Invasion by mats of free-floating plants is among the most important threats to the functioning and biodiversity of freshwater ecosystems ranging from temperate ponds and ditches to tropical lakes. Dark, anoxic conditions under thick floating-plant cover leave little opportunity for animal or plant life, and they can have large negative impacts on fisheries and navigation in tropical lakes. Here, we demonstrate that floating-plant dominance can be a self-stabilizing ecosystem state, which may explain its notorious persistence in many situations. Our results, based on experiments, field data, and models, represent evidence for alternative domains of attraction in ecosystems. An implication of our findings is that nutrient enrichment reduces the resilience of freshwater systems against a shift to floating-plant dominance. On the other hand, our results also suggest that a single drastic harvest of floating plants can induce a permanent shift to an alternative state dominated by rooted, submerged growth forms.D ense mats of free-floating plant have an adverse effect on freshwater ecosystems because they create anoxic conditions that strongly reduce animal biomass and diversity (1). Invasions by introduced exotic species are partly responsible for the increase of floating plant dominance. The problems caused by Eichhornia crassipes, Pistia stratiotes, and Salvinia molesta are notorious: they hamper fish production and navigation in tropical regions around the world (2-4). However, eutrophication is likely to have boosted the spread of free-floating plants, too. In temperate climate zones, it is known that dense beds of duckweeds (Lemnaceae) and small, floating water ferns (Azollaceae) are a symptom of high-nutrient loading in small water bodies, such as ponds and canals (5, 6). Just as in the case of tropical plant beds, the dark and anoxic conditions under thick duckweed cover leave little opportunity for animal or plant life (1).The dependence of free-floating plants on high nutrient concentrations in the water is an obvious consequence of their growth form. They have no direct access to the sediment pool of nutrients, and they have a large portion of their leaf surface exposed to the atmosphere rather than to the water, thereby reducing the possibility of taking up nutrients other than carbon through their leaves. By contrast, rooted submerged macrophytes may take up a large part of their nutrients from the sediment (7, 8) and also use their shoots effectively for nutrient uptake from the water column (9, 10). Although floating plants are obviously superior competitors for light, submerged plants may affect the growth of free-floating plants through a reduction of available nutrients in the water column. Competition is likely to be especially strong for nitrogen. Although phosphorus availability in the water column can be reduced because of uptake by submerged macrophytes (11), many studies show unaltered or even increased ortho-P levels after increased macrophyte cover (12-15). By contrast, submerged nitrogen co...
ABSTRACT1. The European Water Framework Directive requires the determination of ecological status in European fresh and saline waters. This is to be through the establishment of a typology of surface water bodies, the determination of reference (high status) conditions in each element (ecotype) of the typology and of lower grades of status (good, moderate, poor and bad) for each ecotype. It then requires classification of the status of the water bodies and their restoration to at least 'good status' in a specified period.2. Though there are many methods for assessing water quality, none has the scope of that defined in the Directive. The provisions of the Directive require a wide range of variables to be measured and give only general guidance as to how systems of classification should be established. This raises issues of comparability across States and of the costs of making the determinations.3. Using expert workshops and subsequent field testing, a practicable pan-European typology and classification system has been developed for shallow lakes, which can easily be extended to all lakes. It is parsimonious in its choice of determinands, but based on current limnological understanding and therefore as cost-effective as possible.4. A core typology is described, which can be expanded easily in particular States to meet local conditions. The core includes 48 ecotypes across the entire European climate gradient and incorporates climate, lake area, geology of the catchment and conductivity.5. The classification system is founded on a liberal interpretation of Annexes in the Directive and uses variables that are inexpensive to measure and ecologically relevant. The need for taxonomic expertise is minimized.6. The scheme has been through eight iterations, two of which were tested in the field on tranches of 66 lakes. The final version, Version 8, is offered for operational testing and further refinement by statutory authorities.
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