Response of grazing impacts of an algivorous fish (Pseudogastromyzon myersi: Balitoridae) to seasonal disturbance in Hong Kong streams

1. Manipulative experiments were carried out in four Hong Kong streams (two shaded, two unshaded) to investigate the impact of grazing by an algivorous fish, Pseudogastromyzon myersi, on benthic algal biomass and assemblage composition. Experiments were conducted and repeated during both the dry and wet seasons to determine whether spate‐induced disturbance modified any grazing effect. Treatments comprised fish exclusion and inclusion via closed and open cages, with a no‐cage treatment used as a control for the cage effect. Treatments were maintained for 4 weeks in each experimental run. 2. Grazing by P. myersi reduced benthic algal biomass and the organic matter content of periphyton in open cages and the no‐cage treatment relative to closed cages. The similarity between open‐cage and no‐cage treatments was evidence that the overall difference among treatments was caused by limiting fish access to closed cages and not merely an artifact of caging. Grazing effects were broadly similar in all streams, but there was a significant statistical interaction between treatments and seasons. 3. Analysis of dry‐season data matched the overall trend in inter‐treatment differences, confirming the effects of grazing by P. myersi on algal biomass and periphyton organic matter. Significant differences in algal assemblage composition between closed‐cage and no‐cage treatments during the dry season reflected reductions in the abundance of erect, stalked diatoms (Gomphonema) and filamentous cyanobacteria (Homeothrix). Removal of these vulnerable overstorey algae by P. myersi resulted in greater abundance of understorey diatoms (Achnanthes and Cocconeis) in the no‐cage treatment in all streams during the dry season. The composition of algal assemblages in open cages was intermediate between the other two treatments. 4. Although fish densities were greater in all streams during the wet season, spate‐induced disturbance obscured grazing effects and there were no significant differences among treatments attributable to fish grazing. Seasonal variation in impacts of P. myersi grazing provides support for the harsh‐benign hypothesis, and confirms that biotic factors are less important controls of stream algal biomass and assemblage structure during periods (i.e. the wet season in Hong Kong) when abiotic disturbances are frequent or intense.     

Interactions among irradiance,nutrients, and herbivores constrain a stream algal community

Using stream-side, flow-through channels, I tested for the effects of nutrients (NU) (nitrogen plus phosphorus), irradiance (L), and snail grazing (G) on a benthic algal community in a small, forested stream. Grazed communities were-dominated by a chlorophyte (basal cells ofStigeoclonium) and a cyanophyte (Chamaesiphon investiens), whereas ungrazed communities were comprised almost entirely of diatoms, regardless of nutrient and light levels. Snails maintained low algal biomass in all grazed treatments, presumably by consuming increased algal production in treatments to which L and NU were increased. When nutrients were increased, cellular nutrient content increased under ambient conditions (shaded, grazed) and biomass and productivity increased when snails were removed and light was increased. Together, nutrients and light had positive effects and grazing had negative effects on biomass (chlorophylla, AFDM, algal biovolume) and chlorophyll-and areal-specific productivity in ANOVAs. However, in most cases, only means from treatments in which all three factors were manipulated (ungrazed, +NU&L treatments) were significantly different from controls; effects of single factors were generally undetectable. These results indicate that all three factors simultaneously limited algal biomass and productivity in this stream during the summer months. Additionally, the effects of these factors in combination were in some cases different from the effects of single factors. For example, light had slight negative effects on some biomass parameters when added at ambient snail densities and nutrient concentrations, but had strong positive effects in conjunction with nutrient addition and snail removal. This study demonstrates that algal biomass and productivity can be under multiple constraints by irradiance, nutrients, and herbivores and indicates the need to employ multifactor experiments to test for such interactive effects.     

Cascading effects of predatory fish on the composition of benthic algae in high‐altitude streams

Cascading effects of predators can affect ecosystem properties by changing plant biomass, distribution and assemblage composition. Using data from field surveys and whole‐stream experiments we tested the hypothesis that predatory trout change assemblage composition of benthic algae in high‐elevation streams mediated by grazer behavior. Field surveys revealed that the taxonomic composition of algal assemblages differed significantly between streams that contained trout and those that were fishless; but comparisons of palatable versus unpalatable algal taxa between fish and fishless streams were equivocal because of high natural variability. Therefore, we tested for a behavioral (non‐consumptive) trophic cascade experimentally by adding brook trout chemical cues to six naturally fishless streams for 25 days and compared responses of grazers and algae to six reference streams without fish cues added. Algal response variables included rates of change in the abundance of three physiognomic categories, from most palatable (attached erect and prostrate diatoms) to least palatable (non‐diatoms), as determined from food selectivity analyses of the most common grazers (mayflies and caddisflies). Fish cues did not affect the mean densities or changes in densities of total grazers or any individual grazer species. However, in streams where fish cues were added, rates of accrual of attached erect diatoms, which was the preferred algal type for the grazer most vulnerable to trout predation (Baetis), were higher and their densities increased significantly faster with increasing densities of this grazer species than in reference streams. Results of his experiment support the hypothesis that predator induced suppression of grazer foraging behavior, rather than cascading effects of top predators on grazer density, may contribute to variation in the composition of algal assemblages among streams by allowing proliferation of most palatable algal species.     

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.     

Increasing fish taxonomic and functional richness affects ecosystem properties of small headwater prairie streams

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Regulation of algal biomass in a small lowland stream: field experiments on the role of invertebrate grazing, phosphorus and irradiance

1. Field experiments were undertaken in a small Danish lowland stream to study the role of invertebrate grazing, phosphorus concentration and irradiance in the regulation of benthic algal biomass on stones. 2. Algal biomass was regulated by invertebrate grazing. The gastropod Ancylus fluviatilis prevented algal biomass build up in early spring at a density of about 900 ind. m^–2, and reduced algal biomass to very low levels during the algal growth period at a density of about 6000 ind. m^–2. Grazing pressure therefore might regulate the magnitude of peak algal biomass in Gelbæk stream, a finding in agreement with earlier field observations. 3. As phosphorus enrichment to around 152 μg l^–1 during the period when irradiance was not limiting did not result in any further increase in algal biomass, it can be concluded that algal growth in spring was not controlled by phosphorus limitation. 4. Algal biomass development differed significantly in a shaded and in a non-shaded reach of the stream. Only very low algal biomass accumulation was evident in the shaded reach, while in the non-shaded reach an algal biomass peak of about 800 mg chlorophyll m^–2 was evident after 6 weeks of colonization. The shaded reach represented the light condition in Gelbæk stream after leaf proliferation of the overhead canopy and high bank vegetation. The findings therefore suggest that irradiance available to the algal community after shading from riparian vegetation prevents further algal biomass increase and hence determines the timing of peak algal biomass in the stream. 5. The irradiance experiment also suggests that if not regulated by invertebrate grazing or shading by an overhead canopy in the summer, then the magnitude of peak biomass in Gelbæk stream will be regulated by self-shading in the algal community. 6. These field experiments support theories, derived from laboratory experiments and field studies, that regulation of algal biomass is a complex interaction of top-down and bottom-up mechanisms.     

Seasonal shifts in the importance of bottom–up and top–down factors on stream periphyton community structure

We examined the importance of temporal variability in top–down and bottom–up effects on the accumulation of stream periphyton, which are complex associations of autotrophic and heterotrophic microorganisms. Periphyton contributes to primary production and nutrient cycling and serves as a food resource for herbivores (grazers). Periphyton growth is often limited by the availability of nitrogen and phosphorus, and biomass can be controlled by grazers. In this study we experimentally manipulated nutrients and grazers simultaneously to determine the relative contribution of bottom–up and top–down controls on periphyton over time. We used nutrient diffusing substrates to regulate nutrient concentrations and an underwater electric field to exclude grazing insects in three sequential 16–17 day experiments from August to October in montane Colorado, USA. We measured algal biomass, periphyton organic mass, and algal community composition in each experiment and determined densities of streambed insect species, including grazers. Phosphorus was the primary limiting nutrient for algal biomass, but it did not influence periphyton organic mass across all experiments. Effects of nutrient additions on algal biomass and community composition decreased between August and October. Grazed substrates supported reduced periphyton biomass only in the first experiment, corresponding to high benthic abundances of a dominant mayfly grazer (Rhithrogena spp.). Grazed substrates in the first experiment also showed altered algal community composition with reduced diatom relative abundances, presumably in response to selective grazing. We showed that top–down grazing effects were strongest in late summer when grazers were abundant. The effects of phosphorus additions on algal biomass likely decreased over time because temperature became more limiting to growth than nutrients, and because reduced current velocity decreased nutrient uptake rates. These results suggest that investigators should proceed with caution when extending findings based on short‐term experiments. Furthermore, these results support the need for additional seasonal‐scale field research in stream ecology.     

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.     

Response of an algal assemblage to nutrient enrichment and shading in a Hawaiian stream

To investigate the effects of nitrate enrichment, phosphate enrichment, and light availability on benthic algae, nutrient-diffusing clay flowerpots were colonized with algae at two sites in a Hawaiian stream during spring and autumn 2002 using a randomized factorial design. The algal assemblage that developed under the experimental conditions was investigated by determining biomass (ash-free dry mass and chlorophyll a concentrations) and composition of the diatom assemblage. In situ pulse amplitude-modulated fluorometry was also used to model photosynthetic rate of the algal assemblage. Algal biomass and maximum photosynthetic rate were significantly higher at the unshaded site than at the shaded site. These parameters were higher at the unshaded site with either nitrate, or to a lesser degree, nitrate plus phosphate enrichment. Analysis of similarity of diatom assemblages showed significant differences between shaded and unshaded sites, as well as between spring and autumn experiments, but not between nutrient treatments. However, several individual species of diatoms responded significantly to nitrate enrichment. These results demonstrate that light availability (shaded vs. unshaded) is the primary limiting factor to algal growth in this stream, with nitrogen as a secondary limiting factor.     

SPECIES-SPECIFIC CHARACTERISTICS EXPLAIN THE PERSISTENCE OF STIGEOCLONIUM TENUE (CHLOROPHYTA) IN A WOODLAND STREAM1

The heterotrichous alga Stigeoclonium tenue Küetzing is dominant in many streams with high densities of herbivores. Previous in situ studies in Walker Branch (WB), a woodland stream in eastern Tennessee, indicated that dominance by Stigeoclonium basal cells was “grazer-dependent”; however, Stigeoclonium also appeared to have a lower biomass–specific productivity rate than other species that dominated when snails were experimentally removed. Here, an explicit test of the grazing dependence of Stigeoclonium was made with unialgal cultures established in the laboratory. Five different “assemblage types” were tested: 1 and 2) unialgal cultures of Stigeoclonium at low and high biomass, 3 and 4) a mixed assemblage of diatoms at low and high biomass, and 5) a natural stream community. Reduction in chlorophyll a after exposure to snail grazing was dependent on assemblage type (one-way ANOVA, P < 0.0001); low biomass Stigeoclonium tiles and tiles from the stream (on which basal cells of Stigeoclonium were dominant) were most grazer-resistant. In addition, Stigeoclonium had a lower biomass-specific productivity rate (measured as H¹⁴CO₃^? uptake) than a mixed assemblage of diatoms, regardless of biomass level, suggesting an underlying tradeoff between resistance to herbivory and competitive ability. Additional laboratory experiments were conducted to determine the response of Stigeoclonium to high (approx. 150 μmol quanta ·m^?2· s^?1)and low (approx. 25 μmol quanta · m^?2· s^?1) irradiance when nutrients were at 1) ambient WB concentrations and 2) increased 1000× ambient concentrations. There was a positive response of growth to increased irradiance only under high nutrient conditions. This suggests that observed reductions in the abundance of Stigeoclonium under high irradiance/low nutrient conditions that occur on a seasonal basis in WB can be explained in part by autecological resource requirements of this alga. We use these results to model the response of algal communities dominated by basal-regenerating species (e.g. Stigeoclonium) to gradients in herbivory and productivity. The results of our culture studies, combined with an overview of factors affecting communities dominated by grazer-resistant species, illustrate how both broad-scale (e.g. functional form) and species-specific studies can be combined to achieve an understanding of community dynamics.     

Stressor intensity determines antagonistic interactions between species invasion and multiple stressor effects on ecosystem functioning

Biological invasions, nutrient enrichment and ocean warming are known to threaten biodiversity and ecosystem functioning. The independent effects of these ecological stressors are well studied, however, we lack understanding of their cumulative effects, which may be additive, antagonistic or synergistic. For example, the impacts of biological invasions are often determined by environmental context, which suggests that the effects of invasive species may vary with other stressors such as pollution or climate change. This study examined the effects of an invasive seaweed (Sargassum muticum) on the structure and functioning of a synthetic macroalgal assemblage and tested explicitly whether these effects varied with nutrient enrichment and ocean warming. Overall, the presence of S. muticum increased assemblage productivity rates and warming altered algal assemblage structure, which was characterised by a decrease in kelp and an increase in ephemeral green algae. The effects of S. muticum on total algal biomass accumulation, however, varied with nutrient enrichment and warming, producing antagonistic cumulative effects on total algal biomass accumulation. These findings show that the nature of stressor interactions may vary with stressor intensity and among response variables, which leads to less predictable consequences for the structure and functioning of communities.     

Ecosystem Processes Performed by Unionid Mussels in Stream Mesocosms: Species Roles and Effects of Abundance

Unionid mussels are a guild of freshwater, sedentary filter-feeders experiencing a global decline in both species richness and abundance. To predict how these losses may impact stream ecosystems we need to quantify the effects of both overall mussel biomass and individual species on ecosystem processes. In this study we begin addressing these fundamental questions by comparing rates of ecosystem processes for two common mussel species, Amblema plicata and Actinonaias ligamentina, across a range of abundance levels and at two trophic states (low and high productivity) in stream mesocosms. At both low and high productivity, community respiration, water column ammonia, nitrate, and phosphorus concentrations, and algal clearance rates were all linearly related to overall mussel biomass. After removing the effects of biomass with ANCOVA, we found few differences between species. In a separate series of experiments, nutrient excretion (phosphorus, ammonia, and molar N:P) and biodeposition rates were only marginally different between species. For the species studied here, functional effects of unionids in streams were similar between species and linearly related to biomass, indicating the potential for strong effects when overall mussel biomass is high and hydrologic residence times are long.     

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

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Plant—herbivore interactions in streams near Mount St Helens

1. In four separate field experiments near Mount St Helens (Washington, U.S.A.) during 1986, the grazing effects of two large benthic herbivores, tadpoles of the tailed frog Ascaphus truei and larvae of the caddisfly Dicosmoecus gilvipes, were investigated using streamside channels and in-stream manipulations. In the experimental channels, abundances of periphyton and small benthic invertebrates declined significantly with increasing density of these larger herbivores. 2. In eleven small, high-gradient streams affected to varying degrees by the May 1980 eruption, in-stream platforms were used to reduce grazing by A, truei tadpoles on tile substrates. Single platforms erected in each tributary and compared to grazed controls revealed only minor grazing effects, and no significant differences among streams varying in disturbance intensity (and, consequently, tadpole density). However, results probably were confounded by high variability among streams in factors other than tadpole abundance. 3. Grazing effects were further examined in two unshaded streams with different tadpole densities, using five platforms per stream. In the stream with five tadpoles m^?2, grazing reduced periphyton biomass by 98% and chlorophyll a by 82%. In the stream lacking tadpoles, no significant grazing effects were revealed. Low algal abundance on both platforms and controls, and high invertebrate density in that stream (c. 30000m^?2) suggests that grazing by small, vagile invertebrates was approximately equivalent to that of tadpoles. 4. The influence of large benthic herbivores on algal and invertebrate communities in streams of Mount St Helens can be important, but reponses vary spatially in relation to stream disturbance history, local environmental factors, and herbivore distributional patterns and abundance.     

How do grazers affect periphyton heterogeneity in streams?

The effects of grazing by stream invertebrates on algal biomass and spatial heterogeneity were tested experimentally in flow-through microcosms with natural substrates (rocks). One experiment tested the effects of fixed densities of three species of grazers (the caddisfly Allomyia sp. and two mayflies, Epeorus deceptivus and Baetis bicaudatus) on periphyton. Baetis was tested with and without chemical cues from fish predators, which reduced grazer foraging activity to levels similar to the less mobile mayfly (Epeorus). Mean algal biomass (chlorophyll a; chl a) was reduced in grazer treatments compared to ungrazed controls, but there were no differences among grazer treatments. Algal heterogeneity (Morisita index) increased with grazer mobility, with the highest heterogeneity occurring in the Baetis-no fish treatment (most mobile grazer) and the lowest in the caddisfly treatment (most sedentary grazer). A second experiment used a three factorial design, and tested whether initial resource distribution (homogeneous vs. heterogeneous), Baetis density (high vs. low) and fish odor (present vs. absent) affected grazer impact on algal resources. Abundances of Baetis and chl a on individual rocks were recorded to explore the mechanisms responsible for the observed distributions of algae. Initial resource heterogeneity was maintained despite being subjected to grazing. Mean chl a was highest in controls, as in experiment I, and effects of Baetis on algal biomass increased with grazer density. There were no fish effects on algal biomass and no effects of grazer density or fish on algal heterogeneity. At the scale of individual rocks Baetis was unselective when food was homogeneously distributed, but chose high-food rocks when it was heterogeneously distributed. Results of these mechanistic experiments showed that Baetis can track resources at the scale of single rocks; and at moderate densities mobile grazers could potentially maintain periphyton distributions observed in natural streams.     

Herbivorous caddisflies,macroalgae, and epilithic microalgae: dynamic interactions in a stream grazing system

Summary 1. During the low-flow period (April–October) in sunlit pools of Big Sulphur Creek (northern coastal California), the attached algal community predictably changes from an assemblage dominated by lush, upright Cladophora glomerata filaments in spring and early summer to one dominated by epilithic diatoms and blue-green algae (together=microalgae) in late summer through early autumn. Previous studies in this stream indicated that grazing by the caddisflies Helicopsyche borealis and Gumaga nigricula maintain low algal biomass during the latter part of this period. We used a combination of in situ exclusion/enclosure experiments to examine (1) the separate and combined effects of these grazers on Cladophora and microalgal assemblages, and (2) food preferences, growth, and microdistribution patterns of grazers when offered these different algal foods. 2. Grazers exerted strong but divergent effects on algal assemblages. Selective grazing on Cladophora by G. nigricula greatly accelerated the transition from upright Cladophora to epilithic microalgae, whereas selective grazing on microalgae by H. borealis dramatically reduced biomass of these forms. Grazers were largely ineffective at reducing the non-preferred algal food source (i.e. Cladophora by H. borealis, microalgae by G. nigricula). In the case of each grazer, growth was highest on the preferred algal food. Together, the activity of these grazers produced a low-biomass assemblage dominated by microalgal cells. 3. Removal of the Cladophora overstory by G. nigricula resulted in a three-fold increase in the abundance of epilithic microalgae, the preferred food of H. borealis. Elimination of Cladophora by G. nigricula can increase food availability for H. borealis and, in so doing, can indirectly facilitate the growth of this grazer during food-limited conditions. However, microdistribution of G. nigricula shifts from high overlap with H. borealis in spring and early summer when Cladophora is abundant to low overlap in late summer after Cladophora has been eliminated. This may indicate intense competition between these species for limited epilithic algae, and a concomitant movement by G. nigricula to areas in the stream where food resources are more available.     

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.     

Experimental study of trophic cascade effect of silver carp (Hypophthalmichthys molitrixon) in a subtropical lake, Lake Donghu: on plankton community and underlying mechanisms of changes of crustacean community

An enclosure experiment was carried out to test trophic cascade effect of filter-feeding fish on the ecosystem: growth of crustacean zooplankton, and possible mechanism of changes of crustacean community structure. Four fish biomass levels were set as follows: 0, 116, 176 and 316 g m^-2, and lake water (containing ca. 190 g m^-2 of filter-feeding fishes) was comparatively monitored. Nutrient levels were high in all treatments during the experiment. Lowest algal biomass were measured in fishless treatment. Algal biomass decreased during days 21–56 as a function of fish biomass in treatments of low (LF), medium (MF) and high (HF) fish biomass. Crustaceans biomass decreased with increasing fish biomass. Small-bodied cladocerans, Moina micrura, Diaphanosoma brachyurum and Scapholeberis kingii survived when fish biomass was high whilst, large-bodied cladocerans Daphnia spp. and the cyclopoids Theromcyclops taihokuensis, T. brevifuratus, Mescyclops notius and Cyclops vicinus were abundant only in NF enclosures. Evasive calanoid Sinodiaptomus sarsi was significantly enhanced in LF, but decreased significantly with further increase of fish biomass. Demographic data indicated that M. micrura was well developed in all treatments. Our study indicates that algal biomass might be controlled by silver carp biomass in eutrophic environment. Changes of crustacean community are probably affected by the age of the first generation of species. Species with short generation time were dominant and species with long generation time survived less with high fish biomass. Evasive calanoids hardly developed in treatments with high fish biomass because of the (bottle neck) effect of nauplii. Species abundance were positively related to fish predation avoidance. Other than direct predation, zooplankton might also be suppressed by filter-feeding fish via competition.     

Effects of a top invertebrate predator (<Emphasis Type="Italic">Dytiscus alaskanus</Emphasis>; Coleoptera: Dytiscidae) on fishless pond ecosystems

We investigated the predatory effects of Dytiscus alaskanus, a large predaceous diving beetle, on the biomass, species composition and diversity of fishless pond communities. The effects were tested using presence and absence treatments of D. alaskanus in 24 mesocosms distributed among six ponds. We sampled phytoplankton, zooplankton and macroinvertebrates every two weeks for a six week period. Periphyton was sampled from the mesocosm walls on the final day. Total macroinvertebrate biomass decreased in the presence of dytiscids while species richness was not affected. Macroinvertebrate predators, snails and Gammarus lacustris decreased in the dytiscid treatments. Laboratory feeding experiments confirmed feeding preferences consistent with the mesocosm results. Periphyton biomass was six times greater in the dytiscid enclosures, concomitant with the decreased grazing by gastropods and other invertebrate primary consumers indicating a benthic trophic cascade. Top–down effects of dytiscids on other predatory invertebrates led to increased total zooplankton biomass, largely due to increased abundances of large and small cladocerans. Zooplankton species richness increased in the dytiscid enclosures. Inconsistent with trophic cascade theory, phytoplankton did not respond to top–down effects of D. alaskanus within the study period. Overall, the results show D. alaskanus predation caused trophic effects via two distinct food chains, a dytiscid–snail–periphyton trophic cascade, and a dytiscid–predatory macroinvertebrates–zooplankton partial trophic cascade.     

Changes in Stream Primary Producer Communities Resulting from Large-Scale Catastrophic Amphibian Declines: Can Small-Scale Experiments Predict Effects of Tadpole Loss?

Global declines of amphibian populations are well documented, yet effects of these declines on freshwater ecosystem structure and function are poorly understood. Here we examine responses of algal primary producers to tadpole extirpation over differing spatial and temporal scales. We experimentally excluded tadpoles from artificial substrata within localized areas (0.25 m²) of two streams. One stream had an intact community of frogs (frog stream), and the other had recently experienced a catastrophic decline (frogless stream), leaving virtually no tadpoles. In the frog stream, there were significantly greater levels of chlorophyll a (+111%, P = 0.009), ash-free dry mass (AFDM) (+163%, P = 0.02), inorganic sediments (+114%, P = 0.001), and higher mean algal cell biovolume in tadpole exclusion treatments than in the tadpole access treatments. Correspondingly, overall AFDM-specific net primary production (NPP) increased by 38% (P = 0.001) and chlorophyll a-specific NPP increased by 29% (P = 0.001) in tadpole access treatments compared to tadpole exclusion treatments. Areal-specific NPP did not differ between treatments. There were no significant differences in chlorophyll a, AFDM, inorganic sediments, algal cell biovolume, or biomass-specific NPP between treatments in the frogless stream. Fifteen months after our experiments, a massive amphibian decline associated with a fungal pathogen occurred in the frog stream, resulting in the extirpation of over 90% of tadpoles. This extirpation was followed by significant increases in levels of chlorophyll a (269%, P = 0.001), AFDM (+220%, P < 0.001), and inorganic sediments (+140%, P = 0.001). Reach-scale NPP increased from −1587 to −810 mg DO m⁻² d⁻¹. Additionally, algal community composition shifted from a dominance of small adnate diatoms (pre-decline) to a dominance of large upright algal species (post-decline). Our experimental results, combined with algal monitoring at the reach scale, indicate that over the course of our study catastrophic amphibian losses have significant effects on stream ecosystem structure and function. Ecosystem-level impacts of tadpole extirpations were more dramatic than results from our small-scale, short-term experiments, which predicted the direction of change in response variables but underestimated the magnitude. However, the long-term stream ecosystem responses remain unknown. Author Contributions: S.C., C.M.P., M.R.W., K.R.L., and S.K. designed the study, S.C., C.M.P., R.B., M.R.W., K.R.L., and A.D.H. performed research, S.C., C.M.P., R.J.B., M.R.W., and A.D.H. analyzed data. S.C., C.M.P., and R.J.B. wrote the paper.