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Disertační Práce PŘÍRODOVĚDECKÁ FAKULTA Disertační práce Veronika Kalníková Brno 2020 PŘÍRODOVĚDECKÁ FAKULTA Diversity and ecology of the river gravel-bar vegetation Ph.D. Dissertation Veronika Kalníková Vedoucí práce: prof. RNDr. Milan Chytrý, Ph.D. Ústav botaniky a zoologie Brno 2020 Bibliografický záznam Autor: Mgr. Veronika Kalníková Přírodovědecká fakulta, Masarykova univerzita Ústav botaniky a zoologie Název práce: Diverzita a ekologie vegetace štěrkových říčních náplavů Studijní program: Ekologická a evoluční biologie Studijní obor: Botanika Vedoucí práce: prof. RNDr. Milan Chytrý, Ph.D. Akademický rok: 2019/2020 Počet stran: 34+150 Klíčová slova: databáze; disturbance; divočící toky; Evropa; fytocenologické snímky; klasifikace; mechorosty; ochrana biotopů; rostlinná společenstva; sukcese vegetace; štěrkové náplavy; vegetace Bibliographic Entry Author: Mgr. Veronika Kalníková Faculty of Science, Masaryk University Department of Botany and Zoology Title of Thesis: Diversity and ecology of the river gravel-bar vegetation Degree programme: Ecological and Evolutionary Biology Field of Study: Botany Supervisor: prof. RNDr. Milan Chytrý, Ph.D. Academic Year: 2019/2020 Number of Pages: 34+150 Keywords: databases; disturbance; braided rivers; Europe; phytosociological relevés; classification; bryophytes; habitat protection; plant communities; vegetation succession; gravel bars; vegetation Abstrakt Štěrkové náplavy jsou dynamickými a heterogenními biotopy vázanými na horské vodní toky. Existence štěrkových náplavů je závislá na pravidelných záplavách, které jejich vegetaci udržují v různých sukcesních stadiích, na něž jsou vázané specializované a často vzácné druhy rostlin. Štěrkové náplavy se vyskytují na tocích s přirozeným hydromorfologickým režimem a v současné Evropě patří vlivem antropogenních tlaků mezi ohrožené biotopy. Klasifikace vegetace štěrkových náplavů je v Evropě nekonzistentní a v některých zemích není tato vegetace ani rozlišována, což může komplikovat její monitoring, ochranu i management. Cílem práce proto bylo doplnit dosavadní znalosti o rozšíření a ekologii vegetace štěrkových náplavů v Evropě a navrhnout první jednotnou klasifikaci. Na rozlišení sukcesních stadiích vegetace náplavů jsou založeny některé klasifikační přístupy. Sukcesí vegetace štěrkových náplavů, ekologickými aspekty, diverzitou, druhovou skladbou a vývojem jednotlivých stadií jsme se zabývali na čtyřech malých tocích v Západních Karpatech (Česká republika), které zasáhla padesátiletá povodeň. Studie potvrdila jako nejdůležitější faktory ovlivňující směr a rychlost sukcese vegetace nadmořskou výšku a strukturu substrátu. Pro účel celoevropské klasifikace jsme shromáždili a digitalizovali vegetační snímky, které doposud nebyly součástí žádné dostupné vegetační databáze, a založili databázi vegetace štěrkových říčních náplavů (Gravel Bar Vegetation Database). Její součástí se staly vegetační snímky z vlastních terénních výzkumů, které probíhaly v zemích, kde byla tato vegetace méně prozkoumaná nebo údaje chyběly (především z jihovýchodní a severní Evropy a Kavkazu). Pro Gruzii jsme na základě fyziognomie porostů a metody neřízené klasifikace nově vylišili pět vegetačních typů štěrkových náplavů: raně sukcesní vysokohorskou bylinnou vegetaci (asociace Epilobietum colchici), raně sukcesní podhorskou bylinnou vegetaci (společenstvo Petrorhagia saxifraga-Crepis foetida), porosty s Calamagrostis pseudophragmites (Tussilagini farfarae-Calamagrostietum pseudophragmitae), křoviny s Myricaria germanica (Salici purpureae-Myricarietum germanicae) a Hippophaë rhamnoides (Salici incanae-Hippophaëtum rhamnoidis). Vegetaci s Calamagrostis pseudophragmites a křoviny jsme přiřadili k analogickým asociacím popisovaným také ze střední Evropy. U vegetačních jednotek jsme definovali diagnostické druhy a popsali jejich ekologické nároky. Na celoevropské úrovni jsme navrhli jednotnou klasifikaci vegetace štěrkových říčních náplavů. Raně sukcesní a křovinná vegetace byly klasifikovány v rámci dvou tříd, Thlaspietea rotundifolii a Salicetea purpureae. Celkem jsme vytvořili formální definice pro dvě subasociace, jedenáct asociací a čtyři svazy (Calamagrostion pseudophragmitae, Epilobion fleischeri, Salicion cantabricae a Salicion eleagni). Definice jsou založeny na výskytu a pokryvnostech skupin druhů s podobnou ekologií, nebo jednotlivých druhů úzce specializovaných na určitý typ vegetace náplavů. Navržená klasifikace byla ověřena pomocí metod ordinace a neřízené klasifikace. U vegetačních jednotek jsme revidovali nomenklaturu, definovali diagnostické druhy a popsali jejich ekologické nároky a rozšíření. Ukázali jsme, že hlavní variabilita v datech je svázána s nadmořskou výškou, biogeografií, místními hydromorfologickými procesy a sukcesí vegetace. Použitím vegetačních snímků z Balkánu a Kavkazu jsme rozšířili znalosti o druzích mechorostů, které se vyskytují na štěrkových náplavech, a nastínili jejich ekologii a vegetační vazby. Drobný ruderální mech Bryum klinggraeffii byl zjištěn jako nový druh pro Albánii, Černou Horu, Gruzii a Srbsko. Abstract The river gravel bars of mountain streams are spatiotemporally dynamic habitats. Their presence in a landscape depends on regular and relatively frequent flooding disturbances creating a variety of successional stages encompassing rare specialised species. Gravel-bar habitats became rare in Europe due to river regulations and other artificial alterations leading to the interruption of the natural hydro-morphological regime of gravel-bed rivers. The classification of gravel-bar vegetation in Europe, a keystone for the conservation planning, monitoring and management, is inconsistent or even ignored in some national schemes. Therefore, we aimed to extend the knowledge on the distribution and ecology of the vegetation types of European mountain gravel-bar habitat and propose the first unified vegetation classification. Several classification systems emphasized the main criteria for the delimitation of vegetation types of river gravel-bar habitat to be the physiognomy related to the vegetation succession. We performed a case study focused on the ecological aspects of the vegetation succession and changes in plant diversity on river gravel bars of four streams in the Western Carpathians (Czech Republic) which experienced extreme 50- year flood event. In this study, we described the individual successional stages in terms of the species turnover, richness and other characteristics. We identified altitude and size of gravel/stone particles to be the most important factors influencing the vegetation succession. For purpose of development of the pan-European vegetation classification of river gravel-bar habitats, we collected and digitized vegetation-plot data that had not been previously stored in electronic databases and included them in newly created Gravel Bar Vegetation Database. An important part of the database are vegetation plots from our field sampling in the countries where this vegetation had not been studied before or was less explored, especially in south-eastern and northern Europe and the Caucasus. We studied vegetation of gravel bars of several gravel-bed rivers across Georgia. We distinguished five vegetation types based on vegetation physiognomy and unsupervised classification and described them as phytosociological vegetation units. Two of them were described as new to science, i.e. early-successional herbaceous vegetation at higher altitudes (Epilobietum colchici) and that at lower altitudes (Petrorhagia saxifraga-Crepis foetida community), and other were assigned to the associations previously described from Central Europe, i.e. the grasslands dominated by Calamagrostis pseudophragmites (Tussilagini farfarae-Calamagrostietum pseudophragmitae) and scrub vegetation (Salici purpureae-Myricarietum germanicae and Salici incanae-Hippophaëtum rhamnoidis). We established diagnostic plant species for each type and related them to environmental variables. The main aim of the thesis is to determine the vegetation types of gravel-bar habitat in Europe and to assign them to the association and subassociation levels using formalized classification approach. We classified early-successional and scrub gravel- bar vegetation types into two classes, i.e. Thlaspietea rotundifolii and Salicetea purpureae, respectively, and we formally defined two subassocitaions, eleven associations and four alliances (Calamagrostion pseudophragmitae, Epilobion fleischeri, Salicion cantabricae and Salicion eleagni). This distinction was supported by unsupervised classification and ordination modelling. Based on a critical revision, we merged or discarded some associations or alliances which were already defined and described in literature. We established diagnostic plant species for each type and related environmental variables to those types, as well. The main variability within the distinguished vegetation types is connected to the altitudinal gradient, biogeographical variation, local hydro-morphological processes and successional development. As there is a general lack of knowledge on the distribution of bryophyte species growing on gravel bars and on their vegetation affinity, we used the vegetation-plot data collected in several Balkan countries and the Caucasus and described their occurrence in a geographical and ecological context. During the fieldwork, Bryum klinggraeffii, small ruderal moss of frequently disturbed open habitats, was found as a new species for Albania, Georgia, Montenegro and Serbia. Poděkování/Acknowledgements Děkuji, Milanovi,
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