Summary 1.Restoration activities aiming at increasing vegetation diversity often try to stimulate both dispersal and germination. In wetlands, dispersal and germination are coupled as water and water level fluctuations (WLF) simultaneously influence seed transport and germination conditions (soil moisture). Water regime shifts have been shown to affect vegetation composition. However, the interactions between WLF, dispersal and subsequent germination as drivers of such changes are still poorly understood, especially within the complexity of a field situation. 2. We tested the effect of soil moisture on ten riparian species in the greenhouse and sowed these species on 135 field locations in nine wetlands with recently restored WLF. We used quantile regressions to test the effects of WLF on the window of opportunity for germination from sown seeds and other seeds naturally dispersed to our plots, as well as on community diversity. 3. Soil moisture significantly affected germination both in the greenhouse and in the field. In the complexity of a field situation, a flooding depth just below the soil level, an intermediate flooding duration and a high flooding frequency provided the best opportunities for maximal germination. This was because these conditions enhanced germination from the seed bank as well as increasing germination from dispersed seeds. Seedling diversity showed identical patterns. 4. Other known (i.e., light conditions) and unknown factors played a role as we found low and variable germination, even under optimal conditions. We found evidence that WLF can affect vegetation zonation as flooded seedling communities contained more species with high moisture affinity. 5. Synthesis and applications. Water level fluctuations provide clear windows of opportunity for germination both from the seed bank and from dispersed seeds. Water regime changes are therefore likely to strongly affect recruitment opportunities and subsequent community assembly in riparian ecosystems, for instance through climate change or management. Water level fluctuations can be used as management tool to stimulate plant recruitment and seedling diversity in riparian wetlands.
Summary1. Seed dispersal and germination are two primary processes influencing plant community assembly. On freshwater shores, water levels regulate both processes. However, it is still unclear how water levels, shore morphology and species traits interactively affect seed dispersal and germination, and how these interactions determine plant community assembly. We hypothesize that a drawdown water regime enhances seed establishment compared to a year-round stable water level, that this increases species richness and diversity, and that this is modulated by species traits and shore morphology. 2. Germination of 20 wetland plant species with different dispersal capacities (floating capacity expressed as seed floatation half-time) and soil moisture preferences for germination (Ellenberg F) was tested on artificial shores in 24 outdoor ponds in two complementary experiments over 8 weeks.The 'dispersal experiment' tested the effect of water regime on recruitment of hydrochorously dispersing seeds. The 'seed bank experiment' tested the effect of water regime on germination from a sown seed bank, on steep and gradual shores. 3. In the dispersal experiment, the drawdown regime increased recruitment and species richness. Longer floating species colonized a larger shoreline section. Soil moisture preference for germination did not determine colonization patterns. 4. In the seed bank experiment, the drawdown regime increased the number of seedlings on gradual sloping shores, but not on steep shores. The number of germinating seedlings corresponded to the area subjected to the drawdown regime in both shore types. Species richness was not affected by water regime or shore morphology, and species traits did not determine shoreline colonization. Most seeds germinated in moist soil conditions for all species. 5. Synthesis. A spring drawdown instead of stable water regime stimulates establishment of hydrochorously dispersing seeds in temperate wetlands, leading to higher species richness and diversity. Germination from the seed bank is more affected by water regime and shore surface than by the tested species traits. Species traits, water levels and shore morphology together determine wetland plant community assembly, with dispersal as the main driver of seedling community diversity. Water-level regulations and shore morphology can be used to influence plant communities in wetland restoration.
Some shallow lakes switch repeatedly back and forth between a vegetation dominated clear-water state and a contrasting turbid state. Usually such alternations occur quite irregularly, but in some cases the switches between states are remarkably regular. Here we use data from a well-studied Dutch lake and a set of simple models to explore possible explanations for such cyclic behavior. We first demonstrate from a graphical model that cycles may in theory occur if submerged macrophytes promote water clarity in the short run, but simultaneously cause an increased nutrient retention, implying an accumulation of nutrients in the long run. Thus, although submerged plants create a positive feedback on their own growth by clearing the water, they may in the long run undermine their position by creating a slow ''internal eutrophication''. We explore the potential role of two different mechanisms that may play a role in this internal eutrophication process using simulation models: (1) reduction of the P concentration in the water column by macrophytes, leading to less outflow of P, and hence to a higher phosphorus accumulation in the lake sediments and (2) a buildup of organic matter over time resulting in an increased sediment oxygen demand causing anaerobic conditions that boost P release from the sediment. Although the models showed that both mechanisms can produce cyclic behavior, the period of the cycles caused by the build-up of organic material seemed more realistic compared to data of the Dutch Lake Botshol in which regular cycles with a period of approximately 7 years have been observed over the past 17 years.
Till about 1965 a rich characean community occurred in the shallow peat lake Botshol with six species of which the rare Nitellopsis obtusa and Chara major dominated at many sites. In the period 1980-1988 characean biomass strongly decreased and only two species, Chara globularis and C. connivens, remained in small populations at a few localities. Of the macrophyte Najas marina also some small populations remained, while the aquatic moss Fontinalis antipyretica and the filamentous alga Vaucheria dichotoma predominated at many sites. These phenomena may have been due to eutrophication by the input of polluted water. This process of impoverishment was stopped by restoration measures in 1989, resulting in a lower phosphorus concentration (ca 25 ig 1-') and a higher water transparency. Immediately after these measures the Characeae community increased in abundance and number of species. During the summer of 1990, and especially 1991, a spectacular growth occurred of Chara connivens. C. connivens was often accompanied by C. major. Other species with scattered occurrence were C. aculeolata, C. aspera, C. contraria and C. globularis. The reasons for the shift in dominance from Nitellopsis obtusa to Chara connivens are discussed. From growth experiments evidence was obtained that neither the recent higher chloride level, nor the lowered phosphate concentration were the main factors for the domination of Chara connivens.
27The transitions between ecosystems (ecotones) are often biodiversity hotspots, but we know little 28 about the forces that shape them. Today, often sharp boundaries with low diversity are found 29 between terrestrial and aquatic ecosystems. This has been attributed to environmental factors 30 that hamper succession. However, ecosystem properties are often controlled by both bottom-up 31 and top-down forces, but their relative importance in shaping riparian boundaries is not known. 32We hypothesize that 1) herbivores may enforce sharp transitions between terrestrial and 33 aquatic ecosystems by inhibiting emergent vegetation expansion and reducing the width of the 34 transition zone and 2) the vegetation expansion, diversity and species turnover are related to 35 abiotic factors in the absence of herbivores, but not in their presence. We tested these 36 hypotheses in 50 paired grazed and ungrazed plots spread over ten wetlands, during two years. 37Excluding grazers increased vegetation expansion, cover, biomass and species richness. In 38 ungrazed plots, vegetation cover was negatively related to water depth whereas plant species 44We conclude that (aquatic) herbivores can strongly inhibit expansion of the riparian 45 vegetation and reduce vegetation diversity over a range of environmental conditions. 46Consequently, herbivores enforce sharp boundaries between terrestrial and aquatic ecosystems.
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