Timothy E. Essington

Trophic Downgrading of Planet Earth

Until recently, large apex consumers were ubiquitous across the globe and had been for millions of years. The loss of these animals may be humankind’s most pervasive influence on nature. Although such losses are widely viewed as an ethical and aesthetic problem, recent research reveals extensive cascading effects of their disappearance in marine, terrestrial, and freshwater ecosystems worldwide. This empirical work supports long-standing theory about the role of top-down forcing in ecosystems but also highlights the unanticipated impacts of trophic cascades on processes as diverse as the dynamics of disease, wildfire, carbon sequestration, invasive species, and biogeochemical cycles. These findings emphasize the urgent need for interdisciplinary research to forecast the effects of trophic downgrading on process, function, and resilience in global ecosystems.

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...

The Role of Sharks and Longline Fisheries in a Pelagic Ecosystem of the Central Pacific

The increased exploitation of pelagic sharks by longline fisheries raised questions about changes in the food webs that include sharks as apex predators. We used a version of Ecopath/Ecosim models to evaluate changes in trophic interactions due to shark exploitation in the Central North Pacific. Fisheries targeted on blue sharks tend to produce compensatory responses that favor other shark species and billfishes, but they have only modest effects on the majority of food web components. Modest levels of intraguild predation (adult sharks that eat juvenile sharks) produce strong, nonlinear responses in shark populations. In general, analysis of the Central North Pacific model reveals that sharks are not keystone predators, but that increases in longline fisheries can have profound effects on the food webs that support sharks.

An ecosystem model of food web and fisheries interactions in the Baltic Sea

Because fisheries operate within a complex array of species interactions, scientists increasingly recommend multispecies approaches to fisheries management. We created a food web model for the Baltic Sea proper, using the Ecopath with Ecosim software, to evaluate interactions between fisheries and the food web from 1974 to 2000. The model was based largely on values generated by multispecies virtual population analysis (MSVPA). Ecosim outputs closely reproduced MSVPA biomass estimates and catch data for sprat (Sprattus sprattus), herring (Clupea harengus), and cod (Gadus morhua), but only after making adjustments to cod recruitment, to vulnerability to predation of specific species, and to foraging times. Among the necessary adjustments were divergent trophic relationships between cod and clupeids: cod exhibited top-down control on sprat biomass, but had little influence on herring. Fishing, the chief source of mortality for cod and herring, and cod reproduction, as driven by oceanographic conditions as well as unexplained variability, were also key structuring forces. The model generated many hypotheses about relationships between key biota in the Baltic Sea food web and may ultimately provide a basis for estimating community responses to management actions.

SHARKS AND TUNAS: FISHERIES IMPACTS ON PREDATORS WITH CONTRASTING LIFE HISTORIES

Large-scale pelagic fisheries exploit a diversity of apex predators with a wide range of life history strategies. Exploitation of species with different life history strategies has different population and food web consequences. We explored the changes in predation that result from exploitation of a common species with a slow growth and low fecundity life history strategy (blue shark, Prionace glauca) with those that result from exploitation of a common species with fast growth and high fecundity (yellowfin tuna, Thunnus alba- cares) in the central Pacific Ocean. Longline fisheries directed toward billfishes and tunas also capture blue sharks as incidental catch. Mortality rates of sharks had been relatively low prior to the recent surge in finning that has resulted in a substantial rise in mortality of adult and subadult sharks in the last decade. We estimated the magnitude of changes in predation by populations of yellowfin tuna and blue sharks in response to longline fisheries that involve shark finning. Bioenergetics models for sharks and tunas were coupled to simple population models that account for changes in size-structure in response to fishery- induced mortality regimes in order to estimate predation responses to changes in fishing intensities. Our analyses demonstrate that blue shark populations are very sensitive to low exploitation rates, while yellowfin tuna populations are extremely robust across a wide range of exploitation rates by longline fisheries. Although predation rates by yellowfin tuna are 4-5 times higher than by blue sharks, longline fisheries have substantially greater effects on shark predation than on yellowfin tuna predation at the food web scale. Expected food web responses will be strongest where the unexploited biomass of long-lived species is high and predation is relatively specialized compared with other apex predators. Our anal- yses suggest that active management to reduce finning mortality in sharks will play an important role toward minimizing the effects of longline fisheries on the food web structure of the pelagic Pacific Ocean.

Ecological indicators display reduced variation in North American catch share fisheries

A growing push to implement catch share fishery programs is based partly on the recognition that they may provide stronger incentives for ecological stewardship than conventional fisheries management. Using data on population status, quota compliance, discard rates, use of habitat-damaging gear, and landings for 15 catch share programs in North America, I tested the hypothesis that catch share systems lead to improved ecological stewardship and status of exploited populations. Impacts of catch share programs were measured through comparisons of fisheries with catch shares to fisheries without catch shares or by comparing fisheries before and after catch shares were implemented. The average levels of most indicators were unaffected by catch share implementation: only discard rate, which declined significantly in catch share fisheries, showed a significant response. However, catch share fisheries were distinguished by markedly reduced interannual variability in all indicators, being statistically significant for exploitation rate, landings, discard rate, and the ratio of catch to catch quotas. These impacts of catch shares were common between nations and ocean basins and were independent of the number of years that catch share programs had been in place. These findings suggest that for the indicators examined, the primary effect of catch shares was greater consistency over time. This enhanced consistency could be beneficial to fishery systems and might also be an indication of more effective management.

Eco-Label Conveys Reliable Information on Fish Stock Health to Seafood Consumers

Concerns over fishing impacts on marine populations and ecosystems have intensified the need to improve ocean management. One increasingly popular market-based instrument for ecological stewardship is the use of certification and eco-labeling programs to highlight sustainable fisheries with low environmental impacts. The Marine Stewardship Council (MSC) is the most prominent of these programs. Despite widespread discussions about the rigor of the MSC standards, no comprehensive analysis of the performance of MSC-certified fish stocks has yet been conducted. We compared status and abundance trends of 45 certified stocks with those of 179 uncertified stocks, finding that 74% of certified fisheries were above biomass levels that would produce maximum sustainable yield, compared with only 44% of uncertified fisheries. On average, the biomass of certified stocks increased by 46% over the past 10 years, whereas uncertified fisheries increased by just 9%. As part of the MSC process, fisheries initially go through a confidential pre-assessment process. When certified fisheries are compared with those that decline to pursue full certification after pre-assessment, certified stocks had much lower mean exploitation rates (67% of the rate producing maximum sustainable yield vs. 92% for those declining to pursue certification), allowing for more sustainable harvesting and in many cases biomass rebuilding. From a consumer’s point of view this means that MSC-certified seafood is 3–5 times less likely to be subject to harmful fishing than uncertified seafood. Thus, MSC-certification accurately identifies healthy fish stocks and conveys reliable information on stock status to seafood consumers.

Fishing amplifies forage fish population collapses

Significance Forage fish provide substantial benefits to both humans and ocean food webs, but these benefits may be in conflict unless there are effective policies governing human activities, such as fishing. Collapses of forage fish induce widespread ecological effects on dependent predators, but attributing collapses to fishing has been difficult because of natural fluctuations of these stocks. We implicate fishing in forage fish stock collapses by showing that high fishing rates are maintained when stock productivity is in rapid decline. As a consequence, the magnitude and frequency but not duration of stock collapses are far greater than expected from natural fluctuations. Risk-based management policies would provide substantial ecological benefits with little effect on fishery catches. Forage fish support the largest fisheries in the world but also play key roles in marine food webs by transferring energy from plankton to upper trophic-level predators, such as large fish, seabirds, and marine mammals. Fishing can, thereby, have far reaching consequences on marine food webs unless safeguards are in place to avoid depleting forage fish to dangerously low levels, where dependent predators are most vulnerable. However, disentangling the contributions of fishing vs. natural processes on population dynamics has been difficult because of the sensitivity of these stocks to environmental conditions. Here, we overcome this difficulty by collating population time series for forage fish populations that account for nearly two-thirds of global catch of forage fish to identify the fingerprint of fisheries on their population dynamics. Forage fish population collapses shared a set of common and unique characteristics: high fishing pressure for several years before collapse, a sharp drop in natural population productivity, and a lagged response to reduce fishing pressure. Lagged response to natural productivity declines can sharply amplify the magnitude of naturally occurring population fluctuations. Finally, we show that the magnitude and frequency of collapses are greater than expected from natural productivity characteristics and therefore, likely attributed to fishing. The durations of collapses, however, were not different from those expected based on natural productivity shifts. A risk-based management scheme that reduces fishing when populations become scarce would protect forage fish and their predators from collapse with little effect on long-term average catches.

The Effect of Whole-Lake Nutrient Enrichment on Mercury Concentration in Age-1 Yellow Perch

Abstract We evaluated the effect of whole-lake nutrient enrichment on Hg concentration in age-age-1 yellow perch Perca flavescens and assessed whether reduced Hg concentration in fish from enriched lakes could be attributed solely to enhanced fish growth (i.e., growth dilution). A survey of yellow perch in eighteight reference lakes and twotwo experimentally enriched lakes (P input = 2–6 mg · m−2 · d−1; N : P > 25 : 1 by atoms) indicated that yellow perch Hg concentration was highly correlated with lake pH and nutrient enrichment (R 2 = 0.87). Age-1 yellow perch were four to five times larger and had 50% lower Hg concentrations in enriched lakes than in reference lakes of an equivalent pH (reference lakes = 0.24 μg Hg/g wet mass; enriched lakes = 0.11 μg Hg/g). A mass balance model of Hg dynamics indicated that growth dilution could only account for 30–40% of the reduction in yellow perch Hg concentration, suggesting that lake enrichment produced effects on fish Hg concentration that were not explained by...

Nutrient cycling in lakes and streams : Insights from a comparative analysis

ABSTRACT Understanding of general ecosystem principles may be improved by comparing disparate ecosystems. We compared nutrient cycling in lakes and streams to evaluate whether contrasts in hydrologic properties lead to different controls and different rates of internal nutrient cycling. Our primary focus was nutrient cycling that results in increased productivity, so we quantified nutrient cycling by defining the recycling ratio (ρ) as the number of times a nutrient molecule is sequestered by producers before export. An analytic model of nutrient cycling predicted that in lakes ρ is governed by the processes that promote the mineralization and retard the sedimentation of particulate-bound nutrients, whereas in streams, ρ is governed by processes that promote the uptake and retard the export of dissolved nutrients. These differences were the consequence of contrast between lakes and streams in the mass-specific export rates (mass exported · standing stock-1· time-1) of dissolved and particulate nutrients. Although ρ is calculated from readily measured ecosystem variables, we found very few published data sets that provided the necessary data for a given ecosystem. We calculated and compared ρ in two well-studied P-limited ecosystems, Peter Lake and West Fork Walker Branch (WFWB). When ecosystems were scaled so that water residence time was equal between these two ecosystems, ρ was three orders of magnitude greater in WFWB. However, when we scaled by P residence time, ρ was nearly equal between these two ecosystems. This suggests broad similarities in ρ across ecosystem types when ecosystem boundaries are defined so that turnover times of limiting nutrients are the same.