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Evoluční Historie a Rozšíření Bělozubek Druhů Crocidura Hirta a Crocidura Flavescens Ve Východní Africe

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Evoluční Historie a Rozšíření Bělozubek Druhů Crocidura Hirta a Crocidura Flavescens Ve Východní Africe PŘÍRODOVĚDECKÁ FAKULTA EVOLUČNÍ HISTORIE A ROZŠÍŘENÍ BĚLOZUBEK DRUHŮ CROCIDURA HIRTA A CROCIDURA FLAVESCENS VE VÝCHODNÍ AFRICE Diplomová práce Autor práce: Ing. Bc. Michal Bureš Vedoucí práce: Mgr. Adam Konečný, Ph.D. Ústav botaniky a zoologie Brno 2019 Bibliografický záznam Autor: Ing. Bc. Michal Bureš Přírodovědecká fakulta, Masarykova univerzita Ústav botaniky a zoologie Evoluční historie a rozšíření bělozubek druhů Název práce: Crocidura hirta a Crocidura flavescens ve východní Africe Studijní program: Ekologická a evoluční biologie Studijní obor: Zoologie Vedoucí práce: Mgr. Adam Konečný, Ph.D. Akademický rok: 2018/2019 Počet stran: 80+30 stran Klíčová slova: Soricidae, Crocidura, Crocidura hirta, Crocidura flavescens, Afrika, bělozubky, evoluční historie, distribuce, molekulární fylogeneze, cytochrome b, jaderná DNA Bibliographic Entry Author: Ing. Bc. Michal Bureš Faculty of Science, Masaryk University Department of botany and zoology Evolutionary history and distribution of the species Title of Thesis: Crocidura hirta and Crocidura flavescens species in Eastern Africa Degree programme: Ecologicaly and evolutionary biology Field of Study: Zoology Supervisor: Mgr. Adam Konečný, Ph.D. Academic Year: 2018/2019 Number of Pages: 80+30 pages Keywords: Soricidae, Crocidura, Crocidura hirta, Crocidura flavescens, Africa, white-toothed shrews, evolutionary history, distribution, molecular phylogeny, cytochrome b, nuclear DNA 4 Obsah Abstrakt Bělozubky rodu Crocidura jsou v Africe zastoupeny velkým množstvím druhů a jejich fylogenetické vztahy jsou dosud nevyjasněné. Afričtí rejskovití jsou přitom v Africe přenašeči nebezpečných virů a nemocí. Na druhou stranu mohou rejskovití posloužit jako vhodný evoluční model, protože mají vysokou mutační rychlost mitochondriální DNA a jsou vázáni na určitý typ stanoviště. Mají rychlou výměnu generací, omezenou schopnost migrovat a překonávat význačné bariéry (pohoří, velké řeky). A právě jejich studium může doplnit mozaiku vývoje afrických savan a pralesů během několika posledních milionů let. Východoafrické savany v průběhu dob ledových a meziledových procházely dynamickým vývojem, kdy probíhala pulzace savan, která má vliv na současné složení populací savanových hlodavců, kopytníků, primátů a také rejskovitých. V současné době jsou ve východní a jižní Africe široce rozšířeny dva sesterské druhy rejskovitých, Crocidura hirta a Crocidura flavescens. Kvůli jejich značnému rozšíření a fenotypové plasticitě byla v minulosti popsána řada druhů, které v současné době slouží jako synonymní pojmenování pro tyto druhy, ovšem molekulární analýza jednotlivých linií dosud chyběla. Na základě molekulárních dat byly analyzovány vztahy mezi jednotlivými klady linie C. hirta a C. flavescens, datování rozkolu jednotlivých kladů a také porovnání evoluční historie s dalšími savanovými druhy savců. V naší studii se ukázalo, že se pravděpodobně jedná o druhový komplex minimálně 3 druhů, který se vyskytuje od Etiopie až po jižní Afriku. Poslední žijící předek této linie, žil na začátku pleistocénu, kdy sušší a proměnlivější klima pravděpodobně stálo za rozvojem této linie po savanách východní Afriky. Na změny klimatu během pleistocénu zástupci linie C. hirta a C. flavescens reagovali podobně jako ostatní savanové druhy – únikem do refugií, kde proběhla divergence jednotlivých populací, což je patrné na molekulárních datech. Abstract The white-toothed shrew of the genus Crocidura are represented in Africa by a large number of species and their phylogenetic relationships are still unclear. But african shrews are vectors of dangerous viruses and diseases in Africa. On the other hand, shrews can serve as a suitable evolutionary model because they have a high mutational rate of mitochondrial DNA and are bound to a specific type of habitat. They have a rapid exchange of generations, limited ability to migrate and overcome significant barriers (mountain ranges, large rivers). And it is their studies that can complement the mosaic of evolution of African savannas and forest development over the last few million years. East African savannas have undergone a dynamic development during the glacials and interglacials, with the pulsation of the savannas that affects the current composition of savannah rodent populations, ungulates, primates, and shrews. At present, two sister shrew species, Crocidura hirta and Crocidura flavescens, are widespread in eastern and southern Africa. Due to their widespread and phenotypic plasticity, a number of species have been described in the past, which currently serve as synonymous names for these species, but molecular analysis of individual lines has been lacking. On the basis of molecular data, the relationships between the individual clades of the C. hirta and C. flavescens were analyzed, as well as the dating of the split individual clades and the evolutionary history with other mammalian savannah species. Our study has shown that species, C. hirta and C. flavescens are probably a species complex of at least 3 species, occurring from Ethiopia to South Africa. The last living ancestor of this line, lived at the beginning of the Pleistocene, when the climate become drier and more variable. And that was probably behind the development of this line on the savannas of East Africa. Climatic changes during the Pleistocene, similarly to other savannah species, the representatives of the C. hirta and C. flavescens clades reacted by escaping to refugia where individual populations diverged, as shown by molecular data. 8 Obsah Poděkování Na tomto místě bych chtěl poděkovat vedoucímu své diplomové práce Mgr. Adamovi Konečnému, PhD. za jeho trpělivost a pomoc. Konzultantovi doc. Mgr. et Mgr. Josefu Bryjovi, Ph.D. za rady. Dále bych chtěl poděkovat Mgr. Daniele Mizerovské za pomoc při analýzách a celkovou podporu. Rád bych poděkoval také své rodině, zejména manželce a synovi, bez kterých bych diplomku nedopsal vůbec, nebo mnohem rychleji. Prohlášení Prohlašuji, že jsem svoji diplomovou práci vypracoval samostatně s využitím informačních zdrojů, které jsou v práci citovány. Brno, 2. května 2019 …………………………………. Ing. Bc. Michal Bureš 10 Obsah Obsah 1 Úvod a cíle práce 13 1.1 Úvod ....................................................................................................................................... 13 1.2 Cíle práce .............................................................................................................................. 15 2 Úvod do problematiky 16 2.1 Východní Afrika ................................................................................................................. 16 2.1.1 Východoafrické savany ......................................................................................... 21 2.1.2 Klimatické oscilace během pleistocénu ve východní Africe ................... 23 2.1.3 Fylogeografie hlodavců, primátů a kopytníků na východoafrických savanách ...................................................................................................................................... 25 2.2 Linie velkých africký bělozubek kolem komplexu druhů C. olivieri .............. 27 2.2.1 Fylogenetické vztahy v rámci Crocidura olivieri sensu lato ................... 30 2.3 Crocidura hirta ................................................................................................................... 33 2.3.1 Historie fylogenetických vztahů v rámci druhu Crocidura hirta .......... 33 2.3.2 Habitat......................................................................................................................... 40 2.3.3 Ochrana....................................................................................................................... 40 2.4 Crocidura flavescens ......................................................................................................... 41 2.4.1 Historie fylogenetických vztahů v rámci druhu Crocidura flavescens 41 2.4.2 Habitat......................................................................................................................... 46 2.4.3 Ochrana....................................................................................................................... 46 2.5 Molekulární datování v rámci rodu Crocidura ...................................................... 47 3 Materiál a metodika 50 3.1 Terénní výzkum a vzorkování ..................................................................................... 50 3.2 Laboratorní výzkum ........................................................................................................ 50 3.3 Fylogenetická analýza ..................................................................................................... 51 3.3.1 Fylogenetická analýza na základě mitochondriální DNA ........................ 52 3.3.2 Haplotypové sítě ..................................................................................................... 53 3.3.3 Delimitace do druhů na základě mitochondriálního genu a jaderných markerů ....................................................................................................................................... 53 3.3.4 Odhad času divergence jednotlivých linií ..................................................... 54 4 Výsledky 56 4.1 Molekulární analýza Crocidura hirta a Crocidura flavescens ........................... 56 4.1.1 Fylogenetická analýza druhů Crocidura hirta a Crocidura flavescens 56 4.1.2 Haplotypové sítě ....................................................................................................
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