The role of light availability and herbivory on algal responses to nutrient enrichment in a riparian wetland,Alaska

We investigated how the relative availability of solar radiation in the presence or absence of grazing alters the ability of benthic algae to respond to nutrient enrichment in an Alaskan marsh. We used a factorial mesocosm experiment that included nutrient enrichment (enriched or control), grazing (grazed or ungrazed), and light (unshaded or shaded) to simulate shading by macrophytes early and late in the growing season, respectively. We found stronger effects of grazers and nutrients compared to light on benthic algal biomass and taxonomic composition. Algal biomass increased in nutrient‐enriched treatments and was reduced by grazing. Shading did not have an effect on algal biomass or taxonomic composition, but the concentration of chl a per algal biovolume increased with shading, demonstrating the ability of algae to compensate for changes in light availability. Algal taxonomic composition was more affected by grazer presence than nutrients or light. Grazer‐resistant taxa (basal filaments of Stigeoclonium) were replaced by diatoms (Nitzschia) and filamentous green algae (Ulothrix) when herbivores were removed. The interacting and opposing influences of nutrients and grazing indicate that the algal community is under dual control from the bottom‐up (nutrient limitation) and from the top‐down (consumption by herbivores), although grazers had a stronger influence on algal biomass and taxonomic composition than nutrient enrichment. Our results suggest that low light availability will not inhibit the algal response to elevated nutrient concentrations expected with ongoing climate change, but grazers rapidly consume algae following enrichment, masking the effects of elevated nutrients on algal production.     

Why are there few algae on snail shells? The effects of grazing,nutrients and shell chemistry on the algae on shells of Helisoma trivolvis

1. Freshwater snails often lack visible growths of algae on their shells. We tested three possible mechanisms that may account for this (grazing, snail-derived nutrients and chemical defences), using the ramshorn snail Helisoma trivolvis .
2. The experiments were carried out in floating plastic enclosures in a pond and comprised seven treatments. Grazing treatments were: a lone snail (ungrazed, as self-grazing does not occur), Helisoma with conspecifics, Helisoma with the co-occurring pond snail Physa sp., empty shells with Helisoma , and ungrazed empty shells. Nutrient effects were possible in all treatments with occupied shells (lone snail; Helisoma with conspecifics, and with Physa ) versus absent in other treatments. Testing for chemical defences compared algae on fresh empty shells, weathered shells (outer organic periostracum layer absent) and boiled fresh shells (with denaturization of susceptible proteins).
3. Diatoms dominated algal assemblages on snail shells. Although the upright diatom Gomphonema gracile was abundant on all shells, it was dominant on the shells of snails housed with other snail grazers (either Helisoma or Physa ).
4. Only the lone snail (nutrients but no grazing) showed higher algal biomass, so presumably any nutrient effect in the treatments with grazers was masked. Both Helisoma and Physa were observed apparently grazing on Helisoma shells, and consequently algal biomass in multi-snail treatments was similar to that on empty shells. Scanning electron microscopy revealed that algal density was highest near the aperture of live snails, but not empty shells; this is consistent with a nutrient addition effect. There was no evidence of chemical defences against algal growth.
5. In soft-bottomed freshwater habitats with abundant snails, shells of living snails provide nutrient-augmented substrata that may indirectly boost overall snail production.     

Increasing native, but not exotic, biodiversity increases aboveground productivity in ungrazed and intensely grazed grasslands

Species-rich native grasslands are frequently converted to species-poor exotic grasslands or pastures; however, the consequences of these changes for ecosystem functioning remain unclear. Cattle grazing (ungrazed or intensely grazed once), plant species origin (native or exotic), and species richness (4-species mixture or monoculture) treatments were fully crossed and randomly assigned to plots of grassland plants. We tested whether (1) native and exotic plots exhibited different responses to grazing for six ecosystem functions (i.e., aboveground productivity, light interception, fine root biomass, tracer nitrogen uptake, biomass consumption, and aboveground biomass recovery), and (2) biodiversity-ecosystem functioning relationships depended on grazing or species origin. We found that native and exotic species exhibited different responses to grazing for three of the ecosystem functions we considered. Intense grazing decreased fine root biomass by 53% in exotic plots, but had no effect on fine root biomass in native plots. The proportion of standing biomass consumed by cattle was 16% less in exotic than in native grazed plots. Aboveground biomass recovery was 30% less in native than in exotic plots. Intense grazing decreased aboveground productivity by 25%, light interception by 14%, and tracer nitrogen uptake by 54%, and these effects were similar in native and exotic plots. Increasing species richness from one to four species increased aboveground productivity by 42%, and light interception by 44%, in both ungrazed and intensely grazed native plots. In contrast, increasing species richness did not influence biomass production or resource uptake in ungrazed or intensely grazed exotic plots. These results suggest that converting native grasslands to exotic grasslands or pastures changes ecosystem structure and processes, and the relationship between biodiversity and ecosystem functioning.     

The snail <Emphasis Type="Italic">Potamopyrgus antipodarum</Emphasis> grows faster and is more active in the shade,independent of food quality

Ecological stoichiometry has advanced food web ecology by emphasising the importance of food quality over food quantity for herbivores. Here, we focus on the effects of abiotic factors such as nutrients and light (known to influence food quality) on grazer growth rates. As model organism we used the mudsnail Potamopyrgus antipodarum that is native to New Zealand but invasive elsewhere. In a stream channel experiment in New Zealand, we manipulated light (two levels) and nutrients (four levels) to achieve a range of primary producer carbon: nutrient ratios and added mudsnails (3 densities + ungrazed control) to 128 periphyton covered stream channels in a 2 × 4 × 4 full factorial design. We measured snail growth rate and activity, food quality and nutritional imbalance, to test the predictions that (1) less light and more nutrients increase periphyton food quality and thus snail growth rates, and (2) less crowding leads to higher food availability and thus higher snail growth rates. We found that snail growth rates were higher under low light than under high light intensities and this difference increased with increasing nutrient addition. These changes in growth rate were not mediated by food quality in terms of periphyton nutrient ratios. Furthermore, experimental treatments strongly affected snail behaviour. Snails grazed more actively in the low light treatments, and thus it is more likely that snail growth rates were directly affected by light levels, maybe as a result of innate predator avoidance behaviour or as a reaction to high UV intensities. We conclude that in our stream channels snail growth rate was limited by factors other than food quality and quantity such as UV exposure, algal defences or the relatively low ambient water temperature.     

EFFECTS OF IRRADIANCE AND GRAZING ON LOTIC ALGAL ASSEMBLAGES1

A laboratory experiment was conducted for 75 days to examine how irradiance levels and grazing influence algal biomass and community structure. Twelve laboratory streams were used for experimental analyses, with four channels exposed to one of three irradiance levels (15, 100, or 400 μE·m^?2·s^?1). Three of the four stream at each light level were stocked with the snail Juga silicula (250·m^?2), leaving one stream at each light level without snails. Grazed stream exposed to low light levels developed low amounts of algal biomass (<2 g AFDW·m^?2) and were dominated by adnately attached diatoms. Mean algal biomass increased over time in the grazed streams exposed to intermediate light; by day 75, these streams were characterized by moderate algal biomasses (30-40 g AFDW·m^?2) and filamentous chlorophytes. Algal assemblages in high light, grazed channels had high levels of biomass at day 43 (70 g AFDW·m^?2) that declined to 30 g AFDW·m^?2at day 75 and were dominated by chlorophytes. Among ungrazed streams, algal biomass at day 75 was relatively low in the low light streams (<7g AFDW·m^?2) and was dominated by adnately attached diatoms. Ungrazed streams exposed to intermediate and high light levels had moderate biomasses (23 and 19 g AFDW·m^?2, respectively) and were dominated by chlorophytes and large diatoms. Grazing appeared both to delay and alter successional trajectories of algal assemblages, with alterations most noticeable during early seral stages at intermediate and high light levels. Grazing had the least effect on successional trajectories at low light.     

Gastropod grazers and nutrients,but not light,interact in determining periphytic algal diversity

The potential interactions of grazing, nutrients and light in influencing autotroph species diversity have not previously been considered. Earlier studies have shown that grazing and nutrients interact in determining autotroph species diversity, since grazing decreases species diversity when nutrients (i.e. N or P) limit autotroph growth, but increases it when nutrients are replete. We hypothesized that increased light intensities would intensify the interactions between grazing and nutrients on algal species diversity, resulting in even stronger reductions in algal species diversity through grazing under nutrient–poor conditions, and to even stronger increases of algal species diversity through grazing under nutrient-rich conditions. We studied the effects of grazing (absent, present), nutrients (ambient, N + P enriched) and light (low light, high light) on benthic algal diversity and periphyton C:nutrient ratios (which can indicate algal nutrient limitation) in a factorial laboratory experiment, using the gastropod grazer Viviparus viviparus. Grazing decreased algal biomass and algal diversity, but increased C:P and N:P ratios of periphyton. Grazing also affected periphyton species composition, by decreasing the proportion of Spirogyra sp. and increasing the proportion of species in the Chaetophorales. Grazing effects on diversity as well as on periphyton N:P ratios were weakened when nutrients were added (interaction between grazing and nutrients). Chlorophyll a (Chl a) per area increased with nutrient addition and decreased with high light intensities. Light did not increase the strength of the interaction between grazing and nutrients on periphytic algal diversity. This study shows that nutrient addition substantially reduced the negative effects of grazing on periphytic algal diversity, whereas light did not interact with grazing or nutrient enrichment in determining periphytic algal diversity.     

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.     

Responses of stream algae to grazing minnows and nutrients: a field test for interactions

Summary Previous studies have shown that an algivorous grazing minnow (Campostoma anomalum) is the major herbivore in Brier Creek, a hardwater stream in south central Oklahoma. In summer and autumn schools of Campostoma virtually eliminate algae from substrate surfaces in deeper areas of some pools. The pool-to-pool distributions of algae and Campostoma reported for this stream could occur if nutrient limitation permits grazing by Campostoma to outrun algal growth. To test this hypothesis, mesh pens were built to exclude Campostoma from substrates in each of four typical Campostoma pools. N+P+K lawn fertilizer was added daily to two of the four pools; the other two, which received no fertilizer additions and which were not visibly affected by fertilizer transported downstream from the pools enriched with nutrients, served as controls. Algae accumulated rapidly on natural substrates and on unglazed ceramic tiles in grazer-exclusion pens in pools receiving N+P+K additions and more slowly in pens in both control pools. Periphyton biomass on grazed substrates in all four pools remained low throughout the experiment. Hence, Campostoma at normal densities were able to outrun algal growth even when nutrients were added. Eleven days after the experiment started, I determined biomass, biomass-specific net primary productivity, and areal net primary productivity of periphyton on substrates exposed to all combinations of grazer (+,0) and nutrient (+,0) treatments. Grazing increased biomass-specific primary productivity, prevented accumulation of biomass, and decreased areal primary productivity of periphyton. Additions of N+P+K increased biomass-specific net primary productivity of grazed and ungrazed periphyton and markedly increased biomass of periphyton on substrates protected from Campostoma. Although food supply for Campostoma appeared to be greater with nutrient additions, condition of Campostoma in pools receiving N+P+K was not significantly different from Campostoma collected from control pools 35 days after the experiment started. I conclude that although nutrient supply limits biomass-specific primary productivity of periphyton in Brier Creek, nutrient limitation in this stream exacerbates, rather than causes, the visually conspicuous pool-to-pool complimentary distribution of algae and Campostoma: in this stream, grazing by Campostoma at natural densities can outrun periphyton growth even when nutrients are added.     

Alterations in the biomass-specific productivity of periphyton assemblages mediated by fish grazing

1. In an experimental flume, we examined the effects of a biomass reduction and alteration of taxonomic composition, because of grazing by the fish Plecoglossus altivelis, on the net biomass accumulation of periphyton. 2. Grazed and ungrazed assemblages with different biomass and taxonomic composition were first prepared in fish enclosures and exclosures, respectively. These assemblages were then set out in the flume and incubated for 2 days under grazing‐free conditions to examine (i) the relationship between biomass and biomass accumulation rate and (ii) the effect of taxonomic composition on the relationship between these two. 3. The grazed and ungrazed assemblages were dominated by upright filamentous cyanobacteria and diatoms, respectively. The rate of biomass accumulation decreased with increasing periphyton biomass in both the grazed and ungrazed assemblages, and was lower in the grazed than the ungrazed assemblages at any biomass level. 4. The results showed that the reduction in biomass and the alteration of taxonomic composition due to fish grazing have opposite effects on biomass‐specific productivity. Biomass accumulation rate increased in response to biomass reduction, although a shift in dominance from diatoms to upright filamentous cyanobacteria decreased the overall productivity of the periphyton.     

Effects of the herbivorous minnow,southern redbelly dace (<Emphasis Type="Italic">Phoxinus erythrogaster</Emphasis>), on stream productivity and ecosystem structure

We used field and mesocosm experiments to measure effects of southern redbelly dace (Phoxinus erythrogaster), a grazing minnow, on stream ecosystem structure and function. Ecosystem structure was quantified as algal filament length, algal biomass, size distribution of particulate organic matter (POM), algal assemblage structure, and invertebrate assemblage structure, whereas ecosystem function was based on gross and net primary productivity. Our experiments showed that moderate densities of Phoxinus temporarily reduced mean algal filament length and mean size of POM relative to fishless controls. However, there was no detectable effect on algal biomass or ecosystem primary productivity. Several factors could explain the lack of effect of Phoxinus on primary productivity including increased algal production efficiency in grazed treatments or increased grazing by other organisms in fishless treatments. The inability of Phoxinus to reduce algal biomass and system productivity contrasts with experimental results based on other grazing minnows, such as the central stoneroller (Campostoma anomalum), and questions the generality of grazer effects in stream ecosystems. However, environmental venue and the spatial and temporal scale of ecosystem measurements can greatly influence the outcome of these experiments. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users. The experiments described herein comply with the current laws of the country in which the experiments were performed.     

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.     

Light,nutrients and grazing interact to determine stream diatom community composition and functional group structure

1. Benthic algal communities are shaped by the availability of nutrients and light and by herbivore consumption. Many studies have examined how one of these factors affects algal communities, but studies simultaneously addressing all three are rare. 2. We investigated the effects of nutrients, light and a herbivore (the snail Potamopyrgus antipodarum) on benthic stream algae in a fully factorial experiment in 128 circular streamside channels. Four nutrient levels (none added to highly enriched), four snail grazing pressures (no snails to 777 individuals m^?2) and two light levels (ambient and 65% reduced) were applied. Colonising algae were dominated by diatoms (Bacillariophyta), which were determined to species using acid‐cleaned samples and assigned to functional groups according to their physiognomic growth form. 3. Diatom community structure changed considerably in response to our manipulations. Light had the strongest influence (as indicated by manova effect size), whereas nutrients had an intermediate effect and grazing was fairly weak. Relative abundances of six common diatom taxa decreased under reduced light, whereas five others became more prevalent. Eight taxa benefitted from nutrient enrichment, while three became rarer. Grazing affected the relative density of only one common taxon, which increased at higher grazing pressure. 4. Diatom functional groups also responded strongly. ‘Low profile’ taxa dominated at low resource levels (nutrients and especially light), whereas ‘high profile’ and ‘motile’ taxa became markedly more prevalent at higher resource levels. 5. Two‐way interactions between experimental factors were quite common. For example, Planothidium lanceolatum and Rossithidium petersenii responded more strongly to nutrient enrichment at reduced than at ambient light, whereas Cocconeis placentula responded more strongly at ambient light. For diatom functional groups, the benefit of nutrient enrichment for ‘motile’ diatoms was greater at ambient than at reduced light. 6. Our results imply that multifactor experiments are required to determine the main forces driving the composition of benthic algal communities. Further, our findings highlight the considerable potential of using functional algal groups as indicators of changing environmental conditions to complement the traditional taxonomic approach.     

Does an increase in irradiance influence periphyton in a heavily-grazed woodland stream?

Summary Irradiance level and grazer density were manipulated in a factorial design to examine the relative effects of biotic and abiotic factors on periphyton biomass, productivity, and taxonomic structure in a heavily grazed, woodland stream. Irradiance levels were increased from 0.26 to 12.42 mol quanta/m²/d by placing metal halide lamps over the stream. The major grazer in this system was the prosobranch snail Elimia clavaeformis. Its densities were reduced from ca. 750 individuals/m² to near zero by raising platforms off the stream bottom. Experimental treatments were maintained for 48 days. Biomass-specific carbon fixation rates increased significantly in response to higher light levels, indicating that periphyton communities were light-limited at this time of year. However, positive effects of irradiance on areal-specific carbon fixation and biomass were detected only when grazer density was reduced. Basal cells of the chlorophyte Stigeoclonium dominated communities exposed either to low light or high grazing pressure. When light was increased and grazer density reduced, large or upright diatoms became more abundant. Results from this study indicated that limitation of periphyton photosynthesis could be mitigated by increasing the levels of an abiotic resource (light) to this system, but that periphyton biomass was controlled by biotic interactions.     

Grazing increases functional richness but not functional divergence in Tibetan alpine meadow plant communities

Plant community diversity and ecosystem function are conditioned by competition among co-occurring species for multiple resources. Previous studies suggest that removal of standing biomass by grazing decreases competition for light, but coincident grazing effects on competition for soil nutrients remain largely unknown in Tibetan rangelands where grazing tends to deplete soil phosphorus availability. We measured five functional traits indicative of plant productivity and stoichiometry leaf carbon concentration (LCC), leaf nitrogen concentration (LNC), leaf phosphorus concentration (LPC), specific leaf area (SLA), leaf dry matter content (LDMC) for component species of plant communities in grazed and ungrazed plots in five Tibetan alpine meadows. We examined the diversity of traits singly Rao index of functional diversity (FDrao) and in aggregate functional richness (FRic), functional divergence (FDiv), and functional evenness (FEve) in response to grazing. We tested whether foliar trait diversity increases with nutrient competition but decreases with light competition when competitive exclusion is reduced by grazing. The FDrao of LPC significantly increased under grazing, but FDrao for LCC, LNC and SLA tended to decrease. The FDrao of LDMC increased at the drier site but decreased at the wettest site. There was a strong negative association between increase in FDrao of LPC and decrease in soil nutrients, especially soil phosphorus availability. The FRic for all five traits together increased with species diversity following grazing, but neither FDiv nor FEve differed significantly between grazed and ungrazed plots at most sites. Grazing in Tibetan alpine meadows tends to increase competition for soil phosphorus while decreasing competition for light, resulting in an increase in the functional richness in grazed plant communities without any significant changes in the overall functional diversity of foliar traits. Our study highlights the potential importance of grazing mediated competition for multiple resources in alpine meadow ecosystems.     

Grazing history effects on above- and below-ground litter decomposition and nutrient cycling in two co-occurring grasses

Large herbivores may alter carbon and nutrient cycling in soil by changing above- and below-ground litter decomposition dynamics. Grazing effects may reflect changes in plant allocation patterns, and thus litter quality, or the site conditions for decomposition, but the relative roles of these broad mechanisms have rarely been tested. We examined plant and soil mediated effects of grazing history on litter mass loss and nutrient release in two grazing-tolerant grasses, Lolium multiflorum and Paspalum dilatatum, in a humid pampa grassland, Argentina. Shoot and root litters produced in a common garden by conspecific plants collected from grazed and ungrazed sites were incubated under both grazing conditions. We found that grazing history effects on litter decomposition were stronger for shoot than for root material. Root mass loss was neither affected by litter origin nor incubation site, although roots from the grazed origin immobilised more nutrients. Plants from the grazed site produced shoots with higher cell soluble contents and lower lignin:N ratios. Grazing effects mediated by shoot litter origin depended on the species, and were less apparent than incubation site effects. Lolium shoots from the grazed site decomposed and released nutrients faster, whereas Paspalum shoots from the grazed site retained more nutrient than their respective counterparts from the ungrazed site. Such divergent, species-specific dynamics did not translate into consistent differences in soil mineral N beneath decomposing litters. Indeed, shoot mass loss and nutrient release were generally faster in the grazed grassland, where soil N availability was higher. Our results show that grazing influenced nutrient cycling by modifying litter breakdown within species as well as the soil environment for decomposition. They also indicate that grazing effects on decomposition are likely to involve aerial litter pools rather than the more recalcitrant root compartment.     

High nutrient availability leads to weaker top‐down control of stream periphyton: Compensatory feeding in Ancylus fluviatilis

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Forage quality in relation to long-term grazing history,current-year defoliation,and water resource

Forage nitrogen concentrations, nitrogen yields, and in vitro digestibilities were assessed in shortgrass steppe that had been ungrazed, lightly, or heavily grazed for 50 years. Caged plots were defoliated in amounts based upon removals observed in naturallygrazed reference plots or not defoliated. This was done in a year of average precipitation and with a supplemental water treatment to simulate a wet year. In general, current-year defoliation had positive effects, and longterm grazing and supplemental water had negative effects, on forage nitrogen concentrations and digestibilities. However, defoliation interacted with long-term grazing in determning forage nitrogen concentrations, and with grazing and with watering in determining digestibilities. Nitrogen concentration and digestibility increased with defoliation in lightly, but not in heavily, grazed treatments. The dilution effect of supplemental water an digestibilities through increased plant growth was offset by defoliation. The negative effects of long-term grazing on forage quality were small, equally or more than compensated for by defoliation in a year of average precipitation, but more pronounced in the simulated wet year. Nitrogen yields and digestible forage production were usually increased by defoliation, but this depended upon grazing and watering treatments. Increased nitrogen and digestible forage yields and concentrations in response to defoliation were greater than the biomass response in lightly grazed grassland. For both nitrogen and digestibility, yields were greater in grazed than ungrazed treatments in the year of average precipitation, but less in the simulated wet year. Optimizing quantity and year-to-year stability of nitrogen and digestible forage yield may best be achieved with light grazing rather than no or heavy grazing. Clipping was conducted in a manner closely resembling the natural pattern and intensity of defoliation by the cattle, and confirms the potential for a positive feedback of increased forage quality with defoliation observed in pot experiments. Long-term heavy grazing can diminish this response. Quantily (aboveground primary production, ANPP), quantity of quality (digestible and N yields), and quality (concentrations) do not necessarily respond similarly in interactions between current-year defoliation, long-term grazing history, and level of water resource.     

Effects of consumers and enrichment on abundance and diversity of benthic algae in a rocky intertidal community

Human alteration of nutrient cycling and the densities of important consumers have intensified the importance of understanding how nutrients and consumers influence the structure of ecological systems. We examined the effects of both grazing and nutrient enrichment on algal abundance and diversity in a high-intertidal limpet-macroalgal community on the South Island of New Zealand, a relatively nutrient-poor environment. We used a fully factorial design with three levels each of grazing (manipulations of limpet and snail densities) and nutrients (nutrient-diffusers attached to the rock). Top-down control by grazers appears to be the driving organizing mechanism for algal communities in this system, with strong negative effects of grazing on algal diversity and abundance across all levels of nutrient enrichment. However, in contrast to the conclusions drawn from the analysis of the whole algal community, there was an interactive effect of grazing and enrichment on foliose algae, an important component of the algal system. When herbivory was reduced to very low levels, enrichment generated increases in the abundance and biomass of foliose algae. As expected, top-down control was the primary determinant of algal community structure in this system, controlling abundance and diversity of macrophytes on the upper shore. Contrary to expectations, however, increased nutrients had no community-wide effects, although foliose algal abundance increases were greatest with high nutrients and reduced grazing. It seems likely that most of the corticated algal species have limited capacity to respond to nutrient pulses in this nutrient-poor environment.     

Sawdust Addition Reduces the Productivity of Nitrogen‐Enriched Mountain Grasslands

Anthropogenic nutrient enrichment of mountain grasslands has boosted grasses and fast‐growing unpalatable plants at the expense of slow‐growing species, resulting in a significant loss in biodiversity. A potential tool to reduce nutrient availability and aboveground productivity without destroying the perennial vegetation is carbon (C) addition. However, little is known about its suitability under severe climatic conditions. Here, we report the results of a 3‐year field study assessing the effects of sawdust addition on soil nutrients, aboveground productivity, and vegetational composition of 10 grazed and ungrazed mountain grasslands. Of particular interest was the effect of C addition on grasses and on the tall unpalatable weed Veratrum album. After 3 years, soil pH, ammonium, and plant‐available phosphorus were not altered by sawdust application, and nitrate concentrations were marginally higher in treatment plots. However, the biomass of grasses and forbs (without V. album) was 20–25% lower in sawdust‐amended plots, whereas the biomass of V. album was marginally higher. Sawdust addition reduced the cover of grasses but did not affect evenness, vegetation diversity, or plant species richness, although species richness generally increased with decreasing biomass at our sites. Our results suggest that sawdust addition is a potent tool to reduce within a relatively short time the aboveground productivity and grass cover in both grazed and ungrazed mountain grasslands as long as they are not dominated by tall unpalatable weeds. The technique has the advantage that it preserves the topsoil and the perennial soil seed bank.     

Food Limitation Affects Algivory and Grazer Performance for New Zealand Stream Macroinvertebrates

We studied herbivory and grazer performance (i.e., fitness correlates) for the hydrobiid snail Potamopyrgus antipodarum, the leptophlebiid mayfly Deleatidium spp., and the conoesucid caddisfly Pycnocentrodes aeris, common, co-occurring algivores in many New Zealand streams. Grazing effects and costs of coexisting differed among these taxa reared at ambient densities in different combinations in microcosms with algal food conditions (on clay tiles) characteristic of heavily grazed streams. The prostrate diatoms Staurosirella leptostauron, Cymbella novazealandia, and Achnanthidium minutissimum were the dominant algal species on pre- and post-grazed tiles. The relative abundance of erect physiognomic forms, dominated by Synedra ulna and Fragilaria vaucheriae, were 2–3× higher in ungrazed controls and in snail alone treatments than in other grazer treatments. The green filamentous algae Mougeotia sp. and Stigeoclonium lubricum, and the cyanophyte Merismopedia glauca were present only in ungrazed controls. Grazers significantly reduced algal community biomass in treatments by 26–52% relative to controls, except snails alone. Snails (15–30%) burrowed into surrounding sand substrates, dampening their grazing impact on tiles. Caddisflies were more effective than mayflies or snails at removing algae because of higher foraging rates, a larger body size, and an abrasive sand-grained case. Algal biomass reductions did not affect grazer growth. However, pre-pupation rates of caddisflies and emergence rates of subimago mayflies were significantly higher in caddisfly-alone and mayfly-alone treatments, respectively, than in combined-species treatments. These results imply that a limited periphytic food supply ( < 0.3 mg AFDM cm⁻²) even over a relatively brief period ( ≤ 16 d) may have population-scale consequences for co-existing P. aeris and Deleatidium spp.