Trophic Cascades in a Formerly Cod-Dominated Ecosystem Frank, Kenneth T; Petrie, Brian; Choi, Jae S ...
Science (American Association for the Advancement of Science),
06/2005, Letnik:
308, Številka:
5728
Journal Article
Recenzirano
Removal of top predators from ecosystems can result in cascading effects through the trophic levels below, completely restructuring the food web. Cascades have been observed in small-scale or simple ...food webs, but not in large, complex, open-ocean ecosystems. Using data spanning many decades from a once cod-dominated northwest Atlantic ecosystem, we demonstrate a trophic cascade in a large marine ecosystem. Several cod stocks in other geographic areas have also collapsed without recovery, suggesting the existence of trophic cascades in these systems.
Overfishing of large-bodied benthic fishes and their subsequent population collapses on the Scotian Shelf of Canada's east coast and elsewhere resulted in restructuring of entire food webs now ...dominated by planktivorous, forage fish species and macroinvertebrates. Despite the imposition of strict management measures in force since the early 1990s, the Scotian Shelf ecosystem has not reverted back to its former structure. Here we provide evidence of the transient nature of this ecosystem and its current return path towards benthic fish species domination. The prolonged duration of the altered food web, and its current recovery, was and is being governed by the oscillatory, runaway consumption dynamics of the forage fish complex. These erupting forage species, which reached biomass levels 900% greater than those prevalent during the pre-collapse years of large benthic predators, are now in decline, having outstripped their zooplankton food supply. This dampening, and the associated reduction in the intensity of predation, was accompanied by lagged increases in species abundances at both lower and higher trophic levels, first witnessed in zooplankton and then in large-bodied predators, all consistent with a return towards the earlier ecosystem structure. We conclude that the reversibility of perturbed ecosystems can occur and that this bodes well for other collapsed fisheries.
Virtually all studies reporting deepening with increasing size or age by fishes involve commercially harvested species. Studies of North Sea plaice in the early 1900s first documented this phenomenon ...(named Heincke’s law); it occurred at a time of intensive harvesting and rapid technological changes in fishing methods. The possibility that this deepening might be the result of harvesting has never been evaluated. Instead, age- or size-related deepening have been credited to interactions between density-dependent food resources and density-independent environmental factors. Recently, time-dependent depth variations have been ascribed to ocean warming. We use a model, initialized from observations of Atlantic cod (Gadus morhua) on the eastern Scotian Shelf, where an age-dependent deepening of ∼60 m was observed, to assess the effect of size- and depth-selective exploitation on fish distribution. Exploitation restricted to the upper 80 m can account for ∼72% of the observed deepening; by extending exploitation to 120 m, all of the deepening can be accounted for. In the absence of fishing, the model indicated no age-related deepening. Observations of depth distributions of older cod during a moratorium on fishing supported this prediction; however, younger cod exhibited low-amplitude deepening (10–15 m) suggestive of an ontogenetic response. The implications of these findings are manifold, particularly as they relate to hypotheses advanced to explain the ecological and evolutionary basis for ontogenetic deepening and to recent calls for the adoption of evidence of species deepening as a biotic indicator or “footprint” of warming seas.
It is widely believed that consumer control is a weak regulator of marine phytoplankton communities. It remains unclear, however, why this should be the case when marine consumers routinely regulate ...their prey at higher trophic levels. One possibility is that the weak consumer control of phytoplankton communities results from the inability of field researchers to effectively account for consumer–prey trophic relationships operating at the scale of the plankton. We explored this issue by reviewing studies of trophic control in marine plankton. Experimental studies indicate that size is a critical determinant of feeding relationships among plankton. In sharp contrast, of the 51 field studies reviewed, 78% did not distinguish among the sizes or species of phytoplankton and their consumers, but instead assumed a general bulk phytoplankton–zooplankton trophic connection. Such an approach neglects the possibility that several trophic connections may separate the smallest phytoplankton (0.2 μm) from the larger zooplankton (~ 1000 μm), a remarkable size differential exceeding that between a mouse (~10 cm) and an elephant (~2500 cm). The size‐based approach we propose integrates theory, experiments and field observations and has the potential to greatly enhance our understanding of the causes and consequences of recently documented restructuring of plankton communities.
The North Atlantic Fisheries Organization response to a precipitous decline of the Flemish Cap cod (Gadus morhua) stock during the 1990s was the imposition of an 11 year moratorium on directed ...fishing for cod; recovery followed. Over the three decades that encompassed the precollapse, collapse, and recovery stages, we found that the cod stock status was characterized by four traits: spawning stock biomass, maturity-at-age, weight-at-age, and recruitment. The temporal evolution of these traits was consistent with a density-dependent conceptual model suggesting that phenotypic plasticity was at play during the rebuilding of the stock. The temporal progression of the broader fish community paralleled that of cod, underlying its key ecosystem position. The same demographic variables defined the state of the adjacent Northern Cod stock that underwent a similar pattern of decline, an intermittent moratorium but only partial recovery. This partial recovery is possibly related, in part, to declines of prey species brought about by excessive harvesting after the cod collapse and an apparent collapse of capelin, a major dietary component.
Synchronous variations in the abundance of geographically distinct marine fish populations are known to occur across spatial scales on the order of 1,000 km and greater. The prevailing assumption is ...that this large-scale coherent variability is a response to coupled atmosphere–ocean dynamics, commonly represented by climate indexes, such as the Atlantic Multidecadal Oscillation and North Atlantic Oscillation. On the other hand, it has been suggested that exploitation might contribute to this coherent variability. This possibility has been generally ignored or dismissed on the grounds that exploitation is unlikely to operate synchronously at such large spatial scales. Our analysis of adult fishing mortality and spawning stock biomass of 22 North Atlantic cod (Gadus morhua) stocks revealed that both the temporal and spatial scales in fishing mortality and spawning stock biomass were equivalent to those of the climate drivers. From these results, we conclude that greater consideration must be given to the potential of exploitation as a driving force behind broad, coherent variability of heavily exploited fish species.
Globally, overfishing large-bodied groundfish populations has resulted in substantial increases in their prey populations. Where it has been examined, the effects of overfishing have cascaded down ...the food chain. In an intensively fished area on the western Scotian Shelf, Northwest Atlantic, the biomass of prey species increased exponentially (doubling time of 11 years) even though the aggregate biomass of their predators remained stable over 38 years. Concomitant reductions in herbivorous zooplankton and increases in phytoplankton were also evident. This anomalous trophic pattern led us to examine how declines in predator body size (approx. 60% in body mass since the early 1970s) and climatic regime influenced lower trophic levels. The increase in prey biomass was associated primarily with declines in predator body size and secondarily to an increase in stratification. Sea surface temperature and predator biomass had no influence. A regression model explained 65 per cent of prey biomass variability. Trait-mediated effects, namely a reduction in predator size, resulted in a weakening of top predation pressure. Increased stratification may have enhanced growing conditions for prey fish. Size-selective harvesting under changing climatic conditions initiated a trophic restructuring of the food chain, the effects of which may have influenced three trophic levels.
Marine fish populations of the global oceans and particularly large‐bodied, continental shelf‐dwelling groundfish species of the North Atlantic, such as cod (Gadus morhua), have been strongly ...perturbed by over‐fishing, frequently beyond levels that may have altered their capacity to recover. Age and size structure, spatial structure, reproductive potential, and other traits that convey fitness advantages are commonly lost when prolonged and excessive fishing pressure is exerted. Fisheries management protocols implemented to recover collapsed populations have been numerous and varied with all attempting to reduce or eliminate fishing pressure. Such measures, employed singly or in multiple combinations, typically involve quota reductions or fishing moratoria, area closures and other technical measures, as well as enhanced enforcement of fishing practices. A striking geographic pattern exists in the efficacy of such measures to regain lost population attributes and hence recovery. Some regional populations have recovered while others, despite management interventions lasting decades, notably, but not exclusively, those aimed at cod populations of the Northwest Atlantic, have yet to fully recover, an endpoint analogous to the conclusion of the famous nursery rhyme of Humpty Dumpty. Here we examine the dynamics of multiple collapsed populations exhibiting varying responses to recovery initiatives from the perspective of the Humpty Dumpty metaphor.
A key question in ecology is under which conditions ecosystem structure tends to be controlled by resource availability vs. consumer pressure. Several hypotheses derived from theory, experiments and ...observational field studies have been advanced, yet a unified explanation remains elusive. Here, we identify common predictors of trophic control in a synthetic analysis of 52 observational field studies conducted within marine ecosystems across the Northern Hemisphere and published between 1951 and 2014. Spatial regression analysis of 45 candidate variables revealed temperature to be the dominant predictor, with unimodal effects on trophic control operating both directly (r² = 0.32; P < 0.0001) and indirectly through influences on turnover rate and quality of primary production, biodiversity and omnivory. These findings indicate that temperature is an overarching determinant of the trophic dynamics of marine ecosystems, and that variation in ocean temperature will affect the trophic structure of marine ecosystems through both direct and indirect mechanisms.
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