Contemporary ecological landscape planning is often based on the assumption that small isolated habitat patches sustain relatively few species. Here, we suggest that for shallow lakes and ponds, the opposite can be true for some groups of organisms. Fish communities tend to be poor or even absent in small isolated lakes. However, submerged vegetation is often more abundant in such waterbodies. As a consequence of low fish biomass and high vegetation abundance, the richness of aquatic birds, plants, amphibians and invertebrates is often relatively high in small, shallow, isolated lakes. Although the rarity of fish is in line with expectations from the ruling paradigms about effects of habitat fragmentation in landscape ecology, the relative richness of various other groups of organisms in small ponds is opposite to these expectations. The case of shallow lakes illustrates that incorporating ecological interactions is essential to understanding the potential effects of habitat fragmentation. Single‐species meta‐population approaches may be misleading if ecological interactions are strong. A meta‐community approach that explicitly incorporates biotic interactions, also those involving different trophic levels, is needed. Our diagnosis suggests that connection of isolated habitat fragments may in some cases reduce, rather than enhance, landscape‐level biodiversity, and implies that biodiversity at the regional level will be maximized if the local habitat patches vary widely in size and degree of connectivity.

1. Most temperate rivers are heavily regulated and characterised by incised channels, aggradated floodplains and modified hydroperiods. As a consequence, former extensive aquatic /terrestrial transition zones lack most of their basic ecological functions. 2. Along large rivers in Europe and North America, various floodplain restoration or rehabilitation projects have been planned or realised in recent years. However, restoration ecology is still in its infancy and the literature pertinent to river restoration is rather fragmented. (Semi-) aquatic components of floodplains, including secondary channels, disconnected and temporary waters as well as marshes, have received little attention, despite their significant contribution to biological diversity. 3. Many rehabilitation projects were planned or realised without prior knowledge of their potential for success or failure, although, these projects greatly contributed to our present understanding of river-floodplain systems. 4. River rehabilitation benefits from a consideration of river ecosystem concepts in quantitative terms, comparison with reference conditions, historical or others, and the establishment of interdisciplinary partnerships. 5. We present examples from two large European rivers, the Danube and the Rhine, in which the role of aquatic connectivity has been extensively studied. The Danube delta with its diversity of floodplain lakes across an immense transversal gradient (up to 10 km) serves as a reference system for restoration projects along lowland sections of large rivers such as the Rhine in the Netherlands

Summary 1.In recent years, interest has grown in restoring floodplain function of regulated rivers. Successful rehabilitation of riparian systems requires knowledge of how regulation of river flow affects biodiversity and ecosystem function. The effects of changes in the river's low water-level regime on aquatic ecosystems in floodplains has received little attention so far. 2. The aquatic vegetation of 215 floodplain lakes along the Lower Rhine (the Netherlands) was analysed in relation to lake characteristics and lake water-level fluctuations in 1999-2000. 3. Vegetation composition was related to lake morphology and age, cattle access to the shoreline, the amount of time the river was in flood, and lake sediment area exposed at low water level (drawdown). Surprisingly, vegetation composition was correlated more strongly with lake age and occurrence of drawdown than the amount of time the river was in flood. 4. In older lakes, water-level fluctuations are reduced due to an accumulation of clay and silt that 'seals' sediment, preventing drawdown during periods of low river levels. Our results suggest that this clay sealing process is a major driving force for aquatic vegetation succession in floodplain lakes along the Lower Rhine, as succession drives from desiccation-tolerant species (e.g. Chara spp.) in young lakes to desiccation-sensitive species (e.g. Nuphar lutea ) in old lakes. 5. Water levels were stable in lakes along a river branch that was impounded below mean flow only. Here, the original low water-level regime has been replaced by an artificial regime with higher water levels than would be expected naturally. Consequently, in these lakes drawdown was rare or absent, and the aquatic macrophyte vegetation was characterized by low species richness and frequent dominance by the invasive species Elodea nuttallii . 6. Synthesis and applications . Our results show that stabilization of river water levels during low flow may negatively affect vegetation composition and succession in floodplain lakes adjacent to these rivers. A management scheme including incidental temporary lowering of the river water level, which results in drawdown of floodplain lakes, would enhance the ecological status of those rivers with stabilized water levels during low flow.

SUMMARY 1. We analysed the vegetation structure of 215 lakes in the flood plain of the river Lower Rhine in relation to environmental variables related to hydrological connectivity, lake morphometry, lake age and land use on adjacent land. 2. The frequency distribution of the cover of submerged macrophytes was not normal, implying that submerged macrophytes in any one lake were either scarce or abundant. 3. We observed clear water lakes with submerged macrophyte dominance over a wide range of total P concentration (0.020–0.40 mg total P L−1). 4. Multiple logistic regression indicated that the probability of dominance by submerged macrophytes decreased markedly with the surface area, depth and age of the lakes. The surface area effect occurred independently of the depth. Further, there was a negative relationship between submerged macrophyte dominance and the long‐term annual duration of inundation by the river. 5. Nymphaeid cover showed a distinct optimum with respect to mean lake depth, being almost absent in lakes shallower than 0.5 m. In contrast to what was found for submerged plants, the probability of occurrence of nymphaeids increased with lake age. 6. The probability of helophyte occurrence increased with lake age, and decreased with the presence of trees, cattle grazing, surface area, use of manure and mean lake depth. 7. In all cases the critical level of one factor (e.g. mean lake depth) depended on other factors (e.g. surface area or age of lake). Thus, in the present study, small lakes tended to remain dominated by submerged macrophytes up to a greater depth than large lakes, and helophytes colonised smaller lakes in an earlier phase. 8. The effect of inundation by the river was modest. This could be because most of our lakes are rarely inundated during the growing season and experience only moderate current velocities while flooded. 9. The results have practical implications for future management of flood plains for conservation purposes. In new water bodies, macrophyte domination will be promoted if many small shallow lakes, rather than few large deep ones, are excavated.

The characteristic ecology of floodplain lakes is in part due to their relatively strong water-level fluctuations. We analyzed the factors determining water-level fluctuations in 100 floodplain lakes (during nonflooded conditions) in the active floodplains of the Lower Rhine in the Netherlands. Furthermore, we explored the relationship between water-level fluctuations and macrophyte species richness, and analyzed the suitability of artificially created lakes for macrophyte vegetation. During non-flooded conditions along the Rhine, lake water-level fluctuations are largely driven by groundwater connection to the river. Hence, water-level fluctuations are largest in lakes close to the main channel in strongly fluctuating sectors of the river and smallest in isolated lakes. Additionally, water-level fluctuations are usually small in old lakes, mainly due to reduced groundwater hydraulic conductivity resulting from accumulated clay and silt on the bottom. Species richness of floating-leaved and emergent macrophytes was reduced at both small and large water-level fluctuations, whereas species richness of submerged macrophytes was reduced at small waterlevel fluctuations only. In addition, species richness of submerged macrophytes was higher in lakes that experienced drawdown, whereas no similar pattern was detected for floating-leaved and emergent macrophytes. The decline in amplitude of lake water-level with lake age implies that the number of hydrologically dynamic lakes will decrease over time. Therefore, we suggest that excavation of new lakes is essential to conserve the successional sequence of floodplain water bodies including conditions of high biodiversity. Shallow, moderately isolated, lakes with occasional bottom exposure have the highest potential for creating macrophyte-rich floodplain lakes along large lowland rivers. The water-level regime of such lakes can in part be designed, through choice of the location along the river, the distance away from the river and the depth profile of the lake.

Alternative stable states in shallow lakes have received much attention over the past decades, but less is known about transient dynamics of such lakes in the face of stochastic perturbations such as incidental extremes in water levels driven by climatic variability. Here, we report on the ecosystem dynamics of 70 lakes in the floodplains of the Lower Rhine in The Netherlands from 1999 to 2004. In any particular year, most lakes were either in a macrophyte-dominated clear state or in a contrasting state with turbid water and sparse submerged macrophyte cover. Macrophyte dominance was positively related to the occurrence of drawdown, and negatively to lake surface area and mean depth. We did not find a relation with nutrient levels. Remarkably, shifts between the two contrasting states were common, and episodes of low water levels appear to be an important external driver. A dry period before our study and the exceptionally dry summer of 2003 caused widespread drawdown of floodplain lakes, resulting in establishment of submerged macrophytes in the next year upon refill. In the 4 years without drawdown, many lakes returned to a macrophytepoor turbid state. Although some lakes turned turbid again quickly, others took several years to shift into the turbid state. A model analysis suggests that such prolonged transient vegetated states may be explained by the fact that the system dynamics slow down in the vicinity of the ''almost stable'' macrophyte-dominated state. Such a ''ghost'' of an equilibrium causes the system to stick around that state relatively long before slipping into the only true stable state. Our results support the idea that transient dynamics rather than equilibrium may be the key to understanding the overall state of some ecosystems. A practical implication of our findings is that artificial stabilization of the water level in shallow lakes may have been an important factor aggravating the permanent loss of submerged macrophytes due to cultural eutrophication.

A dramatic increase in the breeding population of geese has occurred over the past few decades at Svalbard. This may strongly impact the fragile ecosystems of the Arctic tundra because many of the ultra-oligotrophic freshwater systems experience enrichment from goose feces. We surveyed 21 shallow tundra ponds along a gradient of nutrient enrichment based on exposure to geese. Concentrations of total phosphorus (P) and dissolved inorganic nitrogen (DIN) in the tundra ponds ranged from 2-76 to 2-23 g l ¡1 respectively, yet there was no signiWcant increase in phytoplankton biomass (measured as chlorophyll a; range: 0.6-7.3 g l ¡1 ) along the nutrient gradient. This lack of response may be the result of the trophic structure of these ecosystems, which consists of only a two-trophic level food chain with high biomasses of the eYcient zooplankton grazer Daphnia in the absence of Wsh and scarcity of invertebrate predators. Our results indicate that this may cause a highly eYcient grazing control of phytoplankton in all ponds, supported by the fact that large fractions of the nutrient pools were bound in zooplankton biomass. The median percentage of Daphnia-N and Daphnia-P content to particulate (sestonic) N and P was 338 and 3009%, respectively, which is extremely high compared to temperate lakes. Our data suggest that Daphnia in shallow arctic ponds is heavily subsidized by major inputs of energy from other food sources (bacteria, benthic bioWlm), which may be crucial to the persistence of strong top-down control of pelagic algae by Daphnia.

1. We sampled a set of 93 lakes situated in the floodplains of the lower River Rhine in search for morphometric and other factors that explain their variation in clarity. 2. Lakes with a drop in summer water level were less turbid at the time of sampling, mainly because of a lower concentration of inorganic suspended solids (ISS). 3. We also found that older lakes were more turbid than younger lakes and that this was largely because of an increase in phytoplankton. 4. Water clarity was positively related to lake depth and the presence of vegetation. 5. Model calculations indicated that the underwater light climate was strongly affected by chlorophyll and ISS, the latter being the dominant factor affecting Secchi depth. Dissolved organic carbon (DOC) was less important. 6. The high concentration of ISS suggests that intensive resuspension occurs in most of the lakes. Using a simple wave model, and assuming that vegetation protects sediments against resuspension, we could eliminate wind resuspension as an important process in 90% of the lakes, leaving resuspension by benthivorous fish as probably the most important factor determining transparency. 7. Chlorophyll a concentration showed a strong positive correlation to ISS concentration, suggesting that resuspension may also have a positive effect on phytoplankton biomass in these lakes. 8. In conclusion, in-lake processes, rather than river dynamics, seem to be driving the turbidity of floodplain lakes along the lower River Rhine