Long Transients Near the Ghost of a Stable State in Eutrophic Shallow Lakes with Fluctuating Water Levels

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 macrophyte-poor 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.     

Relations between trophic state indicators and plant biomass in Florida lakes

We collected quantitative data on macrophyte abundance and water quality in 319 mostly shallow, polymictic, Florida lakes to look for relationships between trophic state indicators and the biomasses of plankton algae, periphyton, and macrophytes. The lakes ranged from oligotrophic to hypereutrophic with total algal chlorophylls ranging from 1 to 241 mg m^–3. There were strong positive correlations between planktonic chlorophylls and total phosphorus and total nitrogen, but there were weak inverse relationships between the densities of periphyton and the trophic state indicators total phosphorus, total nitrogen and algal chlorophyll and a positive relationship with Secchi depth. There was no predictable relationship between the abundance of emergent, floating-leaved, and submersed aquatic vegetation and the trophic state indicators. It was only at the highest levels of nutrient concentrations that submersed macrophytes were predictably absent and the lakes were algal dominated. Below these levels, macrophyte abundance could be high or low. The phosphorus–chlorophyll and phosphorus–Secchi depth relationships were not influenced by the amounts of aquatic vegetation present indicating that the role of macrophytes in clearing lakes may be primarily to reduce nutrient concentrations for a given level of loading. Rather than nutrient concentrations controlling macrophyte abundance, it seems that macrophytes acted to modify nutrient concentrations.     

Submerged aquatic vegetation correlations with depth and light attenuating materials in a shallow subtropical lake

A 3-year study was done to quantify the biomass of submerged aquatic vegetation (SAV) and its relationship with environmental attributes in Lake Okeechobee, the largest lake in the southeastern United States. Plants were sampled on 21 occasions at sites located along 15 fixed transects around the shoreline, giving rise to 721 observations of SAV species (Chara spp., Vallisneria americana, Hydrilla verticillata, Potamogeton illinoinensis) dry weight biomass. Environmental sampling focused on factors that attenuate light, including phytoplankton chlorophyll a (chl a), total suspended solids (TSS), non-volatile suspended solids (NVSS) and color. Depth and Secchi transparency also were measured. Based on regression analysis, NVSS was considerably more important in attenuating light than chl a or color. Total biomass of SAV varied from 0 to 271 g dw m^–2, with a mean of 4.7 g dw m^–2, and strong dominance by Chara. The SAV biomass was lower than average for Florida lakes, and may reflect the influence of suspended solids on underwater irradiance, as well as high water level in the late 1990s. Dense SAV was found only where depth was < 2 m and TSS < 20–30 mg l^–1. At locations where high biomass of SAV occurred, the plants may have influenced water quality, because concentrations of TSS, NVSS, and chl a were 2–3 fold lower than at sites with no plants. The potential effects of SAV also were apparent at a regional scale. The shoreline region of the lake displayed a pattern of rising and falling chl a and NVSS with water depth. This occurred both at sites with and without plants, suggesting that it may be driven by physical processes, such as water circulation patterns, which are influenced by depth. However, the pattern was dampened at sites with SAV, indicating a potential to influence these attributes of water quality.     

Impacts of a submerged plant (Elodea canadensis) on interactions between roach (Rutilus rutilus) and its invertebrate prey communities in a lake littoral zone

1. Using 5‐m² field enclosures, we examined the effects of Elodea canadensis on zooplankton communities and on the trophic cascade caused by 4–5 year old (approximately 16 cm) roach. We also tested the hypothesis that roach in Elodea beds use variable food resources as their diet, mainly benthic and epiphytic macroinvertebrates, and feed less efficiently on zooplankton. Switching of the prey preference stabilises the zooplankton community and, in turn, also the fluctuation of algal biomass. The factorial design of the experiment included three levels of Elodea (no‐, sparse‐ and dense‐Elodea) and two levels of fish (present and absent). 2. During the 4‐week experiment, the total biomass of euplanktonic zooplankton, especially that of the dominant cladoceran Daphnia longispina, decreased with increase in Elodea density. The Daphnia biomass was also reduced by roach in all the Elodea treatments. Thus, Elodea provided neither a favourable habitat nor a good refuge for Daphnia against predation by roach. 3. The electivity of roach for cladocerans was high in all the Elodea treatments. Roach were able to prey on cladocerans in Elodea beds, even when the abundance and size of these prey animals were low. In addition to cladocerans, the diet of roach consisted of macroinvertebrates and detrital/plant material. Although the biomass of macroinvertebrates increased during the experiment in all Elodea treatments, they were relatively unimportant in roach diets regardless of the density of Elodea beds. 4. Euplanktonic zooplankton species other than Daphnia were not affected by Elodea or fish and the treatments had no effects on the total clearance rate of euplanktonic zooplankton. However, the chlorophyll a concentration increased with fish in all the Elodea treatments, suggesting that fish enhanced algal growth through regeneration of nutrients. Thus, our results did not unequivocally show that Elodea hampered the trophic cascade of fish via lowered predation on grazing zooplankton. 5. In treatments with dense Elodea beds (750 g FW m^?2), chlorophyll a concentration was always low suggesting that phytoplankton production was controlled by Elodea. Apparently, the top‐down control of phytoplankton biomass by zooplankton was facilitated by the macrophytes and operated simultaneously with control of phytoplankton production by Elodea.     

Effects of fish and nutrient additions on food-web stability in a charophyte-dominated lake

1. The response of major food‐web constituents to combinations of nutrient addition and zooplanktivorous fish abundance was tested during two subsequent years in the shallow charophyte‐dominated lake Naardermeer in the Netherlands, using in situ enclosures. 2. Treatment effects differed sharply between study years. In 1998, when the summer temperature was low (17–21 °C), high algal biomass only developed at high nutrient levels in the presence of fish, but with no major effect on Chara biomass. In 1999, when the summer temperature was relatively high (20–25 °C), algal blooms occurred at high nutrient levels regardless of fish abundance, and were associated with a drastic decline in Chara biomass. 3. Differences between years in temperature and initial zooplankton composition and biomass were likely to contribute to the varying relative importance of top‐down and bottom‐up effects in these enclosure experiments. 4. The results suggest that when nutrient loads are increased towards levels where the macrophyte‐dominated state is being destabilised, a ‘switch’ is more likely to occur in warm summers.     

Effects of waterfowl, large fish and periphyton on the spring growth of Potamogeton pectinatus L. in Lake Mogan, Turkey

It has been argued that waterfowl and fish may threaten growth of submerged macrophytes, especially in spring during the early growth phase when plant biomass is low. A small reduction of biomass at that time might delay growth or decrease subsequent productivity. We investigated the impact of waterfowl and large fish on the spring growth of fennel pondweed (Potamogeton pectinatusL.) by employing an exclosure experiment in the macrophyte-dominated clear-water Lake Mogan, Turkey. Birds and large fish were excluded from eight plots and both in situvegetation and macrophytes kept in pots were compared to eight open plots. Also, to investigate the effect of periphyton on plant growth it was removed from half of the pot plants. Exclusion of waterfowl and fish may decrease predation on macroinvertebrates, which in turn may affect periphyton, and macrophyte growth, why macroinvertebrates also were sampled. Waterfowl density was high (15–70 ind. of coot, Fulica atraL. ha^–1), abundance of submerged plants was also high with a surface coverage of 70–80%, and benthivorous fish were present, mainly tench, (Tinca tincaL.) and carp, (Cyprinus carpioL.). Exclusion of waterfowl and large fish did not significantly affect the spring growth of pondweed; neither plants growing in situnor kept in pots. Removal of periphyton from the plants in the pots did not favour growth. The density of macroinvertebrates was not affected by the exclusion of waterfowl and large fish, but it was positively related to aboveground biomass of fennel pondweed. We suggest that even if waterfowl and large fish are in high densities, their effect on fennel pondweed spring growth in lakes with abundant submerged vegetation, such as Lake Mogan, is low.     

Ecological regime shifts in salinised wetland systems. II. Factors affecting the dominance of benthic microbial communities

This paper is the second in a pair investigating potential mechanisms for ecological regime change in salinising wetlands. The first paper in this series focused on the responses of the salt-tolerant submerged macrophyte community to salinity. In this second paper, we investigated some of the environmental conditions required for initiation and dominance of benthic microbial communities using a combination of experimental and observational data. Two experiments were carried out. One investigated the importance of prior establishment of benthic microbial communities on their ability to maintain prevalence over macrophyte colonisation (‘persistence’ experiment), while the other investigated hydrology and its effect on sediment perturbation, potential nutrient release and subsequent benthic microbial community establishment (‘flooding’ experiment). The ‘persistence’ experiment measured the biomass of benthic microbial communities and emergence of macrophytes from sediments kept either wet or dry for 4 weeks then flooded at a range of salinities. Benthic microbial biomass was similar across all of the salinities tested (15, 45 and 70 ppt), with a slight increase at higher salinities, suggesting that none of these limited benthic microbial community development. Pre-wetting of sediments usually increased benthic microbial community biomass and reduced macrophyte germination, but the latter was attributed to the presence of anoxic sediments rather than the increased benthic microbial community biomass. Germinating macrophytes emerged through benthic microbial communities or dense heterotrophic bacterial blooms, demonstrating that they could become dominant even when another community was already established. Field data supported these results, suggesting that the development of benthic microbial communities is not limited by salinity alone, but includes other factors, such as the water regime. In the ‘flooding’ experiment, the largest differences in nutrient concentrations ultimately lay between the pre-wet and pre-dry treatments (due to the greater release of nutrients and development of anoxia in the latter) rather than those subjected to fast versus slow flooding. In response to this, highest benthic microbial community biomass was in treatments with pre-wet sediment, corresponding with lower phytoplankton biomass.     

Short-term and long-term effects of zooplanktivorous fish removal in a shallow lake: a synthesis of 15 years of data from Lake Zwemlust

SUMMARY 1. Removal of zooplanktivorous fish (mainly bream) in 1987 from a shallow eutrophic lake in the Netherlands, Lake Zwemlust, resulted in a quick switch from a turbid state with cyanobacteria blooms to a clear state dominated by macrophytes.
2. The clear state was not stable in the long term, however, because of high nutrient loadings.
3. In 1999, another removal of zooplanktivorous fish (mainly rudd) had similar effects as in 1987, although macrophytes returned more slowly.
4. In the years directly following both interventions there was a 'transition period' of very clear water with high densities of zooplanktonic grazers in the absence of macrophytes; low oxygen concentrations indicate that during those years primary production was low relative to heterotrophic activity.
5. The transition period appears to provide the light climate necessary for the return of macrophytes.
6. Reduction of nutrient loading is necessary to improve water quality in Lake Zwemlust in the long term. In the short term, repeated fish stock reduction is a reasonable management strategy to keep Lake Zwemlust clear.     

Long‐term dynamics of submerged macrophytes and algae in a small and shallow,eutrophic lake: implications for the stability of macrophyte‐dominance

1. Submerged macrophyte and phytoplankton components of eutrophic, shallow lakes have frequently undergone dynamic changes in composition and abundance with important consequences for lake functioning and stability. However, because of a paucity of long‐term survey data, we know little regarding the nature, direction and sequencing of such changes over decadal–centennial or longer timescales. 2. To circumvent this problem, we analysed multiple (n = 5) chronologically correlated sediment cores for plant macro‐remains and a single core for pollen and diatoms from one small, shallow, English lake (Felbrigg Hall Lake, Norfolk, U.K.), documenting 250 years of change to macrophyte and algal communities. 3. All five cores showed broadly similar stratigraphic changes in macrophyte remains with three distinct phases of macrophyte development: Myriophyllum–Chara–Potamogeton (c. pre‐1900), to Ceratophyllum–Chara–Potamogeton (c. 1900–1960) and finally to Zannichellia–Potamogeton (c. post‐1960). Macrophyte species richness declined from at least 10 species pre‐1900 to just four species at the present day. Additionally, in the final Zannichellia–Potamogeton phase, a directional shift between epi‐benthic and phytoplankton‐based primary production was indicated by the diatom data. 4. Based on macrophyte–seasonality relationships established for the region, concomitant with the final shift to Zannichellia–Potamogeton, we infer a reduction in the seasonal duration of plant dominance (plant‐covered period). Furthermore, we hypothesise that this change in species composition resulted in a situation whereby macrophyte populations were seasonally ‘sandwiched’ between two phytoplankton peaks in spring and late summer as observed in the contemporary lake. 5. We suggest that eutrophication‐induced reductions in macrophyte species richness, especially if the number of plant‐seasonal strategies is reduced, may constrict the plant growing season. In turn, this may render a shallow lake increasingly vulnerable to seasonal invasions of phytoplankton resulting in further species losses in the plant community. Thus, as part of a slow (over perhaps 10–100s of years) and self‐perpetuating process, macrophytes may be gradually pushed out by phytoplankton without the need for a perturbation as required in the alternative stable states model of plant loss.     

Incorporating Natural Variability into the Assessment of Ecological Health in Australian Dryland Rivers

Dryland rivers occur over much of Australia’s inland and have some of the most variable patterns of flow in the world. Despite their dominance in terms of Australia’s river types, dryland rivers have not been the focus of the recent increase in research on indicators of river health, mostly owing to their spatial remoteness and relatively low levels of water resource development and human disturbance. Most rivers in arid and semi-arid regions are ephemeral, and only carry significant flows during the wetter months or following infrequent but intense rainfall events. It is not known which, if any, of the existing approaches to river health assessment can be used to accurately assess the health of these large ephemeral rivers. This paper considers why the standard methods for interpreting the currently-used indicators for river health may need to be adapted for variable systems and suggests the use of trends that recognise natural variation in indicator values for undertaking this.     

How a bottom‐dweller beats the canopy: inhibition of an aquatic weed (Potamogeton pectinatus) by macroalgae (Chara spp.)

1. Bottom‐dwelling charophytes have been observed to replace canopy‐forming pondweeds within a few years in de‐eutrophied shallow lakes. Competition for bicarbonate (HCO₃^?) may explain this shift in vegetation dominance but inhibition of pondweeds by Chara spp. through direct competition has not been shown experimentally. 2. We tested whether charophytes inhibited growth of fennel pondweed (Potamogeton pectinatus) in the absence of belowground competition by growing plants in pots in mesocosms following a replacement series experimental design. To further understand the role of bicarbonate, we studied main and interactive effects of Chara, light and bicarbonate on P. pectinatus growth in a laboratory study. 3. Early in the mesocosm experiment, high charophyte densities had a negative effect on P. pectinatus cover at a time when bicarbonate levels were low. However, bicarbonate levels eventually converged to low levels in all treatments. At final harvest, both species exhibited lower biomasses at higher densities of conspecific pots, indicating that ultimately intraspecific competition was limiting. In a laboratory study, Chara inhibited P. pectinatus the most under a combination of high light and high bicarbonate concentrations, suggesting that Chara may negatively affect P. pectinatus by acting as a general nutrient sink. 4. Our results suggest that Chara growth can reduce bicarbonate levels, delaying but not preventing a P. pectinatus growth pulse. Given the recorded inhibition under ample bicarbonate supply, Chara’s ability to act as nutrient sink may contribute to the decline of P. pectinatus under Chara recovery in shallow lakes.     

The submerged macrophytes in Lake Maarsseveen I: Changes in species composition and biomass over a six year period

The submerged macrophytes of Lake Maarsseveen I were surveyed in 1983 using SCUBA diving techniques. Only 40% of the characeans and 75% of the angiosperms detected in 1977 remained. The area colonized by submerged macrophytes was 0.45% in 1983, compared with 25.10% in 1977. The observed decreases were largely attributable to a shift of the plantcolonized areas to shallower depths. By 1983, most of the earlier predominant vegetation types had disappeared and the biomass had decreased. The decline in submerged vegetation may be attributed to increasing eutrophication, fish populations and recreational activities.     

Existence of a macrophyte-dominated clear water state over a very wide range of nutrient concentrations in a small shallow lake

Little Mere, a small shallow lake, has been monitored for four years, since its main source of nutrients (sewage effluent) was diverted. The lake has provided strong evidence for the persistence of a clear water state over a wide range of nutrient concentrations. It had clear water at extremely high nutrient concentrations prior to effluent diversion, associated with high densities of the large body-sized grazer, Daphnia magna, associated with low fish densities and fish predation. Following sewage effluent diversion in 1991, the nutrient concentrations significantly declined, the oxygen concentrations rose, and fish predation increased. The dominance of large body-sized grazers shifted to one of relatively smaller body-sized animals but the clear water state has been maintained. This is probably due to provision of refuges for grazers by large nymphaeid stands (also found prior to diversion). There has been a continued decrease in nutrient concentrations and expansion of the total macrophyte coverage, largely by submerged plants, following effluent diversion. The grazer community of Little Mere has also responded to this latter change with a decline in daphnids and increase in densities of weed-associated grazers. The presence of large densities of such open water grazers was the apparent main buffer mechanisms of the clear water state until 1994. The lake has, so far, maintained its clear water in the absence of such grazers. Thus, new buffer mechanisms appear to operate to stabilize the ecosystem. Little Mere appears to have shifted from previous top-down controlled clear water state to a bottom-up controlled clear water state.     

The restoration of Lake Apopka in relation to alternative stable states: an alternative view to that of Bachmann et al. (1999)

Bachmann et al. (1999) postulated that wind energy initiated, and has maintained, high turbidity in hypertrophic (mean chlorophyll a = 92 g l^–1) Lake Apopka, Florida (mean depth = 1.6 m; area = 12500 ha). They asserted that the turbid condition was initiated by a hurricane in late 1947 that destroyed submersed plant beds and that high turbidity has since been maintained by wind-driven resuspension of fluid sediments. In their view, there has been sufficient light for re-establishment of submersed plants over about 38% of the lake bottom, but plant growth has been precluded by the fluid character of the sediments. They concluded that the restoration program of the St. Johns River Water Management District, which includes reduction of the phosphorus (P) loading rate, will not restore water clarity or submersed vegetation. An alternative explanation for Lake Apopka's turbid state is that it was initiated, and has been maintained, by excessive P loading that led to algal blooms and elimination of submersed vegetation through light limitation. The transition to the turbid state was contemporaneous with drainage of 7300 ha of the floodplain wetland to create polders for farming, beginning in the early 1940s. Lake P budgets indicate that drainage of the farms caused a seven-fold increase in the P loading rate (0.08 g TP m^–2 yr^–1 to 0.55 g TP m^–2 yr^–1). Paleolimnological analysis of lake sediments also indicates an increase in the P loading rate in mid-century, concomitant with the decline in submersed vegetation and the increase in phytoplankton abundance. After the increase in P loading, wind disturbance may have accelerated the transition to the turbid state; but, before the increase in P loading, wind disturbance was insufficient to elicit the turbid state, as evidenced by the stability of the clear-water state in the face of 14 hurricanes and 41 tropical storms from 1881 to 1946. Measurements of photosynthetically active radiation (PAR) indicate that light limitation has inhibited submersed plant growth except on the shallowest 5% of the lake bottom. Further, the correlation between the diffuse attentuation coefficient (K _PAR) and chlorophyll a (CHLA) indicates that light limitation would be removed over about 82% of the lake bottom with a reduction in CHLA from 92 g l^–1 to 25 g l^–1. Recently, following a 40% reduction in the P loading rate, the mean total P (TP) concentration, mean CHLA, and total suspended solids fell by about 30% while mean Secchi depth increased by more than 20%. Submersed plant beds appeared in areas devoid of macrophytes for nearly 50 years. These improvements, during a period with no change in mean wind speeds measured at Lake Apopka, provide the strongest evidence that the turbid state has been maintained by excessive P loading and that the current restoration program, which combines P load reduction with planting and removal of planktivorous fish, will be effective.     

Vegetation,fire and soil feedbacks of dynamic boundaries between rainforest,savanna and grassland

At fine spatial scales, savanna‐rainforest‐grassland boundary dynamics are thought to be mediated by the interplay between fire, vegetation and soil feedbacks. These processes were investigated by quantifying tree species composition, the light environment, quantities and flammability of fuels, bark thickness, and soil conditions across stable and dynamic rainforest boundaries that adjoin grassland and eucalypt savanna in the highlands of the Bunya Mountains, southeast Queensland, Australia. The size class distribution of savanna and rainforest stems was indicative of the encroachment of rainforest species into savanna and grassland. Increasing dominance of rainforest trees corresponds to an increase in woody canopy cover, the dominance of litter fuels (woody debris and leaf), and decline in grass occurrence. There is marked difference in litter and grass fuel flammability and this result is largely an influence of strongly dissimilar fuel bulk densities. Relative bark thickness, a measure of stem fire resistance, was found to be generally greater in savanna species when compared to that of rainforest species, with notable exceptions being the conifers Araucaria bidwillii and Araucaria cunninghamii. A transect study of soil nutrients across one dynamic rainforest – grassland boundary indicated the mass of carbon and nitrogen, but not phosphorus, increased across the successional gradient. Soil carbon turnover time is shortest in stable rainforest, intermediate in dynamic rainforest and longest in grassland highlighting nutrient cycling differentiation. We conclude that the general absence of fire in the Bunya Mountains, due to a divergence from traditional Aboriginal burning practices, has allowed for the encroachment of fire‐sensitive rainforest species into the flammable biomes of this landscape. Rainforest invasion is likely to have reduced fire risk via changes to fuel composition and microclimatic conditions, and this feedback will be reinforced by altered nutrient cycling. The mechanics of the feedbacks here identified are discussed in terms of landscape change theory.     

The effect of turbidity state and microhabitat on macroinvertebrate assemblages: a pilot study of six shallow lakes

Shallow lakes can occur in two alternative stable states, a clear-water state and a turbid state. This is associated with separate assemblages of fish, zooplankton and plants. Little is known about whether macroinvertebrate assemblages differ across both stable states. This study investigated this in a connected set of three turbid and three clear-water shallow lakes. To overcome confounding effects of differences in spatial structure of macrophytes in turbid and clear-water lakes, we sampled three microhabitats that occurred in both alternative stable states: open water, sago pondweed (Potamogeton pectinatus) and reed (Phragmites australis). Univariate analyses indicated no differences in the number of organisms, taxon richness or diversity between turbid and clear-water lakes. Multivariate analysis, however, showed significant differences in the macroinvertebrate community structure of both stable states. Nine taxa explained a significant amount of the variation between both lake types, of which seven preferred the clear-water lakes. The number of organisms and the taxon richness were higher in reed than in the other microhabitats, but diversity and evenness did not differ among the microhabitats. Multivariate analyses could separate all three microhabitats. Eight taxa, mainly detritus feeders and collector–gatherers, explained most of the variation in the data and preferred the reed microhabitat. The effects of stable state (6.8% explained variance) and microhabitat (13.1% explained variance) on the macroinvertebrate assemblages were largely independent from each other (1.5% shared variance). Although macroinvertebrates are not implemented in the initial theory of stable states, our results show clearly different assemblages across both stable states.     

Cyanobacterial dynamics in shallow Lake Apopka (Florida,U.S.A.) before and after the shift from a macrophyte‐dominated to a phytoplankton‐dominated state

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Response of the flora and macroinvertebrate fauna of a chalk stream site to changes in management

1. Temporal changes in a series of habitats and their macroinvertebrate assemblages were examined on a 50‐m section of a chalk stream in Berkshire, England between June 1975–79 and June 1997–2001. 2. The site was part of a trout fishery in 1975–79, when river management included instream weed cutting together with control of bankside trees and riparian vegetation. Management ceased in the 1980s and by 1997–2001, the site was heavily shaded by trees and riparian vegetation. 3. The mean area of instream macrophytes decreased by 50% between the first and second sampling period. In contrast, gravel and silt increased and invading marginal vegetation formed a new habitat. 4. Changes in macroinvertebrate family richness between sampling periods were scale dependant. Although there were, on average, significantly more families in individual replicates in 1975–79 than in 1997–2001, total family richness for the site in each year did not differ significantly between sampling periods. 5. Sixty families of macroinvertebrates were recorded during the study, 50 in both sampling periods, 53 in 1975–79 and 57 in 1997–2001. This small increase in site family richness may be due to the invading marginal plants. 6. Total macroinvertebrate abundance was significantly lower in the second sampling period. A major drought in 1976 resulted in significantly higher densities of macroinvertebrates, partly through the exploitation of epiphytic diatoms by chironomid larvae. A drought in 1997 failed to elicit a similar response because of the limited macrophytes and diatoms under heavy shading by trees and marginal vegetation. 7. Significant increases in important shredders and decreases in some scrapers between the early and later sampling years largely reflected changes in available food resources. 8. Whereas macroinvertebrate family richness has been conserved under the recent ‘no management’ regime, the site is now less attractive as a fishery because of poor access and lower densities of some macroinvertebrates taken by brown trout.