii vranky a nové poznatky o vnitro- a mezidruhové diverzitě v oblasti Karpatského oblouku. Odhalili jsme dvě vysoce divergentní linie a vymezili území jejich výskytu. Skandinávská linie (I) pokrývá oblast povodí Odry a je pravděpodobně členem North European-West linie ze Švédska, což potvrdila i analýza COI genu (DNA barcoding). Všechny ostatní populace z povodí Dunaje, Visly a Dněstru náleží ke Karpatské linii (II), včetně typové lokality. Uvnitř obou linií byla zjištěna další diferenciace do sublinií Ia - Ib a IIa - IIf, kterou považujeme za projev zvýšené vnitrodruhové variability. Při začlenění linií I a II do euro-asijského komplexu C. poecilopus jsme zjistili nesrovnalosti v současné taxonomii tohoto druhu. K domnělému zástupci evropské vranky byla mylně přiřazena Skandinávská linie s populacemi z povodí Odry. Naopak, Karpatská linie s typovou lokalitou C. poecilopus vytvořila ve fylogenetickém stromě novou, zcela samostatnou větev komplexu. Tuto linii považujeme za korektního zástupce C. poecilopus s. stricto a povodí Dunaje, Visly a Dněstru za areál jejího rozšíření. Vybrané populace vranky pruhoploutvé byly společně s většinou druhů české ichtyofauny zapojeny i do mezinárodní iniciativy FISH-BOL. Na základě porovnání získaných sekvenčních vzorů a sekvencí všech druhů vranek v BoLD databázi bylo zjištěno propojení skandinávského a evropského regionu. Součástí práce byla i analýza hybridních zón z povodí Odry a Moravy. Česká republika je prvním regionem ve střední Evropě, odkud byl popsán, publikován a geneticky charakterizován výskyt nativních hybridních jedinců vranky pruhoploutvé a obecné. Jedná se o hybridní zóny bez reprodukčních bariér, jelikož byli nalezeni, na základě mikrosatelitů, i zpětní kříženci. Dle analýz mtDNA byl zjištěn možný mateřský příspěvek obou vranek - obecné i pruhoploutvé, v hybridních událostech. Byly navrženy některé morfologické znaky k jejich identifikaci v terénu, meristické znaky pro odlišení hybridů od čistých druhů nebyly signifikantní. Pro spolehlivé určení hybridních jedinců byl doporučen komplexní přístup genetické i morfologické analýzy u intenzivně se křížících populací. Pro taxonomický status Skandinávské linie jsme navrhli na základě získaných výsledků tři možné hypotézy. Než bude situace zcela vyjasněna, navrhli jsme u oderské populace zvážit přísnější ochranářský status, pro zamezení možné ztráty nově zjištěné unikátní variability v evropském areálu jejího výskytu. ABSTRACT This dissertation thesis is the continuation of a small local study, which first revealed the high intraspecific variability of the Alpine bullhead. Until then, the Central European region was considered genetically homogeneous. The dissertation extends molecular-genetic analyzes to the entire area of the species and brings new insights into the taxonomy of the bullheads and new insights into the intra and interspecies diversity of the Carpathian Arc. We have revealed two highly divergent lineages and have defined the territory of their occurrence. The Scandinavian lineage (I) covers the Odra basin and is probably a member of the North European-West lineage from Sweden, as confirmed by the DNA barcoding analysis. All other populations of the Danube, Vistula and Dniestr catchment areas belong to the Carpathian lineage (II), including the type locality. Within both lineages, further differentiation was found into sublineages Ia - Ib and IIa - IIf, which we consider to be an expression of increased intraspecific variability. When integrating our lineages into the Euro-Asian C. poecilopus complex, we found discrepancies in current taxonomy of the species. The Scandinavian lineage, with populations from the Odra river basin, was mistakenly associated with the supposed representative of the Alpine bullhead. Conversely, the Carpathian lineage, with the C. poecilopus type locality, created a new, completely separate branch in the phylogenetic tree of the complex. This lineage we consider a correct representative of C. poecilopus s. stricto and the Danube, Vistula and Dniestr basins the areas of its occurrence. Selected populations, together with most species of the Czech Ichthyofauna, have been involved in the international FISH-BOL initiative. Based on the comparison of samples with all species of bullheads in the BoLD database, an interconnection of the Scandinavian and European regions was found. Part of the work was also the analysis of hybrid zones from the Odra and Morava basins. The Czech Republic is the first region in Central Europe where the occurrence of the native hybrids of the Alpine and Common bullheads was described, published and genetically characterized. These are hybrid zones without reproductive barriers as indicated by backcross hybrids based on microsatellites. According to mtDNA analyses, a possible maternal contribution of both – Alpine and Common bullheads, in hybrid events was found. Some morphometric traits have been suggested in order to identify hybrids in the field; meristical signs for distinguishing hybrids from pure species were not significant. However, for the reliable determination of hybrids, a comprehensive approach of genetic and morphological analyses is required in intensively hybridizing populations. For the taxonomic status of the Scandinavian lineage, we suggested three possible hypotheses based on the obtained results from all analyses. Before the situation is fully clarified, we suggested stricter conservation status in order to prevent the possible loss of a newly discovered unique variability in the European area of its occurrence. © Eva Marešová, Masarykova univerzita, 2017 Poděkování Ráda bych poděkovala první školitelce RNDr. Věře Luskové, CSc a doc. Ing. Stanislavu Luskovi, CSc. za podporu a cenné rady do začátku a poté nynějšímu školiteli Mgr. Janu Mendelovi, Ph.D. za ochotu a pomoc při dokončení práce. Manželovi děkuji za trpělivost a vytvoření podmínek k dokončení studia. Tato dizertační práce byla řešena v rámci výzkumných projektů: VaV/SM/6/3/05 Ministerstva životního prostředí ČR a M200930901 finančně podporovaného z Programu interní podpory projektů mezinárodní spolupráce AV ČR, dále projektu VaV SPII2d1/9/07 Ministerstva životního prostředí ČR a projektu Grantové agentury Lesů ČR. Byla podpořena také projektem MSM6215648905 Ministerstva školství ČR. OBSAH 1 ÚVOD .................................................................................................................................. 11 2 PŘEHLED PROBLEMATIKY ........................................................................................ 12 2.1 Stručná biologie a morfologie druhu ....................................................................... 12 2.2 Taxonomie a rozšíření .............................................................................................. 13 2.2.1 Sudeto-karpatský region ............................................................................... 15 2.2.2 Severský region ............................................................................................ 16 2.3 Evoluční diverzifikace sladkovodních vranek ......................................................... 17 2.4 Ochranářský status vranky pruhoploutvé a terminologie ........................................ 17 2.4.1 Jednotky ESU (Evolutionary Significant Unit) a MU (Management Unit) . 18 2.5 Genetická diverzita .................................................................................................. 18 2.5.1 Historie
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