Anders Persson

Effects of Enrichment on Simple Aquatic Food Webs

Simple models, based on Lotka‐Volterra types of interactions between predator and prey, predict that enrichment will have a destabilizing effect on populations and that equilibrium population densities will change at the top trophic level and every second level below. We experimentally tested these predictions in three aquatic food web configurations subjected to either high or low nutrient additions. The results were structured by viewing the systems as either food chains or webs and showed that trophic level biomass increased with enrichment, which contradicts food chain theory. However, within each trophic level, food web configuration affected the extent to which different functional groups responded to enrichment. By dividing trophic levels into functional groups, based on vulnerability to consumption, we were able to identify significant effects that were obscured when systems were viewed as food chains. The results support the prediction that invulnerable prey may stabilize trophic‐level dynamics by replacing other, more vulnerable prey. Furthermore, the vulnerable prey, such as Daphnia and edible algae, responded as predicted by the paradox of enrichment hypothesis; that is, variability in population density increased with enrichment. Hence, by describing ecosystems as a matrix of food web interactions, and by recognizing the interplay between interspecific competition and predation, a more complete description of the ecosystem function was obtained compared to when species were placed into distinct trophic levels.

FORAGING CAPACITY AND RESOURCE SYNCHRONIZATION IN AN ONTOGENETIC DIET SWITCHER, PIKEPERCH (STIZOSTEDION LUCIOPERCA)

Species undergoing ontogenetic diet shifts face a risk of resource competition that delays transitions between feeding stages. Such ontogenetic bottlenecks are common in piscivorous fish because competition with future prey may retard growth and prevent a size advantage. In pikeperch (Stizostedion lucioperca), year class strength of 0+ cohorts is highly variable and positively related to early onset of piscivory. To identify the causes of this pattern, we experimentally quantified size-dependent planktivorous and piscivorous foraging capacity and incorporated the data into a growth model. For any given prey type and size, foraging capacity described a hump-shaped relationship with predator size. Foraging capacity on daphnids peaked at a pikeperch length of 66 mm, suggesting a narrow scope of planktivory. The highest capacity in the piscivorous niche was reached at a predator-to-prey length ratio of 5, where the ratio was an integrated measure of predator size over several prey sizes. With the growth model, we derived size distributions of 0+ cohorts as functions of resource levels. Simulations revealed two major determinants for the year class strength of pikeperch. First, discontinuous availability of prey sizes counteracted switching to piscivory within the first growing season. This was accentuated by prey fish growth, which caused the planktivory and piscivory niches to separate over time and limited the time window when diet shift was possible. Second, the hump-shaped relationship between size and foraging capacity resulted in growth reduction when growing out of the planktivorous niche. Switching to piscivory in our model occurred along a perpendicular relationship between fish prey and zooplankton density. Zooplankton density determined whether pikeperch reached a size advantage over prey, and fish prey density affected whether the foraging return of piscivory was higher than planktivory. Individuals not reaching a size advantage over prey and failing to become piscivorous were stunted at a size when consumption balanced metabolic requirements. Piscivorous individuals, however, continued to grow fast throughout the season by feeding on the wave of the prey cohort. Our results highlight the importance for predators that shift diet to be synchronized with fluctuations in resource availability, such as the pulses of new cohorts of prey fish.

Effects of Fish Predation and Excretion on the Configuration of Aquatic Food Webs

The relative importance for algae of fish excretion and fish predation on zooplankton was studied in experimental pools. Even- (zoo- and phytoplankton) and odd-link systems (planktivorous fish, zoo- and phytoplankton) were established, in which the standing stock of primary producers was expected to follow the predictions from food chain theory sensu Oksanen. Two intermediate treatments (fish predation and nutrient regeneration by fish alone) were used to determine the relative power of fish in affecting the transition from an even-link system to an odd-link system, i.e. their relative power in the trophic cascade. Algal density and production were found to only be enhanced when both fish predation and fish excretion acted together. A reduction in the grazer biomass alone did not increase the algal biomass. Fish excretion alone increased the algal biomass somewhat, but not significantly so compared with the control. The results suggest that both fish predation on zooplankton and the regeneration of nutrients by fish may be necessary to create the difference in algal biomass between an even- and an odd-link system predicted by food chain models, at least in the absence of an external input of nutrients. In addition, the effects of increased nutrient regeneration due to use of non-zooplankton resources by the fish, nutrient maintenance in living biota, and decreased grazing pressure seem to reinforce the cascading effect of planktivorous fish on algae. Consequently, use of a fixed carrying capacity in simple predatorprey models may lead to loss of important information on the dynamics of aquatic food webs, preventing an adequate understanding of cascading trophic interactions from being achieved. (Less)

Consumption patterns, complexity and enrichment in aquatic food chains

The interactions between consumers and prey, and their impact on biomass distribution among trophic levels, are central issues in both empirical and theoretical ecology. In a long–term experiment, where all organisms, including the top predator, were allowed to respond to environmental conditions by reproduction, we tested predictions from ‘prey–dependent’and ‘ratio–dependent’models. Prey–dependent models made correct predictions only in the presence of strong interactors in simple food chains, but failed to predict patterns in more complex situations. Processes such as omnivory, consumer excretion, and unsuitable prey–size windows (invulnerable prey) increased the complexity and created patterns resembling ratio–dependent consumption. However, whereas the prey–dependent patterns were created by the mechanisms predicted by the model, ratio–dependent patterns were not, suggesting that they may be ‘right for the wrong reason’. We show here that despite the enormous complexity of ecosystems, it is possible to identify and disentangle mechanisms responsible for observed patterns in community structure, as well as in biomass development of organisms ranging in size from bacteria to fish.

DDT - FATE IN TROPICAL AND TEMPERATE REGIONS

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Effects of stream predator richness on the prey community and ecosystem attributes

It is important to understand the role that different predators can have to be able to predict how changes in the predator assemblage may affect the prey community and ecosystem attributes. We tested the effects of different stream predators on macroinvertebrates and ecosystem attributes, in terms of benthic algal biomass and accumulation of detritus, in artificial stream channels. Predator richness was manipulated from zero to three predators, using two fish and one crayfish species, while density was kept equal (n = 6) in all treatments with predators. Predators differed in their foraging strategies (benthic vs. drift feeding fish and omnivorous crayfish) but had overlapping food preferences. We found effects of both predator species richness and identity, but the direction of effects differed depending on the response variable. While there was no effect on macroinvertebrate biomass, diversity of predatory macroinvertebrates decreased with increasing predator species richness, which suggests complementarity between predators for this functional feeding group. Moreover, the accumulation of detritus was affected by both predator species richness and predator identity. Increasing predator species richness decreased detritus accumulation and presence of the benthic fish resulted in the lowest amounts of detritus. Predator identity (the benthic fish), but not predator species richness had a positive effect on benthic algal biomass. Furthermore, the results indicate indirect negative effects between the two ecosystem attributes, with a negative correlation between the amount of detritus and algal biomass. Hence, interactions between different predators directly affected stream community structure, while predator identity had the strongest impact on ecosystem attributes.

Synthesis of theoretical and empirical experiences from nutrient and cyprinid reductions in Lake Ringsjön

The reduction in external phosphorus load to Lake Ringsjön during the 1980s, did not result in improved water transparency during the following ten-year period. Furthermore, a fish-kill in the Eastern Basin of the lake, in addition to a cyprinid reduction programme (biomanipulation; 1988–1992), in contrast to theory, did not lead to any increase in zooplankton biomass or size. This absence of response in the pelagic food chain may have been attributed to the increase in abundance of YOY (0+) fish, following the fish reduction programme. Despite the lack of effect on zooplankton, there was a decrease in phytoplankton biomass, a change in species composition and an increase in water transparency following biomanipulation. In 1989, one year after the fish-kill in Eastern Basin, the Secchi depth (summer mean) increased from 60 cm to 110 cm. In the following years, water transparency increased further, despite an increase in phosphorus loading. An unexpected effect of the biomanipulation was an increase in benthic invertebrate and staging waterfowl abundances, which occurred 2–4 years after fish reduction. Hence, the response in the benthic community following biomanipulation was considerably stronger than in the pelagic community. A likely explanation is that reduction in abundance of the benthic feeding fish species bream (Abramis brama), strongly affected the benthic invertebrate fauna. In this paper, we present what we believe happened in Lake Ringsjön, and which processes are likely to have been important at various stages of the restoration process.

The effects of spatial food distribution and group size on foraging behaviour in a benthic fish

Animals foraging in heterogeneous environments benefit from information on local resource density because it allows allocation of foraging effort to rich patches. In foraging groups, this information may be obtained by individuals through sampling or by observing the foraging behaviour of group members. We studied the foraging behaviour of goldfish (Carassius auratus) groups feeding in pools on resources distributed in patches. First, we determined if goldfish use sampling information to distinguish between patches of different qualities, and if this allowed goldfish to benefit from a heterogeneous resource distribution. Then, we tested if group size affected the time dedicated to food searching and ultimately foraging success. The decision of goldfish to leave a patch was affected by whether or not they found food, indicating that goldfish use an assessment rule. Giving-up density was higher when resources were highly heterogeneous, but overall gain was not affected by resource distribution. We did not observe any foraging benefits of larger groups, which indicate that grouping behaviour was driven by risk dilution. In larger groups the proportion searching for food was lower, which suggests interactions among group members. We conclude that competition between group members affects individual investments in food searching by introducing the possibility for alternative strategies, such as scrounging or resource monopolisation.

LINKING PATCH-USE BEHAVIOR, RESOURCE DENSITY, AND GROWTH EXPECTATIONS IN FISH

Optimality theory rests on the assumptions that short-term foraging decisions are driven by variation in environmental quality, and that these decisions have important implications for long-term fitness. These assumptions, however, are rarely tested in a field setting. We linked behavioral foraging decisions in food patches with measures of environmental quality covering larger spatial (resource density) or temporal (growth parameters) scales. In 10 lakes, we measured the food density at which benthic fish give up foraging in experimental food patches (giving-up density, GUD), quantified the biomass of benthic invertebrates, and calculated the maximum individual size (L(infinity)) of bream (Abramis brama L.), a typical benthivore in these lakes. We found positive relationships between resource density and both GUD and L(infinity), and a positive relationship between L(infinity) and GUD. Prey characterized as vulnerable to predation contributed most to the relationships between resource density and either GUD or L(infinity). A path analysis showed that resource density and L(infinity) directly explained 54% and 28%, respectively, of the variation in GUD, whereas 86% of the variation in L(infinity) was explained by resource density, with mostly indirect contribution from GUD. We conclude that the short-term foraging behavior of benthivores matched our expectations based on optimality theory by being positively linked to variables on environmental quality operating at both a larger spatial scale and a longer temporal scale.

Effects of 17α-ethinylestradiol on individual life-history parameters and estimated population growth rates of the freshwater gastropods Radix balthica and Bithynia tentaculata.

Studies of aquatic environments exposed to 17α-ethinylestradiol (EE2) have demonstrated detrimental effects on fish communities. However, much less is known about effects on macro-invertebrates and especially how long-term exposure may affect critical life stages and ultimately population dynamics. We studied the effects of EE2 on relevant endpoints for population growth in two common freshwater gastropods, Radix balthica and Bithynia tentaculata, that differ in reproductive, foraging and anti-predator strategies and endocrine systems. We quantified critical life-history parameters (mortality, somatic growth rate, days to and size at first reproduction, egg production and hatching success) in a concentration–response, life cycle experiment. The two species responded to EE2 exposure in different ways, B. tentaculata showing a significantly lower somatic growth rate and R. balthica a higher somatic growth rate. However, the magnitudes of the effects were small and EE2 exposure did not have any significant effect on estimated population growth rates for the two snail species. The significant effects of EE2 on individual endpoints, but not on population growth rate for both species, emphasise the importance of evaluating higher level effects from long-term exposure studies.