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1. Coexistence of predators that share the same prey is common. This is still the case in size-structured predator communities where predators consume prey species of different sizes (interspecific prey responses) or consume different size classes of the same species of prey (intraspecific prey responses). A mechanism has recently been proposed to explain coexistence between predators that differ in size but share the same prey species, emergent facilitation, which is dependent on strong intraspecific responses from one or more prey species. Under emergent facilitation, predators can depend on each other for invasion, persistence or success in a size-structured prey community. Experimental evidence for intraspecific size-structured responses in prey populations remains rare, and further questions remain about direct interactions between predators that could prevent or limit any positive effects between predators [e.g. intraguild predation (IGP)]. Here, we provide a community-wide experiment on emergent facilitation including natural predators. We investigate both the direct interactions between two predators that differ in body size (fish vs. invertebrate predator), and the indirect interaction between them via their shared prey community (zooplankton). Our evidence supports the most likely expectation of interactions between differently sized predators that IGP rates are high, and interspecific interactions in the shared prey community dominate the response to predation (i.e. predator-mediated competition). The question of whether emergent facilitation occurs frequently in nature requires more empirical and theoretical attention, specifically to address the likelihood that its pre-conditions may co-occur with high rates of IGP.

Patterns in biomass production are determined by resource input (productivity) and trophic transfer efficiency. At fixed resource input, variation in consumer biomass production has been related to food quality, metabolic type and diversity among species. In contrast, intraspecific variation in individual body size because of ontogenetic development, which characterizes the overwhelming majority of taxa, has been largely neglected. Here we show experimentally in a long-term multigenerational study that reallocating constant resource input in a two-stage consumer system from an equal resource delivery to juveniles and adults to an adult-biased resource delivery is sufficient to cause more than a doubling of total consumer biomass. We discuss how such changes in consumer stage-specific resource allocation affect the likelihood for alternative stable states in harvested populations as a consequence of stage-specific overcompensation in consumer biomass and thereby the risk of catastrophic collapses in exploited populations.

Many marine ecosystems have undergone 'regime shifts', i.e. abrupt reorganizations across trophic levels. Establishing whether these constitute shifts between alternative stable states is of key importance for the prospects of ecosystem recovery and for management. We show how mechanisms underlying alternative stable states caused by predator-prey interactions can be revealed in field data, using analyses guided by theory on size-structured community dynamics. This is done by combining data on individual performance (such as growth and fecundity) with information on population size and prey availability. We use Atlantic cod (Gadus morhua) and their prey in the Baltic Sea as an example to discuss and distinguish two types of mechanisms, 'cultivation-depensation' and 'overcompensation', that can cause alternative stable states preventing the recovery of overexploited piscivorous fish populations. Importantly, the type of mechanism can be inferred already from changes in the predators' body growth in different life stages. Our approach can thus be readily applied to monitored stocks of piscivorous fish species, for which this information often can be assembled. Using this tool can help resolve the causes of catastrophic collapses in marine predatory-prey systems and guide fisheries managers on how to successfully restore collapsed piscivorous fish stocks.

Background

In recent decades, many attempts have been made to restore eutrophic lakes through biomanipulation. Reducing the populations of planktivorous and benthivorous fish (either directly or through stocking of piscivorous fish) may induce ecosystem changes that increase water transparency and decrease the risk of algal blooms and fish kills, at least in the short term. However, the generality of biomanipulation effects on water quality across lake types and geographical regions is not known. Therefore, we have undertaken a systematic review of such effects in eutrophic lakes in temperate regions throughout the world.

Methods

Searches for literature were made using online publication databases, search engines, specialist websites and bibliographies of literature reviews. Search terms were developed in English, Danish, Dutch and Swedish. Identified articles were screened for relevance using inclusion criteria set out in an a priori protocol. To reduce the risk of bias, we then critically appraised the combined evidence found on each biomanipulation. Data were extracted on outcomes such as Secchi depth and chlorophyll a concentration before, during and/or after manipulation, and on effect modifiers such as lake properties and amounts of fish removed or stocked.

Results

Our searches identified more than 14,500 articles. After screening for relevance, 233 of them remained. After exclusions based on critical appraisal, our evidence base included useful data on 128 biomanipulations in 123 lakes. Of these interventions, 85% had been made in Europe and 15% in North America. Meta-analysis showed that removal of planktivores and benthivores (with or without piscivore stocking) leads to increased Secchi depth and decreased chlorophyll a concentration during intervention and the first three years afterwards. Piscivore stocking alone has no significant effect. The response of chlorophyll a levels to biomanipulation is stronger in lakes where fish removal is intense, and in lakes which are small and/or have high pre-manipulation concentrations of total phosphorus.

Conclusions

Our review improves on previous reviews of biomanipulation in that we identified a large number of case studies from many parts of the world and used a consistent, repeatable process to screen them for relevance and susceptibility to bias. Our results indicate that removal of planktivorous and benthivorous fish is a useful means of improving water quality in eutrophic lakes. Biomanipulation tends to be particularly successful in relatively small lakes with short retention times and high phosphorus levels. More thorough fish removal increases the efficacy of biomanipulation. Nonetheless successes and failures have occurred across a wide range of conditions.

1. Humic lakes with a high external supply of DOC and low input of nutrients can often support a high biomass of metazoan zooplankton. In such lakes, autotrophic algae compete with bacteria for inorganic nutrients, but bacteria support mixotrophic growth. Consequently, planktonic communities are often dominated by mixotrophic flagellates, while obligate autotrophic phytoplankton occurs in low numbers for extended periods.

2. To test the importance of autotrophic phytoplankton and mixotrophic flagellates as food resources for metazoan grazers and, in turn, the feedback effects of grazers on basal food-web interactions, we conducted a long-term experiment where we simulated abiotic resource relationships of humic lakes (high DOC [glucose] and low P input). We examined the population dynamics of Daphnia galeata when inoculated in systems with autotrophic algae only, mixotrophic algae only and a mixture of autotrophic and mixotrophic algae, and how the systems changed after the inoculation of Daphnia. All combinations were run at high-and low-light conditions to analyse the effects of light on food quantity and quality.

3. Daphnia grew to high densities only when mixotrophs were present at high-light conditions and showed no or only weak growth at low-light conditions or with autotrophs as the only food source.

4. Autotrophic algae and bacteria showed a strong competition for nutrients. Autotrophic algae were released from competition for nutrients after Daphnia grazed on bacteria, which led to a probable change of the bacteria community to less edible but less competitive taxa. As a consequence, there was a mutualistic interaction between autotrophs and mixotrophs before Daphnia were introduced which turned into competition after Daphnia inoculation.

5. We suggest that mixotrophic flagellates can be a critical resource for cladocerans and thereby also have a cascading effect on higher trophic levels, and cladocerans, in turn, have important indirect effects on basal planktonic food webs; hence, both might affect whole lake ecosystems.

Food-dependent growth and size-dependent interactions form cornerstones in the dynamics of fish populations. Using two freshwater examples, we illustrate the importance of considering both these cornerstones for understanding system dynamics. Moreover, a proper understanding of the dynamics requires mechanistic linkages between individual-, population-, and community-level processes based on mass conservation principles. In one example, we further find that quantitative predictions of individual-level energy flows are essential for understanding the community dynamics. This mechanistic approach to understanding system dynamics is generally not reflected in fisheries models as an overview shows that only half of them incorporate food-dependent growth, and none fully observe the principles of mass conservation. As a marine example we examine patterns in the Baltic Sea system and show that no relationship between cod growth and sprat biomass is present related to the low size resolution in prey fish. Linking individual cod performance to its resource base is complicated by the many prey types cod uses over its life cycle. We conclude that an ecological perspective including size-and food-dependent processes is vital for ecosystem-based fisheries management making necessary a proper description of the interactive trophic structure as a result of mechanistic linkages between individual, population, and community processes.

In the study of mechanisms structuring fish communities, mixed competition-predation interactions where large predators feed on prey fish versus those in which small predators compete with prey fish for a shared prey have been the focus of substantial research. We used a long-term data set from a system inhabited by brown trout (Salmo trutta) (predator) and Arctic char (Salvelinus alpinus) (prey) to evaluate whether mixed interspecific interactions were present in this system as suggested in other studies focusing on this species pair. We found no evidence for a negative interspecific density dependence in individual performance in either Arctic char or brown trout. In contrast, a negative intraspecific density dependence was present, especially in Arctic char. Furthermore, large brown trout condition showed a positive response to encounter rate with Arctic char (related to the density of small Arctic char). The most parsimonious interaction module to explain the Arctic char - brown trout interaction patterns in the studied system does therefore not need to include interspecific competition. We suggest that size-structured mixed competition-predation interactions in different systems are realized as being either mainly structured through interspecific predation or by competition depending on species life history characteristics and environmental conditions.

Size-dependent interactions and habitat complexity have been identified as important factors affecting the persistence of intraguild predation (IGP) systems. Habitat complexity has been suggested to promote intraguild (IG) prey and intraguild predator coexistence through weakening trophic interactions particularly the predation link. Here, we experimentally investigate the effects of habitat complexity on coexistence and invasion success of differently sized IG-predators in a size-structured IGP system consisting of the IG-predator Poecilia reticulata and a resident Heterandria formosa IG-prey population. The experiments included medium-long and long-term invasion experiments, predator-prey experiments and competition experiments to elucidate the mechanisms underlying the effect of prey refuges. Habitat complexity did not promote the coexistence of IG-predator and IG-prey, although the predation link was substantially weakened. However, the presence of habitat structure affected the invasion success of large IG-predators negatively and the invasion success of small IG-predators positively. The effect of refuges on size-dependent invasion success could be related to a major decrease in the IG-predator's capture rate and a shift in the size distribution of IG-predator juveniles. In summary, habitat complexity had two main effects: (i) the predation link was diminished, resulting in a more competition driven system and (ii) the overall competitive abilities of the two species were equalized, but coexistence was not promoted. Our results suggest that in a size-structured IGP system, individual level mechanisms may gain in importance over species level mechanisms in the presence of habitat complexity.

Body size ( biomass) is the dominant determinant of population dynamical processes such as giving birth or dying in almost all species, with often drastically different behaviour occurring in different parts of the growth trajectory, while the latter is largely determined by food availability at the different life stages. This leads to the question under what conditions unstructured population models, formulated in terms of total population biomass, still do a fair job. To contribute to answering this question we first analyze the conditions under which a size-structured model collapses to a dynamically equivalent unstructured one in terms of total biomass. The only biologically meaningful case where this occurs is when body size does not affect any of the population dynamic processes, this is the case if and only if the mass-specific ingestion rate, the mass-specific biomass production and the mortality rate of the individuals are independent of size, a condition to which we refer as "ontogenetic symmetry". Intriguingly, under ontogenetic symmetry the equilibrium biomass-body size spectrum is proportional to 1/size, a form that has been conjectured for marine size spectra and subsequently has been used as prior assumption in theoretical papers dealing with the latter. As a next step we consider an archetypical class of models in which reproduction takes over from growth upon reaching an adult body size, in order to determine how quickly discrepancies from ontogenetic symmetry lead to relevant novel population dynamical phenomena. The phenomena considered are biomass overcompensation, when additional imposed mortality leads, rather unexpectedly, to an increase in the equilibrium biomass of either the juveniles or the adults (a phenomenon with potentially big consequences for predators of the species), and the occurrence of two types of size-structure driven oscillations, juvenile-driven cycles with separated extended cohorts, and adult-driven cycles in which periodically a front of relatively steeply decreasing frequencies moves up the size distribution. A small discrepancy from symmetry can already lead to biomass overcompensation; size-structure driven cycles only occur for somewhat larger discrepancies.

Most organisms show substantial changes in size or morphology after they become independent of their parents and have to find their own food. Furthermore, the rate at which these changes occur generally depends on the amount of food they ingest. In this book, André de Roos and Lennart Persson advance a synthetic and individual-based theory of the effects of this plastic ontogenetic development on the dynamics of populations and communities. De Roos and Persson show how the effects of ontogenetic development on ecological dynamics critically depend on the efficiency with which differently sized individuals convert food into new biomass. Differences in this efficiency--or ontogenetic asymmetry--lead to bottlenecks in and thus population regulation by either maturation or reproduction. De Roos and Persson investigate the community consequences of these bottlenecks for trophic configurations that vary in the number and type of interacting species and in the degree of ontogenetic niche shifts exhibited by their individuals. They also demonstrate how insights into the effects of maturation and reproduction limitation on community equilibrium carry over to the dynamics of size-structured populations and give rise to different types of cohort-driven cycles.