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Biology, Ecology and Invasion Characteristics Of Charles University in Prague, Faculty of Science Department of Botany Biology, ecology and invasion characteristics of Campylopus introflexus in the Czech Republic Biologie, ekologie a charakteristiky invaze druhu Campylopus introflexus v Ceskéˇ Republice Mgr. Eva Mikulášková Study programme: Biology; Botany Ph.D. thesis Brno 2012 Supervisor: RNDr. ZdenˇekSoldán, CSc. DECLARATION I hereby declare, that I made this thesis independently, using the listed refer- ences, or in the cooperation with other authors of the papers (for my contri- bution to particular papers see chapter 6 Author´s contribution). I did submit neither the thesis nor its any part to acquire any other academic title. Brno, 2012 Mgr. Eva Mikulášková ABSTRACT Ecological and economic impact of invasive plants to natural ecosystems is the subject of many studies; however, invasive bryophytes have been stud- ied only marginally. Campylopus introflexus (Hedw.) Brid. is one of the most strongly invasive bryophyte species in Europe. The species appears to be native in the Southern Hemisphere. In Europe, it was collected for the first time in the British Isles in 1941. The moss has expanded eastward and the first collection in the Czech Republic is dated to 1988. This thesis found that more than 70 localities were known known in the Czech Republic in 2006, and more than 100 localities became known by 2011. It has been further demonstrated that the Czech Republic was colonized repeatedly by generative spores and all populations have a unique genetic composition. Genetic variation of the populations is low, the genetic diver- sity of populations within the Czech Republic is not correlated with their geographic position or with any of the monitored environmental variables. At a fine scale within particular localities, the species disperses by vege- tative diaspores, while it uses generative spores for spreading over longer distances. In Central Europe, C. introflexus prefers open coniferous forests, especially plantations of either spruce or pine. It colonizes clearings, pathsides, forests edges and disturbed peatbogs. It may form large and compact stands at these habitats. It is able to colonize microhabitats with very heterogeneous ecologi- cal conditions. It inhabits acidic, nutrient-poor, bare soil without strong com- petition with vascular plants, lichens and other bryophytes. Cultivation ex- periments as well as vegetation-data analyses showed that the species thrives best on organic and sandy soils and avoids limestone and strongly water- logged soils. C. introflexus represents no significant risk to natural plant communities in Central Europe presently, because it colonizes mainly human-disturbed habitats. It can form stable mixed stands with the domestic pioneer species. It occupies free patches more rapidly than some native species, but it not able to outcompete them directly. The ratio of coverage of C. introflexus and native pioneer species is partially dependent on the amount of both generative and vegetative diaspores during initial colonization. In some cases C. introflexus can block the natural succession. v ABSTRAKT(INCZECH) Ekologický a ekonomický dopad invazních rostlin na pˇrirozené ekosystémy je pˇredmˇetemmnoha studií, avšak invazní mechorosty jsou studovány pouze okrajovˇe. Campylopus introflexus (Hedw.) Brid. je nejvýznamnˇejšíinvazní druh mechorostu v Evropˇe.Pochází z jižní polokoule a v Evropˇebyl poprvé zaz- namenán v roce 1941 na Britských ostrovech, odkud se postupnˇešíˇrísmˇerem na východ. První údaj z Ceskéˇ republiky pochází z roku 1988. V rámci této práce bylo zjištˇeno,že v roce 2006 bylo v Ceskéˇ republice známo pˇres 70 lokalit, v roce 2011 už bylo známo více než 100 lokalit. Dále je ukazováno, že Ceskᡠrepublika byla kolonizována opakovanˇepomocí gener- ativních spor, všechny populace mají unikátní genotyp. Populace mají malou genetickou variabilitu, genetická diverzita v rámci Ceskéˇ republiky není ko- relována s geografickou pozicí ani s žádnou ze sledovaných promˇenných prostˇredí. V rámci jemného mˇeˇrítkajedné dílˇcílokality se rozšiˇrujepomocí vegetativních diaspor, zatímco pro šíˇrení na vˇetšívzdálenosti využívá gener- ativní spory. Ve stˇrední Evropˇe C. introflexus preferuje jehliˇcnatéprosvˇetlenélesy, zvláštˇe monokultury smrku ˇciborovice. Druh kolonizuje paseky, bˇrehy cest, okraje porost ˚ua narušená rašeliništˇe.Na tˇechtomístech m ˚užetvoˇrit rozsáhlé a kompaktní porosty. Je schopný osídlit mikrostanovištˇes velice r ˚uznorodými ekologickými podmínkami. Osidluje holou, kyselou, na živiny chudou p ˚udu bez veliké kompetice cévnatých rostlin, lišejník ˚ua ostatních mechorost ˚u. Kultivaˇcníexperimenty stejnˇejako analýzy vegetaˇcníchdat ukázaly, že druh nejlépe prosperuje na organických a písˇcitých p ˚udácha vyhýbá se p ˚udám vápenatým a silnˇepodmáˇceným. C. introflexus v souˇcasnédobˇenepˇredstavuje významné riziko pro pˇrirozená spoleˇcenstva stˇrední Evropy, protože kolonizuje pˇrevážnˇeantropogennˇenarušená stanovištˇe.S domácími pionýrskými druhy m ˚užetvoˇritstabilní smˇesnéporosty. Volný neosídlený prostor osidluje mnohem rychleji než ostatní domácí druhy, nicménˇeje není schopen pˇrímovytlaˇcovat. Pomˇerpokryvnosti druhu C. in- troflexus a domácích pionýrských druh ˚uje ˇcásteˇcnˇe závislý na množství pohlavních i nepohlavních diaspor pˇripoˇcáteˇcníkolonizaci. V nˇekterých pˇrí- padech m ˚uže C. introflexus blokovat pˇrirozenou sukcesi na stanovištích. vi Nature preserves the species, and cares but very little for individuals. — Voltaire ACKNOWLEDGMENTS First of all, I would like to thank Michal Hájek, Jon and Blanka Shaw for motivation, support, and many indispensable advices. I thank to my Thesis adviser, ZdenˇekSoldán for help starting my work and for help in the field. I am grateful to Michal Hájek, Zuzana Fajmonová and Tomáš Fér for their comments on the manuscript of this thesis. My thanks are due to Matthew Johnson and Jon Shaw for the revision of English style. For some consultations and valuable information for on work in the DNA lab, I thank to Veronika Kuˇcabová, and I also thank to JiˇríVáˇnafor consulta- tions about bryology. I thank Ondˇrej Hájek for the creation of maps. For the precise assistance and kind atmosphere, I thank the staff of cryp- togamic section and the staff of the DNA lab of Department of Botany, Fac- ulty of Science, Charles University and staff of the Mire ecology group of Department of Botany and Zoology, Faculty of Science, Masaryk University. I would like to thank my colleagues and friends from the Czech and Slo- vak lichenological community for nice experiences and kind atmosphere throughout the years. I deeply thank JiˇríMikulášek, Jindˇriškaand my family for their patience and all support. I thank to all babysitters who took care about Jindˇriškaduring writing this thesis. My work was supported by Grant Agency of Charles University (project no. 258/2004/B-BIO/PrF) and institutional support of Masaryk University. vii CONTENTS i introduction1 1 general introduction3 1.1 Bryophytes . 5 1.2 Heath star moss Campylopus introflexus (Hedw.) Brid. 6 1.2.1 Taxonomy . 6 1.2.2 Anatomy, morphology and reproduction . 7 1.2.3 Ecology . 9 1.2.4 Phytosociology . 11 1.2.5 Biogeography – native and invasive distribution . 11 ii main thesis 15 2 the object and the aims of the study 17 3 outline of the thesis 19 3.1 The main ideas and results of papers . 19 3.1.1 Habitats colonized by C. introflexus in the Central Eu- rope (Paper 1)......................... 19 3.1.2 C. introflexus dispersal and genetic diversity (Paper 2). 20 3.1.3 Molecular methods optimalization (Paper 3)....... 21 3.1.4 The role of interspecific competition in C. introflexus spreading (Paper 4)...................... 21 3.1.5 Distribution of C. introflexus in the Czech Republic (Pa- per 5).............................. 22 3.2 The results of papers in the broader context of species biology . 22 3.2.1 Dispersal mode . 22 3.2.2 Invaded habitats . 23 3.2.3 Distribution . 25 4 conclusion 27 iii papers 37 5 list of papers 39 6 author´s contribution 41 7 habitats colonized by c. introflexus in the central europe 43 7.1 Introduction . 44 7.2 Methods . 46 7.2.1 Data sampling, vegetation data . 46 7.2.2 Data analyses . 48 7.3 Results . 50 7.3.1 Distribution of Campylopus introflexus in the Czech Re- public . 50 7.3.2 Vegetation composition of C. introflexus habitats . 51 7.3.3 Ecological characteristic of the habitats – field data . 53 7.3.4 Ecological characteristic of the habitats – cultivation ex- periment . 57 7.3.5 Invasibility of vegetation by C. introflexus ........ 58 ix x contents 7.4 Discussion . 59 7.4.1 Distribution pattern in the Czech Republic . 59 7.4.2 Recently invaded habitats and their ecological charac- teristics . 61 7.4.3 Differences in habitat affiliation between the Czech Re- public and Western Europe . 61 7.4.4 Future invasion potential . 63 7.4.5 Conclusions . 63 8 c. introflexus dispersal and genetic diversity (paper 2) 71 8.1 Introduction . 72 8.2 Methods . 74 8.2.1 Samples . 74 8.2.2 DNA extraction and AFLP fingerprinting . 77 8.2.3 Isozyme analysis . 77 8.2.4 Data analysis . 79 8.3 Results . 80 8.3.1 Clonal structure . 82 8.3.2 Genetic diversity . 83 8.3.3 Genetic structure . 83 8.3.4 Genetic relationships at the population scale . 85 8.4 Discussion . 85 8.4.1 Distribution mechanisms of C. introflexus ......... 89 8.4.2 Genetic variation among and within populations . 90 8.4.3 Spatial genetic structure in the Czech Republic . 91 8.4.4 Conclusions . 92 9 molecular methods optimalization (paper 3) 101 9.1 Introduction . 101 9.2 Methods . 103 9.2.1 Sample collection . 103 9.2.2 DNA extraction . 103 9.2.3 AFLP fingerprinting . 104 9.2.4 Assessment of AFLP error rate . 106 9.3 Results . 106 9.3.1 DNA extraction . 106 9.3.2 AFLP fingerprinting . 107 9.4 Discussion . 108 9.4.1 Optimization of DNA extraction . 108 9.4.2 AFLP fingerprinting . 113 9.4.3 Reproducibility tests . 114 10 the role of interspecific competition in c. introflexus spreading 119 10.1 Introduction . 120 10.2 Methods . 122 10.2.1 Field design . 122 10.2.2 Data analysis . 123 10.3 Results . 123 10.3.1 Colonization of unvegetated patches .
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