Periphyton responds differentially to nutrients recycled in dissolved or faecal pellet form by the snail grazer Theodoxus fluviatilis

1. We aimed to separate the effects of grazers on periphyton via grazing from that of nutrient recycling from their faecal pellets. 2. We set up three different experimental treatments (snails/no snails/faecal pellets) and sampled over 16 days. The ‘snail’ treatment contained a low density (snail biomass c. 14 g^?2) of the gastropod grazer Theodoxus fluviatilis and the ‘faecal pellet’ treatment received the same amount of faecal pellets as were produced in the ‘snail’ treatment. Whereas the ‘faecal pellet’ treatment provided extra nutrients to periphyton from the faeces, the ‘snail’ treatment provided nutrients in the form of both faeces and in excreta. There was also direct grazing on periphyton in the ‘snail’ treatment. The ‘no snail’ was not grazed and received no nutrients in faeces or excreta. 3. We measured periphyton C, N and P content, chlorophyll‐a (chl‐a), primary production, bacterial biomass, bacterial production and bacterial respiratory activity. In the water column we measured dissolved inorganic N and soluble reactive P. 4. Snails increased the amount of dissolved inorganic N in the water. On day 16, the periphyton N : P ratio in the ‘faecal pellet’ treatment was lower, and periphyton P content was higher, than in the other two treatments. N : P ratios decreased over time in the ‘faecal pellet’ treatment. Primary and bacterial production were positively correlated in all treatments. 5. Algal chl‐a and primary production of periphyton per unit area and periphyton chl‐a : C ratios increased over the 16 day in the ‘snail’ treatment, and thus excretion of dissolved N by snails had a stronger positive effect on the periphyton community than N and P in faecal pellets. 6. Our data show that excretion and egestion can have different effects on periphyton, probably because of the higher proportion of dissolved N in excreta and the higher proportion of P recycled in faecal pellets. The relative effect of nutrients recycled in egesta or in excretions, probably depends on the form of nutrient limitation of the periphyton. Further, the different components of the periphyton matrix could react differently to the different forms of nutrient recycling. 7. We conclude that direct grazing effects are less important than nutrient effects when nutrients are limiting and grazing pressure is low. Further, the spatial separation of different grazing effects can lead to differences in periphyton production and nutrient stoichiometry. This might be an explanation for the patchiness of periphyton in nature.     

Parasite and host elemental content and parasite effects on host nutrient excretion and metabolic rate

Ecological stoichiometry uses the mass balance of elements to predict energy and elemental fluxes across different levels of ecological organization. A specific prediction of ecological stoichiometry is the growth rate hypothesis (GRH), which states that organisms with faster growth or reproductive rates will require higher phosphorus content for nucleic acid and protein synthesis. Although parasites are found ubiquitously throughout ecosystems, little is understood about how they affect nutrient imbalances in ecosystems. We (1) tested the GRH by determining the carbon (C), nitrogen (N), and phosphorus (P) content of parasitic trematodes and their intermediate host, the freshwater snail Elimia livescens, and (2) used this framework to determine the trematode effects on host nutrient excretion and metabolism. Snail and parasite tissues were analyzed for elemental content using a CHN analyzer and soluble reactive phosphorus (SRP) methods. Ammonium and SRP assays were used to estimate N and P excretion rates. A respirometer was used to calculate individual snail metabolism. Trematode tissues contained lower C:P and N:P (more P per unit C and N) than the snail tissues. Snail gonadal tissues more closely resembled the elemental content of parasite tissues, although P content was 13% higher in the gonad than the trematode tissues. Despite differences in elemental content, N and P excretion rates of snails were not affected by the presence of parasites. Parasitized snails maintained faster metabolic rates than nonparasitized snails. However, the species of parasite did not affect metabolic rate. Together, this elemental imbalance between parasite and host, and the altered metabolic rate of infected snails may lead to broader parasite effects in stream ecosystems.     

Effects of macrograzers and light on periphyton stoichiometry

Ecological stoichiometry describes the biochemical constraints of trophic interactions emerging from the different nutrient content and nutrient demand of producers and consumers, respectively. Most research on this topic originates from well-mixed pelagic food webs, whereas the idea has received far less attention in spatially structured habitats. Here, we test how light as well as grazing and nutrient regeneration by consumers affects growth and biomass of benthic primary producers. In the first laboratory experiment, we manipulated grazer presence (two different snail species plus ungrazed control), in the second experiment we factorially combined manipulation of grazer presence and light intensity. We monitored snail and periphyton biomass as well as dissolved and particulate nutrients (nitrogen and phosphorus) over time. Grazers significantly reduced algal biomass in both experiments. Grazers affected periphyton nutrient content depending on the prevailing nutrient limitation and their own body stoichiometry. In the nitrogen (N-) limited first experiment, grazers increased N both in the periphyton and in the water column. The effect was stronger for grazers with lower N-content. In the phosphorus (P-) limited second experiment, grazers increased the P-content of the periphyton, but the grazer with lower N-content had additionally positive effects on algal N. Light reduction did not affect periphyton biomass, but increased chlorophyll-, N- and P-content of the periphyton. These experiments revealed that the indirect effects of grazers on periphyton were bound by stoichiometric constraints of nutrient incorporation and excretion.     

Feedbacks of consumer nutrient recycling on producer biomass and stoichiometry: separating direct and indirect effects

Herbivores can have both direct (consumptive) and indirect (nutrient‐mediated) effects on primary producer biomass and nutrient stoichiometry. Ecological stoichiometry theory predicts that herbivores of contrasting body stoichiometry will differentially remineralize nutrients, resulting in feedbacks on producer stoichiometry. We experimentally separated direct and indirect effects of aquatic vertebrate grazers on periphyton by manipulating grazer abundance and identity in mesocosms, and using grazer exclusion cages to expose periphyton to recycled nutrients in the absence of direct grazing. In experiment 1, we used a catfish with high body phosphorus (low body N:P), Ancistrus triradiatus, to assess consumptive versus nutrient‐mediated effects of grazer density on periphyton. In experiment 2, we compared the nutrient‐mediated effects of grazing by Ancistrus triradiatus and Rana palmipes, a tadpole with low body phosphorus and high body N:P. In experiment 1, we found that increasing catfish density led to lower biomass and particulate nutrients in periphyton through direct consumptive effects, but that nutrient‐mediated indirect effects enhanced periphyton biomass when grazers were experimentally separated from direct contact with periphyton. As predicted by stoichiometry theory, nutrient recycling by this P‐rich grazer tended to increase algal C:P and N:P (although effects were not statistically significant), while their consumptive effects reduced algal C:P and N:P. In experiment 2, grazer identity had strong effects on dissolved water nutrient concentrations, N recycling (measured with a ¹⁵N tracer), and periphyton stoichiometry. In accordance with stoichiometry theory, catfish increased N concentrations and recycling rates leading to higher periphyton N:P, while tadpoles had greater effects on P availability leading to lower periphyton N:P. Our experiments elucidate the importance of both the density and identity of grazers in controlling periphyton biomass and stoichiometry through consumptive and nutrient‐mediated effects, and support the power of ecological stoichiometry theory to predict feedbacks on producer stroichiometry arising from consumer stoichiometry through nutrient recycling.     

Competition and facilitation within and between a snail and a mayfly larva and the effect on the grazing process

We studied the competitive effects within and between two taxonomically distant freshwater herbivores, a snail and a mayfly, common in Swedish lakes, Lymnaea peregra and Cloeon dipterum, respectively, and their effect on grazing in a laboratory experiment. The experimental set-up consisted of 2-l aquaria, each containing a periphyton covered tile. Intra- and interspecific effects were tested by increasing the density of one species at a time in four different treatments, (1) snails (intraspecific treatment), (2) mayflies (intraspecific treatment), (3) mixed-snails (interspecific treatments, snails kept constant) and (4) mixed-mayflies (interspecific treatments, mayflies kept constant). Intraspecific competition affected both snails and mayflies negatively, i.e. increasing mortality with increasing con-specific density. Furthermore, there was a decrease in snail growth with increasing snail density. In the mixed-species treatments both species changed their microhabitat use indicating interspecific competition. Despite this, we also found a positive effect of mayfly density on snail growth, most likely due to indirect commensalism. No density-dependent effect of grazing on periphyton was found, probably due to interference competition between grazers. However, there was a significant difference in periphyton biomass, due to species composition of grazers. Irrespective of their densities, if they co-existed, the two grazer species decreased the periphyton biomass significantly compared with both single-species treatments. We considered this as a joint action of facilitation and interaction. Our results suggest that competition can be an important structuring factor in macroinvertebrate communities and that species composition can be significant for ecosystem processes within lentic environments.     

Nutrient loading and grazing by the minnow Phoxinus erythrogaster shift periphyton abundance and stoichiometry in mesocosms

1. Anthropogenic activities in prairie streams are increasing nutrient inputs and altering stream communities. Understanding the role of large consumers such as fish in regulating periphyton structure and nutritional content is necessary to predict how changing diversity will interact with nutrient enrichment to regulate stream nutrient processing and retention. 2. We characterised the importance of grazing fish on stream nutrient storage and cycling following a simulated flood under different nutrient regimes by crossing six nutrient concentrations with six densities of a grazing minnow (southern redbelly dace, Phoxinus erythrogaster) in large outdoor mesocosms. We measured the biomass and stoichiometry of overstory and understory periphyton layers, the stoichiometry of fish tissue and excretion, and compared fish diet composition with available algal assemblages in pools and riffles to evaluate whether fish were selectively foraging within or among habitats. 3. Model selection indicated nutrient loading and fish density were important to algal composition and periphyton carbon (C): nitrogen (N). Nutrient loading increased algal biomass, favoured diatom growth over green algae and decreased periphyton C : N. Increasing grazer density did not affect biomass and reduced the C : N of overstory, but not understory periphyton. Algal composition of dace diet was correlated with available algae, but there were proportionately more diatoms present in dace guts. We found no correlation between fish egestion/excretion nutrient ratios and nutrient loading or fish density despite varying N content of periphyton. 4. Large grazers and nutrient availability can have a spatially distinct influence at a microhabitat scale on the nutrient status of primary producers in streams.     

A manipulative study of macroinvertebrate grazers in Hong Kong streams: do snails compete with insects?

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Nitrogen processing by grazers in a headwater stream: riparian connections

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Do submerged aquatic plants influence their periphyton to enhance the growth and reproduction of invertebrate mutualists?

It has been suggested that submerged aquatic plants can influence the nutritional quality of the periphyton which grows on their surfaces, making it more nutritious for grazing invertebrates, particularly snails. In return, these grazers might preferentially feed on the periphyton and clear the plants of a potential competitor, with the plants and grazers both gaining from this mutualistic relationship. A highly replicated experiment was conducted, in which the nature of the plant (isoetid and elodeid types compared with similar-shaped inert substrata), the nutrient loading, and the influence of periphyton grazers (the bladder snail, Physa fontinalis) of similar size and history were controlled. Plant growth and survival significantly increased in the presence of the periphyton grazer. Whilst the presence of the grazers had the largest influence on periphyton abundance, nutrient availability and plant type also had effects. Plant type had little influence on the nutritional quality of the periphyton measured as carbohydrate, protein and C:N. Effects of treatment on snail growth, and the timing and extent of snail reproduction disappeared when they were compared with the quantity of periphyton available. There was no evidence of enhanced grazer success in the presence of the live plants compared with inert substrata. Although submerged plants affect the growth and reproduction of the grazers which feed on their surfaces, through differences in the amount of periphyton which grows there, we found no evidence that they manipulate the periphyton to encourage such grazers. Received: 1 September 1998 / Accepted: 12 May 1999     

Ecological stoichiometry of indirect grazer effects on periphyton nutrient content

Ecological stoichiometry has been successful in enhancing our understanding of trophic interactions between consumer and prey species. Consumer and prey dynamics have been shown to depend on the nutrient composition of the prey relative to the nutrient demand of the consumer. Since most experiments on this topic used a single consumer species, little is known about the validity of stoichiometric constraints on trophic interactions across consumers and ecosystems. We conducted a quantitative meta-analysis on grazer–periphyton experiments to test (1) if benthic grazers have consistent effects on the nutrient composition of their prey, and (2) whether these effects can be aligned to the nutrient stoichiometry of grazer and periphyton, other environmental factors, or experimental constraints. Grazers significantly lowered periphyton C:N and C:P ratios, indicating higher N- and P-content of grazed periphyton across studies. Grazer presence on average increased periphyton N:P ratios, but across studies the effect size did not differ significantly from zero. The sign and strength of grazer effects on periphyton nutrient ratios was strongly dependent on the nutrient content of grazers and their food, but also on grazer biomass, the amount of biomass removal and water column nutrients. Grazer with low P-content tended to reduce periphyton P-content, whereas grazers with high P-content increased periphyton P-content. This result suggests that low grazer P-content can be an indication of physiological P-limitation rather than a result of having relatively low and fixed P-requirements. At the across-system scale of this meta-analysis, predictions from stoichiometric theory are corroborated, but the plasticity of the consumer nutrient composition has to be acknowledged. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Current velocity influences the facilitation and removal of algae by stream grazers

The modification of flows in lotic ecosystems can have dramatic effects on abiotic and biotic processes and change the structure of basal trophic levels. In high-gradient streams, most of the biota are benthic, and decreased flow may homogenize and reduce benthic current velocity, potentially changing stream ecosystem function. Grazing by macroinvertebrates is an important component of stream function because grazers regulate energy flow from primary producers to higher trophic levels. We conducted an experiment to examine how macroinvertebrate grazers facilitated or removed algal biomass across a gradient of benthic current velocity (0–40 cm s^?1). We chose three grazers (Drunella coloradensis, Cinygmula spp., and Epeorus deceptivus) from a montane stream and conducted our experiment using 24 artificial stream channels that had three treatments: no grazers (control), single-grazer, and combined-grazer treatments. In the absence of grazers, algal biomass increased with benthic current velocity. Grazer treatments differed from the control in that more algal biomass was removed at higher velocities, whereas algal accrual was largely facilitated at low velocities. The transition from facilitation to removal ranged from 4.5 to 5.9 cm s^?1 for individual grazer treatments and occurred at 11.7 cm s^?1 for the combined-grazer treatment. Our data suggest that velocity plays a significant role in the facilitation and removal of algae by macroinvertebrate grazers. Additionally, the patterns revealed here could have general implications for algal accrual in systems where flow is reduced.     

Nuisance biomass levels of periphytic algae in streams

Relative coverage of filamentous periphytic algae increased with chlorophyll a (chl a) biomass on natural substrata in 22 northwestern United States and Swedish streams. A biomass range of 100–150 mg chl a m⁻² may represent a critical level for an aesthetic nuisance; below those levels, filamentous coverage was less than 20%. Other indices of water quality (dissolved oxygen content and measures of benthic macroinvertebrate diversity) were apparently unaffected by periphytic biomass or filamentous coverage in these streams. Neither was biomass related to limiting nutrient content (soluble reactive phosphorus, SRP), as has been observed in previous experiments using bare rocks in streams and slides in artificial channels. Ambient SRP concentration may not be a useful predicter of periphyton accrual on natural substrates, due to uptake and recycling of P throughout the stream and undetermined losses such as sloughing and grazing.     

Invertebrate grazing and riparian shade as controllers of nuisance algae in a eutrophic river

1. Algal growth in lotic systems is controlled either from the bottom‐up (e.g. nutrients and light, which determine growth rates) or from the top‐down (e.g. grazing pressure, which reduces accumulated biomass). Nutrient‐enriched streams that support large and diverse grazing macroinvertebrate populations and those with shaded riparian corridors rarely suffer from excessive algal growth. 2. In this study, the density of benthic algivorous macroinvertebrates was experimentally manipulated in shaded and open nutrient‐enriched stream habitats of the Owennagearagh River, south‐west Ireland. The ability of macroinvertebrate grazers and riparian shade to control benthic algal growth [particularly the nuisance alga, Cladophora glomerata (L. Kütz)] was investigated. Three sites with markedly different concentrations of plant nutrients (one site upstream and two sites downstream of the sewage outfall) were selected. The density of grazing invertebrates colonising ceramic tiles was reduced using high‐voltage localised electric pulses. Replicates of treatment (grazer‐excluded) and control (grazed) tiles were deployed in open and shaded (<25 and >80% canopy cover, respectively) patches of stream bed, in each site. 3. After 2‐week Cladophora cover, periphytic chlorophyll a and biofilm ash‐free dry mass (AFDM) were quantified for all experimental tiles. Values for all three parameters were highest on grazer‐excluded tiles from open patches. Grazed tiles from open patches accrued little Cladophora and had significantly lower levels of chlorophyll a and AFDM. Nutrient inputs were found to have an impact on the density of grazing invertebrates, with higher densities of Baetis nymphs at the most nutrient‐enriched site. 4. Our results demonstrate that in eutrophic, high‐light streams, filamentous algae can quickly accumulate to nuisance levels in the absence of invertebrate grazers. In future, greater attention should be paid to the role of grazing invertebrates in controlling nuisance algae in streams, in addition to algal–nutrient relationships.     

Upward cascading effects of nutrients: shifts in a benthic microalgal community and a negative herbivore response

We evaluated the effects of nutrient addition on interactions between the benthic microalgal community and a dominant herbivorous gastropod, Cerithidea californica (California horn snail), on tidal flats in Mugu Lagoon, southern California, USA. We crossed snail and nutrient (N and P) addition treatments in enclosures on two tidal flats varying from 71 to 92% sand content in a temporally replicated experiment (summer 2000, fall 2000, spring 2001). Diatom biomass increased slightly (~30%) in response to nutrient treatments but was not affected by snails. Blooms of cyanobacteria (up to 200%) and purple sulfur bacteria (up to 400%) occurred in response to nutrient enrichment, particularly in the sandier site, but only cyanobacterial biomass decreased in response to snail grazing. Snail mortality was 2–5 times higher in response to nutrient addition, especially in the sandier site, corresponding to a relative increase in cyanobacterial biomass. Nutrient-related snail mortality occurred only in the spring and summer, when the snails were most actively feeding on the microalgal community. Inactive snails in the fall showed no response to nutrient-induced cyanobacterial growths. This study demonstrated strongly negative upward cascading effects of nutrient enrichment through the food chain. The strength of this upward cascade was closely linked to sediment type and microalgal community composition.     

Snail responses to cues produced by an invasive decapod predator

Abstract. Cues released by predators and injured prey often induce shifts in prey behavior that allow prey to evade predators, but also affect prey resource use. I investigated the effects of chemical and mechanical signals produced by injured snails (Physella gyrina) and predatory crayfish (Procambarus clarkii) on microdistributions of P. gyrina. In an initial experiment, I observed snail responses to the presence of a caged crayfish predator, to injured conspecifics, or to both. There were significant effects of time and the treatment × time interaction on the proportion of snails moving above the water line, with greater proportions of snails above the water line at night than during the day and with weak snail crawl‐out behavior being elicited by caged crayfish at night, but not during the day. In a second experiment, I examined snail microdistributions when exposed to crayfish confined to a small cage within each aquarium, crayfish confined to half of each aquarium, and crayfish ranging freely throughout each aquarium. Snails responded most strongly to free‐ranging crayfish by moving above the water line, but also demonstrated significant, but reduced, crawl‐out responses to crayfish confined to half of each aquarium; however, snails did not respond behaviorally to crayfish confined to small cages. In both experiments, there were marginally significant effects of unfed caged crayfish on the proportions of snail populations hiding under benthic shelters, with this response being the strongest at the start of the experiments but weak overall (with only 4–5% of P. gyrina responding in each experiment). These results indicate that cues (e.g., chemical or mechanical) produced by predators altered prey microdistributions, but that the exact prey responses (e.g., moving above the water line or into horizontal or benthic refugia) depended on the intensity and nature of cues.     

Effects of phosphorus enrichment and grazing snails on modern stromatolitic microbial communities

1. The effects of phosphorus enrichment and grazing snails on a benthic microbial community that builds stromatolic oncolites were examined in an experiment at Rio Mesquites, Cuatro Ciénegas, Mexico. Chemical analyses of stream water samples indicated that overall atomic ratios of total nitrogen (N) to total phosphorus (P) were approximately 110, indicating a strong potential for P‐limitation of microbial growth. 2. Phosphorus enrichment involved addition of 5 μmol Na₂HPO₄ L^?1 to streamside microcosms receiving intermittent inputs of stream water while grazer manipulation involved removal of the dominant grazer, the snail Mexithauma quadripaludium. After 7 weeks, we examined responses in organic matter content, C : N : P ratios, metabolism (P removal, primary production, dark respiration, and calcification), and microbial community structure using molecular fingerprinting of 16S rRNA genes. 3. Manipulation of snails did not affect response variables measured in these treatments (organic matter, C : P ratio, P removal rate). However, P enrichment significantly decreased the C : P and N : P ratios of surficial materials in the oncolites (organic matter content was unchanged), increased net and gross photosynthesis (oxygen consumption in the dark was unchanged), increased rates of calcification, and increased diatoms relative to cyanobacteria. Heterotrophic Eubacteria and Archaea were only modestly affected. Thus, our results indicate weak grazing effects but strong impacts of P in this benthic system. 4. We hypothesise that a state of severe P‐limitation is imposed on autotrophic production in this food web due, at least in part, to co‐precipitation of phosphate during calcite deposition. This produces severe P‐limitation of the benthic algae and cyanobacteria, resulting in high C : P ratio of microbial mats relative to the biomass of photoautotrophs (phytoplankton, terrestrial foliage) in other ecosystems. In turn, this high C : P ratio is likely to generate severe stoichiometric constraints on the herbivores, thus limiting their populations and resulting in weak overall grazing impacts.     

Spatial and seasonal variation in nutrient excretion by benthic invertebrates in a eutrophic reservoir

1. Nitrogen (N) and phosphorus (P) fluxes via excretion by benthic invertebrates were quantified in a eutrophic reservoir (Acton Lake, Ohio, U.S.A.). We quantified variation in nutrient fluxes seasonally (June until November 1997), spatially (three sites) and among taxa (chironomids, tubificid oligochaetes and Chaoborus). 2. The three taxa differed in spatial distribution and contribution to nutrient fluxes. Tubificids were the most abundant taxon at two oxic sites (1.5 and 4 m depth), and were exceedingly rare at an anoxic, hypolimnetic site (8 m). Chironomids were abundant only at the shallowest oxic site. Chaoborus was the only abundant taxon at the anoxic site. Total benthic invertebrate biomass was greatest at the shallowest site and lowest at the anoxic, hypolimnetic site. 3. Mass‐specific excretion rate [μmol NH₄–N or soluble reactive P (SRP) excreted mg dry mass^–1 h^–1] varied among experiments and was influenced by temperature. Differences among taxa were not significant. Thus, nutrient flux through benthic invertebrates was affected more by total invertebrate biomass and temperature than by species composition. 4. Fluxes of N and P via benthic invertebrate excretion (μmol NH₄–N or SRP m^–2 day^–1) were greatest at the oxic sites, where fluxes were dominated by the excretion of tubificids and chironomids. The N and P fluxes at the anoxic site were much lower, and were dominated by excretion by Chaoborus. The ratio of N and P excreted by the benthic invertebrate assemblage varied seasonally and was lowest at the anoxic site. 5. Comparison with other measured inputs shows that excretion by benthic invertebrates could be an important source of nutrients, especially of P. However, the relative importance of nutrient excretion by the benthos varies greatly spatially and temporally.     

Role of the polychaete <Emphasis Type="Italic">Neanthes succinea</Emphasis> in phosphorus regeneration from sediments in the Salton Sea,California

The Salton Sea currently suffers from several well-documented water quality problems associated with high nutrient loading. However, the importance of phosphorus regeneration from sediments has not been established. Sediment phosphorus regeneration rates may be affected by benthic macroinvertebrate activity (e.g. bioturbation and excretion). The polychaete Neanthes succinea (Frey and Leuckart) is the dominant benthic macroinvertebrate in the Salton Sea. It is widely distributed during periods of mixing (winter and spring), and inhabits only shallow water areas following development of anoxia in summer. The contribution of N. succinea to sediment phosphorus regeneration was investigated using laboratory incubations of cores under lake temperatures and dissolved oxygen concentrations typical of the Salton Sea. Regeneration rates of soluble reactive phosphorus (SRP) were lowest (−0.23–1.03 mg P m⁻² day⁻¹) under saturated oxygen conditions, and highest (1.23–4.67 mg P m⁻² day⁻¹) under reduced oxygen levels. N. succinea most likely stimulated phosphorus regeneration under reduced oxygen levels via increased burrow ventilation rates. Phosphorus excretion rates by N. succinea were 60–70% more rapid under reduced oxygen levels than under saturated or hypoxic conditions. SRP accounted for 71–80% of the dissolved phosphorus excreted under all conditions. Whole-lake SRP regeneration rates predicted from N. succinea biomass densities are highest in early spring, when the lake is mixing frequently and mid-lake phytoplankton populations are maximal. Thus, any additional phosphorus regenerated from the sediments at that time has potential for contributing to the overall production of the lake. Guest Editor: John M. Melack Saline Water and their Biota     

Grazer control of nutrient availability in the periphyton

Summary Benthic algal assemblages are regulated by both abiotic (e.g., nutrient) and biotic (e.g., grazing) constraint. The objective of this study was to determine how changes in these two factors affected the structure of an algal assemblage in an ephemeral stream. Coverslips were incubated for 21 days in enclosures containing one of three nutrient environments (ambient, phosphorus-enriched, or phosphorus and nitrogen enriched) and one of four densities of the snail Gonibasis (0, 40, 80, or 120 snails/m²) and examined directly to enumerate the algal assemblage. The effect of grazing on algal biomass was dependent on the nutrient environment. An overstory of diatoms was susceptible to removal by grazing and was not strongly affected by nutrient enrichment. An understory of Stigeoclonium was more resistant to grazing and responded strongly to nutrient enrichment only in the presence of grazers. Snail grazers may mediate nutrient availability to the understory indirectly by removing overlying cells or by direct excretion of nutrients. Multiple interactions occur between benthic herbivores and algae, and, as shown here, some of them are positive and involve modifications of the nutrient environment.