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Genetická Struktura Populací Dvou Druhů Sympatricky Se Vyskytujících Luňáků (Milvus Spp.) Diplomová Práce Matej Dolinay

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Genetická Struktura Populací Dvou Druhů Sympatricky Se Vyskytujících Luňáků (Milvus Spp.) Diplomová Práce Matej Dolinay MASARYKOVA UNIVERZITA ŘÍRODOVĚDECKÁ FAKULTA P ÚSTAV BOTANIKY A ZOOLOGIE Genetická struktura populací dvou druhů sympatricky se vyskytujících luňáků (Milvus spp.) Diplomová práce Matej Dolinay Vedoucí práce: Prof. MVDr. Ivan Literák, CSc. Brno 2015 Bibliografický záznam Autor: Matej Dolinay Přírodovědecká fakulta, Masarykova univerzita Ústav botaniky a zoologie Genetická struktura populací dvou druhů Název práce: sympatricky se vyskytujících luňáků (Milvus spp.) Studijní program: Biologie Studijní obor: Zoologie Vedoucí práce: Prof. MVDr. Ivan Literák, CSc. Akademický rok: 2014/2015 Počet stran: 67 Klíčová slova: Luňáci; Milvus; Hybridizace; Genetická struktura populace; Sympatrické druhy; Speciace Bibliographic Entry Author Matej Dolinay Faculty of Science, Masaryk University Department of Botany and Zoology Title of Thesis: Genetic structure of populations in two sympatric species of kites (Milvus spp.) Degree programme: Biology Field of Study: Zoology Supervisor: Prof. MVDr. Ivan Literák, CSc. Academic Year: 2014/2015 Number of Pages: 67 Keywords: Kites; Milvus; Hybridization; Genetic structure of population; Sympatric species; Speciation Abstrakt U dvou sympatricky se vyskytujících evolučně mladých druhů může docházet k hybridizaci, která ovlivňuje genetickou variabilitu jejich populací. Tato práce se zabývá genetickou strukturu populací dvou druhů luňáků (Milvus spp.) ve střední Evropě. Luňák červený (Milvus milvus) a luňák hnědý (Milvus migrans) jsou od sebe dobře rozlišitelné druhy na základě morfologie, chování a migrační ekologie. Navzdory tomu byla v minulosti v několika studiích dokumentována existence smíšených hnízdících párů a hybridních jedinců. Molekulární data byla v této práci vůbec poprvé použita pro experimentální potvrzení hybridizace u těchto dvou druhů luňáků. Datový soubor obsahoval sekvence 266 jedinců, kteří byli analyzováni na dvou mitochondriálních lokusech (CO1 a Cytb). Byli nalezeni tři hybridi. Menší diverzita haplotypů byla zjištěna u druhu M. milvus, což je pravděpodobně důsledek menšího areálu rozšíření a prudkých poklesů populační velikosti v minulosti. U obou druhů bylo nalezeno jenom několik transverzí, což ukazuje na recentní vznik jejich polymorfismů. Haplotypy zastoupené u nejvíce jedinců byly rovnoměrně rozloženy ve všech regionech a nebyla u nich nalezena žádná geografická struktura. Interspecifická vzdálenost mezi těmito dvěma druhy je velice malá a v souladu s hypotézou o jejich nedávné divergenci. Hybridizace mezi druhy M. migrans a M. milvus je pořád možná a skutečně k ní dochází. Abstract The hybridization between two evolutionary young species can occur in areas where they are sympatric and influences the genetic variability of their populations. In this thesis, I investigated the genetic structure of Central European populations of two species of kites (Milvus spp.). The Red Kite (M. milvus) and Black Kite (M. migrans) are clearly distinguished species by morphology, behaviour and migration ecology. However, the existence of hybrid individuals and mixed breeding pairs was documented by previous observational studies. The molecular data were used to confirm experimentally the hybridization between M. migrans and M. milvus for the first time here. The dataset comprised of 266 individuals analysed for two mitochondrial loci (CO1 and Cytb). Three hybrid individuals were found. The haplotype diversity was lower in M. milvus, which is probably caused by bottlenecks in the past and smaller distribution than M. migrans. Only a small number of transversions was found for both species, indicating a recent origin of polymorphisms. The most abundant haplotypes were equally present in all regions and no geographical structure was found. Only a minute interspecific distance was found between these two species and it supports the hypothesis that these species have diverged very recently. The hybridization between M. milvus and M. migrans is still possible and it also ocasionally happens. Poděkování Na tomto místě bych chtěl poděkovat Prof. MVDr. Ivanu Literákovi, CSc. za zkušené vytvoření dizajnu studie, její financování a cenné rady ohledně základních postupů pro praktické uchopení vědeckého projektu. Dále děkuji RNDr. Petru Henebergovi, Ph.D., který mi poskytl zázemí v molekulární laboratoři, neocenitelné praktické zkušenosti a znalosti o použití molekulárních technik a následných analýz dat. Děkuji i Mgr. Jiřímu Bizosovi, který byl mým společníkem v laboratoři a vždy mi ochotně pomohl s jakýmkoliv pracovním úkonem. Mé poděkování patří také Mgr. Adamovi Konečnému, Ph.D. za pomoc při zařizování formalit a oficiální zastřešení studie na Masarykově univerzitě. Rád bych poděkoval také Martinu Macečkovi za pomoc s vytvořením geografických map s hnízdy luňáků. Na závěr nesmím zapomenout na všechny kroužkovatele, kteří přispěli svým dílem k získání úctyhodného množství vzorků pro tuto studii, jmenovitě děkuji Hynkovi Matušíkovi, Karlu Makoňovi, Jakubovi Mrázovi, Václavovi Šenovi, Vladimírovi Pečeňákovi, Thomasovi Pfeifferovi a Winfriedovi Nachtigallovi. Za podporu děkuji svým rodičům a partnerce. Prohlášení Prohlašuji, že jsem svoji diplomovou práci vypracoval samostatně s využitím informačních zdrojů, které jsou v práci citovány. Brno 04. měsíce 2015 ……………………………… Jméno Příjmení TABLE OF CONTENTS 1. Introduction…………………..………………………………………………..9 1.1. Speciation and recognition of species and subspecies……….…......….9 1.2. Introduction to ecology and migration of kites (Milvus spp.).….............12 1.3. Systematics of kites (Milvus spp.)…………………………………....…...14 1.4. Kites on Cape Verde islands……………………………..……….…...….17 1.5. Documented cases of hybridization in kites…………………...……...…18 1.6. The first case of mixed breeding pair of Red Kite and Black Kite in Czech Republic……………………………………..………………………….…….…..20 1.7. Genetic structure of populations………………………………..….....…..25 1.8. Aims……………………………………………………………………..…...26 2. Materials and methods…………………………….…….….…….………..27 2.1. Sample collection and DNA extraction…………………………..…….....27 2. 2. PCR and sequencing………………………………………………....…...29 2. 3. MtDNA sequence analyses and phylogeny…………………….…...…..42 3. Results…………………………………….…………….……………………..46 3. 1. Analyses of CO1 locus………………………………….………...…….…46 3. 2. Analyses of Cytb locus……………………………………………….……50 4. Discussion……………………………..…………………….………….....….56 4. 1. Reproductive isolation between M. milvus and M. migrans populations in Central Europe………………………………………………………………...….56 4. 2. Intraspecific variability in M. milvus and M. migrans populations in Central Europe and their migratory behaviour……………………………………..…...57 4. 3. Hybridization between M. milvus and M. migrans in Central Europe....59 5. References……………………………………………………………........….62 1 1. INTRODUCTION Evolutionists generally consider biological species more or less distant natural entities. In fact, there are very complex mechanisms which precede the final diversification of related lineages and study of them resulted to creation of several species concepts. If there is an absence of strong isolating mechanisms the boundary between species is blured by hybridization. To study the hybridization in sympatric bird species, I investigated the genetic structure of populations in two European kites (Milvus spp.). 1.1. Speciation and recognition of species and subspecies The definition of species as the main unit in classification of life forms has been a problem for many biologists. To recognize the ”clean lineages“ several species concepts (Tab. 1) based on different biological attributes were created (MAYDEN 1997, DE QUEIROZ 2007). The most popular one called as Biological species concept by MAYR (1942) describes species as ”groups of actually or potentially interbreeding natural populations, which are reproductively isolated from other such groups“. However, it has been widely confused and mixed with the general metapopulation lineage species concept of the same author. This fact was mentioned by DE QUEIROZ (2005) and replenished with a proposal of ”unified species concept“. It claimed, that ”metapopulation lineages do not have to be phenetically distinguishable, or diagnosable, or monophyletic, or reproductively isolated, or ecologically divergent, to be species. They only have to be evolving separately from other such lineages.“ The reproductive isolation is usually considered to be the main feature of good species (MAYR 1942). It consists of mechanisms which prevent two different species from gene flow between their populations (BARTON & BENGTSSOB 1986). If the barrier between two taxa is strong enough, any offspring produced by hybridization will not be fertile (CRAPON DE CAPRONA & FRITZSCH 1984). MAYR (1963) has classified the mechanisms of reproductive isolation to pre-zygotic (before fertilisation) and post-zygotic (after fertilisation). However, it is possible to observe examples where these 2 mechanisms are not strong enough, suggesting that there is the process of speciation going on. In such cases the two geographicaly not very divergent species may hybridize occasionally with possibility to produce fertile offspring and they can be described as ”semispecies“ (MALLET 2001). Tab. 1 Alternative species concepts and criteria on which they are based. Based on DE QUEIROZ (2007). Species concept Property(ies) References Biological Interbreeding (natural reproduction resulting in WRIGHT (1940) viable and fertile offspring) MAYR (1942) Isolation: Intrinsic reproductive isolation DOBZHANSKY (1950) (absence of interbreeding betw een heterospecific organisms based on intrinsic
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