Microcystin-producing blooms of Anabaenopsis arnoldi in a potable mountain lake in Saudi Arabia

This study reports for the first time the presence of Anabaenopsis arnoldi blooms in Saudi freshwaters. This species has been investigated with high cell densities (3.8 × 10³–264 × 10³ cells mL⁻¹) during June–November 2007 in Tendaha Lake, one of the major freshwater sources in Saudi Arabia. High temperature and conductivity, and a high concentration of phosphate, and low nitrate concentrations may have contributed to the formation of these blooms. The blooms were found to produce microcystins (MCYSTs) at concentrations up to 364 μg g⁻¹ dry weight as detected by an enzyme-linked immunosorbent assay. MCYSTs were also detected in the raw and treated water of the lake at concentrations (1.6–8.3 and 0.33–1.6 μg L⁻¹, respectively) exceeding the World Health Organization guideline level of 1 μg L⁻¹ for these toxins. HPLC analysis revealed that the extracts of A. arnoldi blooms contained MCYST-RR, -YR and two unidentified MCYSTs, but a pure culture of A. arnoldi isolated from Tendaha Lake during the present study produced MCYST-RR and –YR only. This is the first study to report MCYST production by A. arnoldi . Therefore, this cyanobacterium should be taken into consideration during monitoring of toxic cyanobacterial blooms in drinking and recreational water sources in the world, particularly arid and semi-arid countries including Saudi Arabia.     

Whitemere, a lake that defies some conventions about nutrients

1. Whitemere, a kettle-hole lake in north-west England (z_m,14 m, area 22.5 ha) has extremely high maximal concentrations (around 1 mg L⁻¹) of total phosphorus (TP) and soluble reactive phosphorus (SRP), and comparatively low maximal concentrations of dissolved inorganic nitrogen (<0.5 mg N L⁻¹). Bioassays indicate that its phytoplankton is nitrogen limited, and it has surface blue-green algal blooms in summer. Palaeoecological investigations have shown that high populations of cyanophytes occurred more than 6000 years ago, even before human settlement of the catchment.
2. The reasons for this combination of features are not immediately clear. This paper attempts to distinguish between two hypotheses: that the high phosphorus concentrations arise from input of phosphate-rich groundwater; and that the high phosphorus concentrations arise from concentration and recycling mechanisms within the lake.
3. The lake is entirely fed by ground water, direct rainfall and surface sheet flow. The concentrations of phosphorus in the ground water are much lower than in the lake water. The lake is stratified and has high phosphorus concentrations in the anaerobic hypolimnion in summer as a result of substantial release from the sediment. Vertical turbulence in the relatively weakly stratified water column may transfer substantial amounts to the epilimnion.There is also substantial phosphorus release from the sediments under the aerobic epilimnion.
4. There is little outflow of water and phosphorus. Most of the phosphorus is returned to the sediment under winter isothermal conditions, to be released again the following summer. The lake has probably maintained these mechanisms for millennia. Recent eutrophication is likely to have been driven by nitrogen inputs from greater intensification of agriculture in the catchment.     

Breakdown of tussock grass in streams along a gradient of agricultural development in New Zealand

Summary 1. We measured the breakdown rate of tussock grass in 12 New Zealand streams in catchments that provided a gradient of agricultural development. We also examined the microbial and invertebrate communities associated with decomposing tussock litter.
2. Pristine streams in the study had low concentrations of dissolved inorganic nitrogen (<10  μ g L⁻¹) and dissolved reactive phosphate (<3  μ g L⁻¹), whereas streams in the most developed catchments had high concentrations of nitrate (>2500  μ g L⁻¹) and phosphate (35  μ g L⁻¹), as well as greater amounts of suspended sediment and fine sediment covering the streambed.
3. Breakdown rate and microbial respiration were significantly related across the sites, and both were positively related to concentrations of nitrate and phosphate. Fungal biomass, measured as ergosterol, was positively related to microbial respiration and was also higher at sites with higher concentrations of nutrients. Total and shredding invertebrates were most abundant at the sites with high nutrient concentrations, but abundance of shredding invertebrates was not significantly related to breakdown rate. Amphipods were the most common shredding invertebrate at most sites, but probably did not contribute greatly to high rates of breakdown in streams in agricultural catchments.
4. With the exception of one site, nutrients from agricultural development appeared to have larger positive effects on litter breakdown than negative effects from sedimentation. Litter breakdown can serve as a functional measure of ecosystem health in streams, but caution should be exercised when a stress, such as land use, can have both positive (nutrients) and negative (sedimentation) effects.     

Impact of sediment and nutrient inputs on growth and survival of tadpoles of the Western Toad

1. Sediment and nutrient loading in freshwater systems are leading causes of aquatic habitat degradation globally. We investigated the impacts of fine-sediment and nutrient additions on the growth and survival of western toad ( Bufo boreas ) tadpoles and emergent metamorphs in mesocosm and exclosure experiments.
2. Mesocosm tanks received weekly pulses of fine sediments to create initial concentrations of 0, 130 and 260 mg L⁻¹ of suspended sediment and either bi-weekly additions of nutrients (N = 160 μg L⁻¹, P = 10 μg L⁻¹) or no additions in a factorial design. Within mesocosms, tadpole exclosures allowed for quantification of tadpole grazing pressure on periphyton biomass, chlorophyll- a and sediment deposition.
3. Tadpoles receiving sediment additions experienced slower growth rates and reduced survival to metamorphosis, although no effects of treatment were detected on size at metamorphosis or time to metamorphosis. Nutrient additions also lowered survival, but had no impact on other measured parameters of tadpole fitness. Dissections and gut content analysis revealed that tadpoles ingested sediment in large quantities altering the proportion of the organic content of ingested food.
4. Together these results suggest that although sediment was readily consumed by tadpoles, its presence in the larval environment had an overall negative effect on tadpole growth and survival, although not as severe as predicted.     

Constitutive mineralization of low concentrations of the herbicide linuron by a Variovorax sp. strain

The mineralization of the herbicide linuron at concentrations of μg and mg L⁻¹ was studied in liquid batch experiments with Variovorax sp. strain SRS16. The strain was highly efficient at mineralizing a range of linuron concentrations (0.002–10 mg L⁻¹) with 20–60% of the added ¹⁴C-ring-labeled linuron metabolized to ¹⁴CO₂ within hours to days depending on the initial linuron concentration and incubation period. At mg L⁻¹ linuron concentrations the mineralization activity by SRS16 was inducible and a shift to constitutive mineralization activity was apparent with a reduction in the linuron concentration to μg L⁻¹ levels. This study revealed that strain SRS16 is a promising candidate for bioaugmentation of water or soil resources contaminated with low linuron concentrations.     

Potassium dependence and phytoplankton ecology: an experimental study

SUMMARY 1. Unialgal cultures of three species common in the freshwater phytoplankton were used to test limitation of specific growth rate and final yield in defined media of low K⁺ concentration (range <0.3–6 μmol L⁻¹ or mmol m⁻³).
2. Growth rate of the diatom Asterionella formosa was independent of K⁺ concentration above 0.7 μmol L⁻¹. Final yield was dependent on initial concentration when accompanied by K⁺ depletion below this concentration, but not by lesser depletion with more residual K⁺. Analyses of particulate K in the biomass indicated a mean final cell content of 2.8 μmol K 10⁻⁸ cells, approximately 1.0% of the organic dry weight.
3. Less detailed work with the diatom Diatoma elongatum showed no dependence of growth rate or final yield upon the initial K⁺ concentration in the range 0.8–3.2 μmol L⁻¹. The phytoflagellate Plagioselmis nannoplanctica suffered net mortality in the lowest concentration tested, 0.8 μmol L⁻¹.
4. Comparison with the range of K⁺ concentration in natural fresh waters, including a depletion induced by an aquatic macrophyte, suggests that K⁺ is unlikely to limit growth of phytoplankton. Nevertheless, there can be correlation of K⁺ with lake trophy.     

The nutrient budget of a small eutrophic loch and the effectiveness of straw bales in controlling algal blooms

1. A 1-year intensive study of nutrient flows in Airthrey Loch, central Scotland, a small eutrophic [mean total phosphorus (TP) = 61.1 μg P l^–1] well-flushed freshwater body (area, 6.9 ha; mean depth, 1.85 m; volume, 1.274 × 10⁵ m³; retention time, 0.44 yr), was undertaken.
2. The nutrient budget was dominated by large allochthonous P inputs, equivalent to an areal load to the loch of 8.56 kg P ha^–1 yr^–1, which occurred predominantly during winter. In the summer, when TP inputs were low, water column levels of TP still increased, as a result of aerobic P release from sediments.
3. Sorption experiments indicated potential for sediment P release at water P concentrations of up to 200 μg P l^–1.
4. Aerobic release rate of P from sediments to the water column of Airthrey Loch was estimated to be of the order of 1 mg P m^–2 day^–1, and occurred during periods of elevated water column pH.
5. Straw bales placed in the loch to retard algal blooms were found not to have any demonstrable impact on algal concentrations observed during the study.     

Empirical relationships between profundal macroinvertebrate biomass and environmental variables in boreal lakes of Alberta, Canada

1. We empirically modelled profundal macroinvertebrate biomass (PMB) from collections of macroinvertebrates and a suite of trophic, water chemistry and morphometric variables from 26 lakes located within the Boreal Mixedwood and Boreal Subarctic ecoregions of Alberta, Canada.
2. Hypolimnetic dissolved oxygen (DO) concentrations (mg L⁻¹) explained 37% of the variance in PMB ((g m⁻²)^0.1) as a quadratic function, whereas water temperature (°C) explained 23% of variance in a linear relationship.
3. In contrast with other studies based on oligotrophic systems, indirect indicators of food availability (i.e. chlorophyll a and total phosphorus concentrations and Secchi depth) were not significant predictors of PMB in this study.
4. The predictive power of our models may be improved by obtaining more frequent (seasonal) estimates of PMB and hypolimnetic DO concentration and by quantifying biotic factors such as competition, predation, life history phenologies and recruitment success.     

The nutrient budget of a small eutrophic loch and the effectiveness of straw bales in controlling algal blooms

1. A 1-year intensive study of nutrient flows in Airthrey Loch, central Scotland, a small eutrophic [mean total phosphorus (TP) = 61.1 μg P l^–1] well-flushed freshwater body (area, 6.9 ha; mean depth, 1.85 m; volume, 1.274 × 10⁵ m³; retention time, 0.44 yr), was undertaken.
2. The nutrient budget was dominated by large allochthonous P inputs, equivalent to an areal load to the loch of 8.56 kg P ha^–1 yr^–1, which occurred predominantly during winter. In the summer, when TP inputs were low, water column levels of TP still increased, as a result of aerobic P release from sediments.
3. Sorption experiments indicated potential for sediment P release at water P concentrations of up to 200 μg P l^–1.
4. Aerobic release rate of P from sediments to the water column of Airthrey Loch was estimated to be of the order of 1 mg P m^–2 day^–1, and occurred during periods of elevated water column pH.
5. Straw bales placed in the loch to retard algal blooms were found not to have any demonstrable impact on algal concentrations observed during the study.     

Inorganic carbon limitation of photosynthesis in lake phytoplankton

1. Inorganic carbon availability influences species composition of phytoplankton in acidic and highly alkaline lakes, whereas the overall influence on community photosynthesis and growth is subject to debate.
2. The influence of total dissolved inorganic carbon (DIC) and free CO₂ on community photosynthesis was studied in six Danish lakes during the summer of 1995. The lakes were selected to ensure a wide range of chlorophyll a concentrations (1–120 μg l^–1), pH (5.6–9.6) and DIC concentration (0.02–2.5 m m ). Photosynthesis experiments were performed using the ¹⁴C technique in CO₂-manipulated water samples, either by changing the pH or by adding/removing CO₂.
3. Lake waters were naturally CO₂ supersaturated during most of the experimental period and inorganic carbon limitation of photosynthetic rates did not occur under ambient conditions. However, photosynthesis by phytoplankton in lakes with low and intermediate DIC concentrations was seriously restricted when CO₂ concentrations declined. Similarly, photosynthesis was limited by low CO₂ concentrations during phytoplankton blooms in the hardwater alkaline lakes.     

Contribution of rotifers to the diet and fitness of Boeckella (Copepoda: Calanoida)

1. The calanoid copepods, Boeckella triarticulata Sars and Boeckella hamata Brehm, are major components of the freshwater zooplankton of New Zealand. It was not known whether these copepods ingest rotifers, nor whether the inclusion of rotifers in their diets might improve their fitness. The present study aimed to identify rotifer taxa which are eaten by each copepod species, and to examine the fitness consequences of the inclusion of one species of rotifer in the diet of B. triarticulata .
2. In feeding experiments using natural rotifer assemblages, both species of copepod ingested the rotifer Anuraeopsis fissa (0.4–4% of daily carbon intake), and B. triarticulata also ingested Polyarthra dolichoptera (6–30% of daily carbon intake) and Keratella cochlearis tecta (1% of daily carbon intake).
3. The contribution of rotifers to the fitness of adult female B. triarticulata was assessed by comparing survival and reproduction among five diets that contained varying densities of algae ( Cryptomonas sp.) and/or Polyarthra dolichoptera (4 μg C L⁻¹). Boeckella triarticulata produced fewer clutches on a rotifer-only diet than on a solely algal diet, and addition of rotifers to a threshold algal diet did not affect copepod fitness relative to the solely algal diets. The present results suggest that Polyarthra at 4 μg C L⁻¹ is not a high-quality food for B. triarticulata .     

Spatio-temporal variation in macroinvertebrate assemblages of glacial streams in the Swiss Alps

1. Changes in water chemistry, benthic organic matter (BOM), and macroinvertebrates were examined in four different glacial streams over an annual cycle. The streams experienced strong seasonal changes in water chemistry that reflected temporal changes in the influence from the source glacier, especially in water turbidity, particulate phosphorus and conductivity.
2. Nitrogen concentrations were high (nitrate-N values were 130–274 μg L^–1), especially during spring snowmelt runoff. Benthic organic matter attained >600 g m^–2 dry mass at certain times, peaks being associated with seasonal blooms of the alga Hydrurus foetidus .
3. Macroinvertebrate taxon richness was two to three times higher (also numbers and biomass) in winter than summer suggesting winter may be a more favourable period for these animals. Benthic densities averaged 1140–3820 ind. m^–2, although peaking as high as 9000 ind. m^–2. Average annual biomass ranged from 102 to 721 mg m^–2, and reached >2000 mg m^–2 at one site in autumn.
4. Taxa common to all sites included the dipterans Diamesa spp. and Rhypholophus sp., the plecopterans Leuctra spp. and Rhabdiopteryx alpina , and the ephemeropterans Baetis alpinus and Rhithrogena spp. Principal components analysis clearly separated winter assemblages from those found in summer.     

Differing effects of suspended sediments on the performance of native and exotic Daphnia

1. Although large cladocerans are usually uncommon in large rivers, Daphnia lumholtzi (an exotic species in North America) is widespread and occasionally abundant. We collected zooplankton on the Illinois River (Illinois, U.S.A.) in 1995 and 1996 and found that the peak density of D. lumholtzi (25 L⁻¹) typically exceeded that of all native species combined. Maximum density occurred during warm periods (up to 27 °C) when concentrations of inorganic suspended sediments were high (>50 mg L⁻¹).
2. Using a life table experiment, we examined the effects of variation in suspended sediment (0 and 80 mg L⁻¹) and food (10⁴ and 10⁵ Ankistrodesmus cells mL⁻¹) on fitness of D. lumholtzi and the native Daphnia parvula. Daphnia lumholtzi had greater survivorship than D. parvula in most treatments and higher life-time fertility than D. parvula in all treatments. Both species achieved their fastest intrinsic rates of growth in treatments with high food, but their responses to suspended solids differed. The growth rate of D. lumholtzi in high food was slightly increased by higher turbidity, whereas that of D. parvula was depressed.
3. Results suggest that the ability of D. lumholtzi to tolerate suspended solids is an important factor contributing to its success in invading North American rivers.     

EFFECTS OF CADMIUM TOXICITY ON GROWTH AND ELEMENTAL COMPOSITION OF MARINE PHYTOPLANKTON

Some classes of marine phytoplankton are believed to be more tolerant of high concentrations of trace metals than others, but the results of experimental tests of this hypothesis are ambiguous. Eleven species of phytoplankton representing five classes were grown in Aquil medium containing Cd concentrations between 10⁻⁸ and 10⁻⁵ M ([Cd²⁺]= 10^−9.85 to 10^−6.84 M), and growth rates and intracellular concentrations of Cd, C, N, and S were measured. The mean Cd²⁺ concentration (pCd⁵⁰) that reduced the growth rate of each species to 50% of its maximum varied by 2.5 orders of magnitude, from 10^−6.23 for Emiliania huxleyi to 10^−8.79 for Synechococcus sp. Taxonomic trends in Cd resistance were not apparent in these data. Cadmium quotas (mol Cd·L⁻¹ cell volume) were lowest in species of Bacillariophyceae (ANOVA, P < 0.001), suggesting that they might regulate Cd transport differently than other taxa. Cellular S:C molar ratios increased in four of seven phytoplankton grown at high pCd (7.37–6.84) compared to low Cd ion concentrations (no added Cd), a result of increases in S·L⁻¹ cell volume. Nitrogen:carbon molar ratios were also higher in Cd-exposed phytoplankton, as changes in N and S were highly correlated ( r = 0.98, P < 0.0001). In two species that were examined, S:C ratios increased as a linear function of increasing Cd concentration. The results demonstrate large variability in Cd resistance among phytoplankton that is primarily a function of interspecific differences in Cd detoxification.     

160 Copepodology for the Phycologist with Apologies to G. E. Hutchenson

Heterocapsa triquetra is one of the most common bloom forming dinoflagellates found in estuaries and near shore regions around the world. In order to bloom, H. triquetra optimizes a suite of factors including low grazing pressure, increased nutrient inputs, alternative nutrient sources, and favorable salinity and hydrodynamic conditions, as well as the negative factors of temperature-limited growth, short day lengths, and periods of transient light limitation. The prevailing environmental conditions associated its wintertime blooms are largely the result of atmospheric forcing. Low-pressure systems moved through coastal area at frequent intervals and are accompanied by low air temperatures and rainfall. Runoff following the rainfall events supplies nutrients critical for bloom initiation and development. Heterocapsa triquetra blooms can reach chl a levels >100 mg L⁻¹ and cell densities between 1 to 6×106 L⁻¹. As the blooms develop, nutrient inputs from the river became insufficient to meet growth demand and H. triquetra feeds mixotrophically, reducing competition from co-occurring phytoplankton. Cloud cover associated with the low-pressure systems light limit H. triquetra growth as do low temperatures. More importantly though, low temperatures limit micro and macrozooplankton populations to such an extent that grazing losses are minimal.     

The occurrence and ecological importance of dissolved ATP in fresh water

SUMMARY. Dissolved ATP, defined as ATP which passes through 0.2 μm filters, was found in fresh water. During the spring diatom bloom in two eutrophic Danish lakes, concentrations of dissolved ATP varied between 0.1 and 3.8 μgl⁻¹, constituting 14–76% of the total ATP (particulate plus dissolved ATP). The kinetics of the light emission obtained from mixing firefly enzyme with dissolved ATP demonstrated that the major proportion of the dissolved ATP was in fact ATP. Despite some variations, the seasonal changes in dissolved ATP paralleled the changes in the increasing phytoplankton population during the rise of the diatom blooms. The dissolved ATP increased after the diatom peak, indicating that release of ATP from the phytoplankton due to mortality may be a major source of dissolved ATP.
Consumption of dissolved ATP was evaluated in uptake experiments using ³H-ATP. Rates of uptake of ³H-ATP by micro-organisms (diameter 0.2–0.6 μm) proved to be close to the rates for ³H-D-glucose uptake. The variations in ³H-ATP uptake during the diatom blooms showed non-systematic changes and ranged between 1.0 and 15.8% h⁻¹ (mean = 4.9% h⁻¹) of the quantity added. Turnover rates for dissolved ATP varied between 12 and 730 ng l⁻¹ h⁻¹ (mean = 175 ng l⁻¹). These rather high rates of turnover suggest that dissolved ATP is an important compound in the metabolism of freshwater bacteria.     

DO PHYSICAL FACTORS REGULATE PHYTOPLANKTON DISTRIBUTION PATTERNS IN LARGE, SHALLOW LAKES?

Factors that regulate phytoplankton dynamics in shallow, productive lakes are poorly understood, due to their predisposition for frequent algal blooms and sediment resuspension events. In Lake Apopka, greatest phytoplankton biomass reflects wind-induced resuspension of algae (meroplankton) that exists on the aphotic lake bottom in a layer approximately 5 cm thick; this assemblage is dominated by diatoms (>60% of total biomass) that can occur in resting stages. Once exposed to moderate light, meroplankton are capable of growth and photosynthetic rates comparable with surface populations. In Lake Okeechobee, remote sensing was used to assess the basin-wide distribution of suspended particles. Satellite reflectance values agreed well with in situ particle densities at 20 in-lake stations (average r²; LANDSAT = 0.81, AVHRR = 0.53), and maps of algal blooms (r² = 0.79, p ≤ 0.01). The greatest chlorophyll concentrations occurred in the vicinity of tributary nutrient inputs at the lake's perimeter, while turbidity increased towards the center of the lake reflecting predominant water circulation patterns. These results underscore the importance of physical-biological interactions in lakes.     

Taxonomic composition and biomass of heterotrophic flagellates in relation to lake trophy and season

1. Taxonomic composition and abundance of heterotrophic flagellates (HF) were studied in 55 lakes with different trophy in northern Germany using a live-counting technique.
2. Mean abundances and biomasses of HF ranged from 169 cells L⁻¹ and 22 μg L⁻¹ in mesotrophic lakes to 2439 cells mL^–1 and 475 μg L⁻¹ in hypertrophic lakes, respectively. Highest values were generally observed in spring, but mesotrophic lakes showed maximum values in early summer.
3. The taxonomic composition of HF was not significantly influenced by lake trophy and season. The major fraction of HF consisted of chrysomonads and Protista incertae sedis; other important groups were choanoflagellates and bicosoecids. The size distribution of HF changed with lake trophy and season, with a higher proportion of large HF (> 10 μm) in hypertrophic lakes and in spring, respectively.
4. Correlation analyses revealed a strong negative impact of cladocerans on total HF biomass and especially on large HF. Ciliates and large bacteria (> 10 μm) were strongly positively correlated with HF biomass; small bacteria (< 2 μm) showed a weak positive correlation.
5. Analyses at the level of species and genera revealed distinct distribution patterns of some taxa. Paraphysomonas , Aulacomonas and Quadricilia as large-bodied HF showed highest abundance in hypertrophic lakes and in spring. Attached taxa (e.g. Monosiga , Salpingoeca amphoridium ) were highly abundant in late summer and autumn, whereas Spumella and Kathablepharis occurred frequently in most samples.     

Effects of cadmium stress on periphytic diatom communities in indoor artificial streams

SUMMARY 1. Two experiments were performed with periphytic diatoms originating from the River Lot (France) and allowed to grow on clean substrata within indoor artificial streams. Three cadmium (Cd) levels (control, low Cd=10 μg L⁻¹ and high Cd=100 μg L⁻¹) were used to test the effects of Cd on (i) the settlement and development of diatom communities (Experiment 1, 4 weeks Cd exposure) and (ii) predeveloped communities grown over a 2-week period without contamination (Experiment 2, 2 weeks Cd exposure).
2. Experiment 1 revealed that growth and taxonomic composition of diatom communities clearly differed with Cd exposure. Biofilms were more adpressed to substrata under low Cd concentration, and were thinner and patchy under high Cd concentration.
3. Exposure of developed diatom communities (Experiment 2) to Cd revealed only minor variations in taxonomic composition, possibly linked to the protective role of the developed organic matrix against metal stress.
4. These results support the validity of periphytic diatom communities as indicators of metal pollution, although significant structural changes would take longer within developed communities.     

Stoichiometry of consumer-driven nutrient recycling across nutrient regimes in streams

Stoichiometric constraints within ecological interactions and their ecosystem consequences may depend on characteristics of the abiotic environment such as background nutrient levels. We assessed whether consumer identity, via differing body stoichiometry, could regulate periphyton stoichiometry across nutrient regimes in open systems. In 60 flow-through artificial streams, we factorially crossed dissolved inorganic nitrogen levels (elevated = 294  μ g L⁻¹, ambient = 26  μ g L⁻¹) with dissolved inorganic phosphorus levels (DIP: elevated = 15  μ g L⁻¹, ambient = 3  μ g L⁻¹) and consumer type [crayfish (body N : P = 18), snails (body N : P = 28) or a control]. At ambient DIP, periphyton in the crayfish treatment had a lower %P and a lower C : P than periphyton in the snail treatment suggesting that consumer identity, probably mediated by differing P-excretion, regulated periphyton P content. At high DIP, consumer identity no longer affected periphyton elemental composition. Therefore, the stoichiometry of consumer-driven nutrient recycling and consumer identity may be less important to ecosystem functioning in environments with elevated nutrient levels.