James F. Kitchell

Trophic cascades revealed in diverse ecosystems

New studies are documenting trophic cascades in theoretically unlikely systems such as tropical forests and the open ocean. Together with increasing evidence of cascades, there is a deepening understanding of the conditions that promote and inhibit the transmission of predatory effects. These conditions include the relative productivity of ecosystems, presence of refuges and the potential for compensation. However, trophic cascades are also altered by humans. Analyses of the extirpation of large animals reveal loss of cascades, and the potential of conservation to restore not only predator populations but also the ecosystem-level effects that ramify from their presence.

The trophic cascade in lakes

1. Cascading trophic interactions 2. Experimental lakes, manipulations and measurements 3. Statistical analysis of the ecosystem experiments 4. The fish populations 5. Fish behavioral and community responses to manipulation 6. Roles of fish predation: piscivory and planktivory 7. Dynamics of the phantom midge: implications for zooplankton 8. Zooplankton community dynamics 9. Effects of predators and food supply and diel vertical migration of Daphnia 10. Zooplankton biomass and body size 11. Phytoplankton community dynamics 12. Metalimnetic phytoplankton 13. Primary production and its interactions with nutrients and light transmission 14. Heterotrophic microbial processes 15. Annual fossil record of food-web manipulation 16. Simulation models of the trophic cascade: predictions and evaluations 17. Synthesis and new directions Index.

Whole-lake carbon-13 additions reveal terrestrial support of aquatic food webs

Ecosystems are supported by organic carbon from two distinct sources. Endogenous carbon is produced by photosynthesis within an ecosystem by autotrophic organisms. Exogenous carbon is produced elsewhere and transported into ecosystems. Consumers may use exogenous carbon with consequent influences on population dynamics, predator–prey relationships and ecosystem processes. For example, exogenous inputs provide resources that may enhance consumer abundance beyond levels supported by within-system primary production. Exogenous fluxes of organic carbon to ecosystems are often large, but this material is recalcitrant and difficult to assimilate, in contrast to endogenously produced organic matter, which is used more easily. Here we show, by the experimental manipulation of dissolved inorganic 13C in two lakes, that internal primary production is insufficient to support the food webs of these ecosystems. Additions of NaH13CO3 enriched the 13C content of dissolved inorganic carbon, particulate organic carbon, zooplankton and fish. Dynamics of 13C indicate that 40–55% of particulate organic carbon and 22–50% of zooplankton carbon are derived from terrestrial sources, showing that there is significant subsidy of these ecosystems by organic carbon produced outside their boundaries.

Early Warnings of Regime Shifts: A Whole-Ecosystem Experiment

High-frequency monitoring of manipulated and reference lakes enabled early detection of subsequent catastrophic regime shift. Catastrophic ecological regime shifts may be announced in advance by statistical early warning signals such as slowing return rates from perturbation and rising variance. The theoretical background for these indicators is rich, but real-world tests are rare, especially for whole ecosystems. We tested the hypothesis that these statistics would be early warning signals for an experimentally induced regime shift in an aquatic food web. We gradually added top predators to a lake over 3 years to destabilize its food web. An adjacent lake was monitored simultaneously as a reference ecosystem. Warning signals of a regime shift were evident in the manipulated lake during reorganization of the food web more than a year before the food web transition was complete, corroborating theory for leading indicators of ecological regime shifts.

TROPHIC CASCADES, NUTRIENTS, AND LAKE PRODUCTIVITY: WHOLE‐LAKE EXPERIMENTS

Responses of zooplankton, pelagic primary producers, planktonic bacteria, and CO2 exchange with the atmosphere were measured in four lakes with contrasting food webs under a range of nutrient enrichments during a seven-year period. Prior to enrichment, food webs were manipulated to create contrasts between piscivore dominance and planktivore dominance. Nutrient enrichments of inorganic nitrogen and phosphorus exhibited ratios of N:P > 17:1, by atoms, to maintain P limitation. An unmanipulated reference lake, Paul Lake, revealed baseline variability but showed no trends that could confound the interpretation of changes in the nearby manipulated lakes. Herbivorous zooplankton of West Long Lake (piscivorous fishes) were large-bodied Daphnia spp., in contrast to the small-bodied grazers that predominated in Peter Lake (planktivorous fishes). At comparable levels of nutrient enrichment, Peter Lakes areal chlorophyll and areal primary production rates exceeded those of West Long Lake by factors of approximatel...

Consumer Control of Lake ProductivityLarge-scale experimental manipulations reveal complex interactions among lake organisms

here is no common currency for ecological interactions. For example, the consumption of a small fish by a larger one entails all the following characteristics: behavioral interplay during pursuit and capture, an instantaneous reduction of the prey population, greater reproductive potential for the predator, a flux of organic energy, and a transfer of mineral nutrients such as phosphorus and nitrogen. Thus the same event is viewed differ-

ECOSYSTEM SUBSIDIES: TERRESTRIAL SUPPORT OF AQUATIC FOOD WEBS FROM 13C ADDITION TO CONTRASTING LAKES

Whole-lake additions of dissolved inorganic C-13 were used to measure allochthony (the terrestrial contribution of organic carbon to aquatic consumers) in two unproductive lakes (Paul and Peter Lakes in 2001), a nutrient-enriched lake (Peter Lake in 2002), and a dystrophic lake (Tuesday Lake in 2002). Three kinds of dynamic models were used to estimate allochthony: a process-rich, dual-isotope flow model based on mass balances of two carbon isotopes in 12 carbon pools; simple univariate time-series models driven by observed time courses of delta(13)CO(2); and multivariate autoregression models that combined information from time series of delta(13)C in several interacting carbon pools. All three models gave similar estimates of allochthony. In the three experiments without nutrient enrichment, flows of terrestrial carbon to dissolved and particulate organic carbon, zooplankton, Chaoborus, and fishes were substantial. For example, terrestrial sources accounted for more than half the carbon flow to juvenile and adult largemouth bass, pumpkinseed sunfish, golden shiners, brook sticklebacks, and fathead minnows in the unenriched experiments. Allochthony was highest in the dystrophic lake and lowest in the nutrient-enriched lake. Nutrient enrichment of Peter Lake decreased allochthony of zooplankton from 0.34-0.48 to 0-0.12, and of fishes from 0.51-0.80 to 0.25-0.55. These experiments show that lake ecosystem carbon cycles, including carbon flows to consumers, are heavily subsidized by organic carbon from the surrounding landscape.

Forage Fishes and Their Salmonid Predators in Lake Michigan

Abstract Alewife Alosa pseudoharengus and rainbow smelt Osmerus mordax dominate the planktivorous fish fauna of Lake Michigan and are now the primary food of lake trout Salvelinus namaycush and introduced salmonids. Their fluctuations in abundance have been a concern due to their effects on native species and their present role as forage species. Each has been implicated as an important factor in the local reduction or extinction of important native species. Mechanisms for these interactions include competition for food and predation on eggs and larvae. Bioenergetic modeling simulations of alewife consumption by stocked salmonids suggest that as much as 20 to 33% of the annual alewife production may be consumed in some years. Increasing stocking rates of salmonids in Lake Michigan yield a predator-prey system in which predator numbers become relatively independent of prey dynamics. This suggests possible declines in alewife production, changes in major forage available to predators, and perhaps destabiliz...

Prey selection naticid gastropods; experimental tests and application to the fossil record

Because predation by drilling gastropods is uniquely preservable in the fossil record, it represents important evidence for the study of coevolution. Previous studies of drilling gastropod predation have been largely descriptive and sometimes contradictory. We formulate and test a model of prey selection by naticid drilling gastropods. The model adequately predicts both prey species selection and prey size selection. Prey preferences parallel prey profitabilities, determined by calculating prey species-specific and predator size-specific cost-benefit functions. The model also specifically suggests the evolution of potential refugia from predation and the evolution of potential predatory attributes. Application of the model to several Miocene and Pliocene assemblages studied by Thomas (1976) corroborates the feasibility and utility of this approach in examining the evolutionary record of naticid predation, which extends from the Late Mesozoic. Apparent evolutionary stasis and convergent morphological trends among prey species may be consistent with continuous selection pressures against predation.

Representing Density Dependent Consequences of Life History Strategies in Aquatic Ecosystems: EcoSim II

ABSTRACT EcoSim II uses results from the Ecopath procedure for trophic mass-balance analysis to define biomass dynamics models for predicting temporal change in exploited ecosystems. Key populations can be represented in further detail by using delay-difference models to account for both biomass and numbers dynamics. A major problem revealed by linking the population and biomass dynamics models is in representation of population responses to changes in food supply; simple proportional growth and reproductive responses lead to unrealistic predictions of changes in mean body size with changes in fishing mortality. EcoSim II allows users to specify life history mechanisms to avoid such unrealistic predictions: animals may translate changes in feeding rate into changes in reproductive rather than growth rates, or they may translate changes in food availability into changes in foraging time that in turn affects predation risk. These options, along with model relationships for limits on prey availability caused by predation avoidance tactics, tend to cause strong compensatory responses in modeled populations. It is likely that such compensatory responses are responsible for our inability to find obvious correlations between interacting trophic components in fisheries time-series data. But Ecosim II does not just predict strong compensatory responses: it also suggests that large piscivores may be vulnerable to delayed recruitment collapses caused by increases in prey species that are in turn competitors/predators of juvenile piscivores.