Human-induced alterations of the environment are causing biotic changes worldwide, including the extinction of species and a mixing of once disparate floras and faunas. One type of biological communities that is expected to be particularly affected by environmental alterations are herb layer plant communities of fragmented forests such as those in the west European lowlands. However, our knowledge about current changes in species diversity and composition in these communities is limited due to a lack of adequate long-term studies. In this thesis, I resurveyed the herb layer communities of ancient forest patches in the Weser-Elbe region (NW Germany) after two decades using 175 semi-permanent plots. The general objectives were (i) to quantify changes in plant species diversity considering also between-community (β) and functional diversity, (ii) to determine shifts in species composition in terms of species’ niche breadth and functional traits and (iii) to find indications on the most likely environmental drivers for the observed changes. These objectives were pursued with four independent research papers (Chapters 1-4) whose results were brought together in a General Discussion. Alpha diversity (species richness) increased by almost four species on average, whereas β diversity tended to decrease (Chapter 1). The latter is interpreted as a beginning floristic homogenization. The observed changes were primarily the result of a spread of native habitat generalists that are able to tolerate broad pH and moisture ranges. The changes in α and β diversity were only significant when species abundances were neglected (Chapters 1 and 2), demonstrating that the diversity changes resulted mainly from gains and losses of low-abundance species. This study is one of the first studies in temperate Europe that demonstrates floristic homogenization of forest plant communities at a larger than local scale. The diversity changes found at the taxonomic level did not result in similar changes at the functional level (Chapter 2). The likely reason is that these communities are functionally “buffered”. Single communities involve most of the functional diversity of the regional pool, i.e., they are already functionally rich, while they are functionally redundant among each other, i.e., they are already homogeneous. Independent of taxonomic homogenization, the abundance of 30 species decreased significantly (Chapter 4). These species included 12 ancient forest species (i.e., species closely tied to forest patches with a habitat continuity > 200 years) and seven species listed on the Red List of endangered plant species in NW Germany. If these decreases continue over the next decades, local extinctions may result. This biotic impoverishment would seriously conflict with regional conservation goals. Community assembly mechanisms changed at the local level particularly at sites that experienced disturbance by forest management activities between the sampling periods (Chapter 3). Disturbance altered community assembly mechanisms in two ways: (i) it relaxed environmental filters and allowed the coexistence of different reproduction strategies, as reflected by a higher diversity of reproductive traits at the time of the resurvey, and (ii) it enhanced light availability and tightened competitive filters. These limited the functional diversity with respect to canopy height and selected for taller species. Thirty-one winner and 30 loser species, which had significantly increased or decreased in abundance, respectively, were characterized by various functional traits and ecological performances to find indications on the most likely environmental drivers for the observed floristic changes (Chapter 4). Winner species had higher seed longevity, flowered later in the season and had more often an oceanic distribution compared to loser species. Loser species tended to have a higher specific leaf area, to be more susceptible to deer browsing and to have a performance optimum at higher soil pH values compared to winner species. Multiple logistic regression analyses indicated that disturbances due to forest management interventions were the primary cause of the species shifts. As one of the first European resurvey studies, this study provides indications that an enhanced browsing pressure due to increased deer densities and increasingly warmer winters are important drivers. The study failed to demonstrate that eutrophication and acidification due to atmospheric deposition substantially drive herb layer changes. The restriction of the sample to the most base-rich sites in the region is discussed as a likely reason. Furthermore, the decline of several ancient forest species is discussed as an indication that the forest patches are still paying off their “extinction debt”, i.e., exhibit a delayed response to forest fragmentation. / Umweltveränderungen beeinträchtigen weltweit die Artenvielfalt. Die Lebensgemeinschaften fragmentierter Lebensräume gelten als besonders anfällig für Veränderungen. In dieser Arbeit wurden Untersuchungen an Krautschichtgemeinschaften historisch alter Waldfragmente im Elbe-Weser-Dreieck nach zwei Jahrzehnten wiederholt. Ziel war es anhand von 175 semi-permanenten Aufnahmeflächen (i) die Veränderungen der Pflanzenartendiversität zu quantifizieren, (ii) Artenverschiebungen in Bezug auf Nischenbreite und funktionale Merkmale festzustellen und (iii) Hinweise auf die verantwortlichen Umweltveränderungen zu finden. Die α-Diversität (Artenzahl) stieg durchschnittlich um vier Arten an. Die β-Diversität (Artenturnover zwischen den Flächen) nahm tendenziell ab. Letzteres wird als Beginn einer floristischen Homogenisierung interpretiert. Diese Studie ist eine der ersten im gemäßigten Europa, die eine floristische Homogenisierung von Waldpflanzengemeinschaften auf einer größeren als der lokalen Ebene aufzeigt. Die Diversitätsveränderungen auf taxonomischer Ebene führten nicht zu ähnlichen Veränderungen auf funktionaler Ebene. Bereits einzelne Gemeinschaften wiesen den Großteil der funktionalen Vielfalt des regionalen Artenpools, also ein Maximum an funktionaler Diversität auf. Gleichzeitig waren sie untereinander funktional redundant, also bereits homogen. Die mit der beginnenden taxonomischen Homogenisierung verbundene floristische Verarmung wird als gering eingestuft, da die Homogenisierung primär das Ergebnis der Zuwanderung häufig vorkommender Standortgeneralisten war. Unabhängig von der Homogenisierung gingen 30 Arten signifikant in ihrer Abundanz zurück, darunter 12 an historisch alte Wälder gebundene Arten sowie sieben Rote-Liste-Arten. Ein weiterer Rückgang oder ein lokales Aussterben dieser Arten stünde im Widerspruch zu regionalen Naturschutzzielen. Nullmodelltests und der Vergleich funktionaler und taxonomischer Diversitätskomponenten lassen auf regionaler Ebene auf eine zeitliche Konstanz der grundlegenden Mechanismen der Artenvergesellschaftung schließen. Auf der lokalen Ebene veränderten sich die Vergesellschaftungsmechanismen erheblich, insbesondere auf forstwirtschaftlich gestörten Standorten. Einerseits ermöglichte dort eine Abschwächung der Umweltfilter die Koexistenz von Arten mit unterschiedlichen Reproduktionsstrategien. Andererseits führte die erhöhte Lichtverfügbarkeit zu einer Verstärkung der Konkurrenzfilter und einer Selektion hochwüchsiger Arten. Gewinner- und Verliererarten wurden anhand funktionaler Merkmale und ihres ökologischen Verhaltens charakterisiert, um Hinweise auf die verantwortlichen Umweltveränderungen zu finden. Gewinnerarten wiesen eine höhere Langlebigkeit der Samen auf, blühten später in der Vegetationsperiode und hatten öfter eine ozeanische Verbreitung. Verliererarten hatten eine höhere spezifische Blattfläche, einen höheren Attraktivitätswert als Wildäsung und ein ökologisches Optimum bei höheren pH-Werten. Logistische Regressionsanalysen zeigen, dass Störung durch forstwirtschaftliche Eingriffe hauptverantwortlich für die Artenverschiebungen war. Zusätzlich liefert diese Wiederholungsstudie als eine der ersten in Europa Hinweise darauf, dass ein erhöhter Äsungsdruck sowie zunehmend mildere Winter entscheidende Einflussfaktoren darstellen. Der Rückgang mehrerer an historisch alte Wälder gebundener Arten wird als Anzeichen für eine verspätete Reaktion auf die Waldfragmentierung diskutiert.

Abstract Boreal and arctic plant communities are responding to anthropogenic climate change that has been exceptionally rapid during the recent decades. General responses include increased productivity, range expansions and biodiversity changes, all of which affect ecosystem functions. Vegetation dynamics are however controlled by multiple drivers, and the outcomes under the changing climate are not yet fully clear. As high latitude areas often lack long-term monitoring of vegetation, alternative methods are required to observe and understand vegetation changes and dynamics. Recently, resurveying historical vegetation data has become a valuable method of studying vegetation changes over the past few decades. In this thesis, I studied multidecadal (23–60 years) vegetation changes in forest and treeless heath and tundra plant communities along a latitudinal gradient in northern Fennoscandia using both vegetation resurveys and long-term experimental data. In addition to examining climate-driven vegetation changes, I related changes in plant communities to key local drivers of each context including mesotopography, grazing, soil moisture and soil fertility. General trends among the resurveyed treeless heath sites were the pronounced increase of the dwarf shrub Empetrum nigrum ssp. hermaphroditum in snow-protected habitats and the decrease of lichens throughout. Southernmost heath communities showed strong responses to multidriver effects and had shifted towards new community states. The long-term experiment in the tundra confirmed that depending on driver combinations, tundra communities evolve towards divergent alternative states, highlighting the importance of local drivers in modifying tundra vegetation over time. Communities in fertile forest sites experienced greater temporal turnover compared to infertile forest sites, suggesting that the soil fertility level is a key predictor of vegetation changes under climate change. This particularly important finding previously relied mainly on experimental evidence. Despite these generalities, changes in diversity, plant groups and species varied under a rather uniform climatic warming trend and were often habitat- or region-specific. Thus, the results of my thesis highly motivate continued monitoring and resurveying of vegetation under rapid environmental change and also form baseline time-series data for future studies. / Tiivistelmä Poikkeuksellisen nopea ilmastonmuutos on johtanut viime vuosikymmenten aikana muutoksiin boreaalisissa ja arktisissa kasviyhteisöissä. Muutoksiin lukeutuvat tuottavuuden lisääntyminen, levinneisyysrajojen siirtyminen sekä muutokset biodiversiteetissä, mitkä kaikki muuttavat ekosysteemien toimintaa. Kasvillisuuden dynamiikkaa säätelevät kuitenkin useat paikallistason tekijät, minkä seurauksena ei ole täysin selvää, miten kasvillisuus on eri alueilla ja habitaateissa muuttunut. Koska kasvillisuuden jatkuva monitorointi on harvinaista pohjoisilla alueilla, vanhojen kasvillisuusaineistojen uudelleenkartoituksista on tullut tärkeä menetelmä muutosten havaitsemiseksi. Tutkin väitöskirjassani vuosikymmenten kuluessa tapahtuneita (23–60 vuotta) kasvillisuusmuutoksia Pohjois-Fennoskandian metsissä, puuttomilla kankailla ja tundralla uudelleenkartoitusten ja kokeellisen tutkimuksen avulla, ja kytkin ne ilmastonmuutokseen sekä tärkeimpiin paikallisiin tekijöihin. Yleisiä trendejä uudelleenkartoitetuilla puuttomilla kankailla olivat variksenmarjan (Empetrum nigrum ssp. hermaphroditum) voimakas lisääntyminen lumensuojaisissa habitaateissa sekä jäkälien väheneminen kaikkialla. Yhteisöjen kokonaismuutos oli voimakkainta eteläisillä puuttomilla kankailla, jossa se korreloi yhtä aikaa lisääntyneiden lämpötilojen ja laidunpaineen kanssa. Kokeellinen tutkimus tundralla osoitti, että kasviyhteisöt kehittyvät hyvin erilaisiksi paikallisten tekijöiden voimakkuussuhteista riippuen, jotka voivat joko hidastaa tai nopeuttaa ympäristömuutoksista johtuvia kasvillisuusmuutoksia. Metsien uudelleenkartoitus osoitti yhteisöjen kokonaismuutoksen olevan pitkällä aikavälillä suurempaa tuottavilla maaperillä lehtometsissä verrattuna karumpiin kangasmetsiin. Tutkimuksen mukaan maaperän tuottavuus on avaintekijä, joka ennustaa kasvillisuusmuutosten voimakkuutta ilmastonmuutoksen aikana. Tästä tärkeästä löydöstä oli aiemmin pääasiassa vain kokeellista tutkimustietoa. Yleisistä trendeistä huolimatta, muutokset diversiteetissä, kasviryhmissä ja yksittäisissä lajeissa olivat kuitenkin vaihtelevia ja usein habitaatti- tai aluesidonnaisia. Väitöskirjani tulokset, jotka muodostavat myös aikasarjan tuleville tutkimuksille, osoittavat kasvillisuuden monitoroinnin ja uudelleenkartoitusten olevan ensisijaisen tärkeitä, jotta kasvillisuuden dynamiikkaa voidaan ymmärtää paremmin nopeasti muuttuvissa olosuhteissa.

Vegetation changes in Czech lowland forests over the past decades Martin Kopecký ABSTRACT To explore decadal changes in plant species diversity and composition of typical lowland forests in the Czech Republic, I compared vegetation on plots sampled decades ago with vegetation on the same plots sampled recently. First I evaluated the robustness of the approach used in my thesis. In Chapter 1, we provided the first direct test of the effect of uncertainty in original plot location on results from vegetation resurvey. We found that temporal trends in vegetation diversity and composition were comparable between exactly relocated permanent and only approximately relocated semi-permanent plots. Therefore, we conclude that the resurvey of semi-permanent plots is robust to the uncertainty in original plot location. Then, we showed that vegetation in lowland oak forest shifted from species-rich communities of thermophilous forest toward species poorer communities of mesic forest (Chapter 2). The species typical for thermophilous oak forests and nationally endangered species suffered the most significant decline. We identified as the main driver behind these changes shift from traditional coppicing toward high forest management after WWII. Further, we explored the processes behind temporal shifts in species diversity...