Reconstructing historical marine ecosystems using food web models: Northern British Columbia from Pre-European contact to present

Mass-balance trophic models (Ecopath with Ecosim) are developed for the marine ecosystem of northern British Columbia (BC) for the historical periods 1750, 1900, 1950 and 2000 AD. Time series data are compiled for catch, fishing mortality and biomass using fisheries statistics and literature values. Using the assembled dataset, dynamics of the 1950-based simulations are fitted to agree with observations over 50 years to 2000 through the manipulation of trophic flow parameters and the addition of climate factors: a primary production anomaly and herring recruitment anomaly. The predicted climate anomalies reflect documented environmental series, most strongly sea surface temperature and the Pacific Decadal Oscillation index. The best-fit predator–prey interaction parameters indicate mixed trophic control of the ecosystem. Trophic flow parameters from the fitted 1950 model are transferred to the other historical periods assuming stationarity in density-dependent foraging tactics. The 1900 model exhibited an improved fit to data using this approach, which suggests that the pattern of trophic control may have remained constant over much of the last century. The 1950 model is driven forward 50 years using climate and historical fishing drivers. The resulting ecosystem is compared to the 2000 model, and the dynamics of these models are compared in a predictive forecast to 2050. The models suggest similar restoration trajectories after a hypothetical release from fishing.     

Integrative trophic network assessments of a lentic ecosystem by key ecological approaches of water chemistry, trophic guilds, and ecosystem health assessments along with an ECOPATH model

The objective of this study was to describe the trophic structure and energy flow in a lentic ecosystem in South Korea. Physicochemical water conditions were evaluated along with the reservoir ecosystem health using a multimetric IBI model. Nutrient analyses of the reservoir showed a nutrient rich and hypereutrophic system. Guild analysis revealed that tolerant and omnivorous species dominated the ecosystem. Tolerant fish, as a proportion of the number of individuals, were associated (R² > 0.90, p < 0.01) with TN and TP, the key indicators of trophic state in lentic ecosystems. The mean Reservoir Ecosystem Health Assessment (REHA) score was 19.3 during the study, which was judged as in ‘fair to poor’ condition. A trophic analysis of the reservoir estimated by the ECOPATH model shows that most activity in terms of energy flow occurred in the lower part of the trophic web, where there was intensive use of primary producers as a food source. Consequently, of the 10 consumer groups, nine fell within trophic levels <2.8. Trophic levels (TL) estimated from the weighted average of prey trophic levels varied from 1.0 for phytoplankton, macrophytes, and detritus to 3.25 for the top predator, Pseudobagrus fulvidraco. Our integrated approach to trophic network analysis may provide a key tool for determining the effects of nutrient influx on energy flow pathways in lentic ecosystems.     

Which forcing factors fit? Using ecosystem models to investigate the relative influence of fishing and changes in primary productivity on the dynamics of marine ecosystems

Fishing mortality and primary production (or proxy for) were used to drive the dynamics of fish assemblages in 9 trophodynamic models of contrasting marine ecosystems. Historical trends in abundance were reconstructed by fitting model predictions to observations from stock assessments and fisheries independent survey data. The model fitting exercise derives values for otherwise unknown parameters that specify the relative strength of trophic interactions and, in some instances, a time series anomaly for changes in primary production. We measured how much better or worse were model predictions when bottom-up forcing by primary production were added to top-down forcing by fishing. Searching for cross system patterns, the relative contribution of fishing and changes in primary production, mediated through trophic interactions, are evaluated for the ecosystems as a whole and for selected similar species in different ecosystems. The analysis provides a simple qualitative way to explain which forcing factors have most influence on modeled dynamics. Both fishing and primary production forcing were required to obtain the best model fits to data. Fishing effects more strongly influenced 6 of 9 of the ecosystems, but primary production was more often found to be the main factor influencing the selected pelagic and demersal fish stock trends. Examination of sensitivity to ecological and model parameters suggests that the results are the product of complex food-web interactions rather than simple deterministic responses of the models.     

Decadal changes in a NW Mediterranean Sea food web in relation to fishing exploitation

We analysed changes in the ecological roles of species, trophic structure and ecosystem functioning using four standardized mass-balance models of the South Catalan Sea (North-western Mediterranean). Models represented the ecosystem during the late 1970s, mid 1990s, early 2000s, and a simulated no-fishing scenario. The underlying hypothesis was that ecosystem models should quantitatively capture the increasing exploitation in the ecosystem from the 1970s to 2000s, as well as differences between the exploited and non-exploited scenarios. Biomass showed a general decrease, while there was an increase in biomass at lower trophic levels (TL) from the 1970s to 2000s. The efficiency of energy transfer (TE) from lower to higher TLs significantly increased with time. The ecosystem during the 1990s showed higher biomass and flows than during the 1970s and 2000s due to an increase in small pelagic fish biomass (especially sardines). Exploited food webs also showed similarities in terms of general structure and functioning due to high intensity of fishing already in the 1970s. This intensity was highlighted with low trophic levels in the catch, high consumption of production by fisheries, medium to high primary production required to sustain the catches and high losses in secondary production due to fishing. Significant differences on ecosystem structure and functioning were highlighted between the exploited and no-fishing scenarios. Biomass of higher TLs increased under the no-fishing scenario and the mean trophic level of the community and the fish/invertebrate biomass ratios were substantially lower in exploited food webs. The efficiency of energy transfer (TE) from lower to higher TLs was lower under the no-fishing scenario, and it showed a continuous decrease with increasing TL. Marine mammals, large hake, anglerfish and large pelagic fish were identified as keystone species of the ecosystem when there was no fishing, while their ecological importance notably decreased under the exploited periods. On the contrary, the importance of small-sized organisms such as benthic invertebrates and small pelagic fish was higher in exploited food webs.     

Field evidence for pervasive indirect effects of fishing on prey foraging behavior

The indirect, ecosystem-level consequences of ocean fishing, and particularly the mechanisms driving them, are poorly understood. Most studies focus on density-mediated trophic cascades, where removal of predators alternately causes increases and decreases in abundances of lower trophic levels. However, cascades could also be driven by where and when prey forage rather than solely by prey abundance. Over a large gradient of fishing intensity in the central Pacific's remote northern Line Islands, including a nearly pristine, baseline coral reef system, we found that changes in predation risk elicit strong behavioral responses in foraging patterns across multiple prey fish species. These responses were observed as a function of both short-term ("acute") risk and longer-term ("chronic") risk, as well as when prey were exposed to model predators to isolate the effect of perceived predation risk from other potentially confounding factors. Compared to numerical prey responses, antipredator behavioral responses such as these can potentially have far greater net impacts (by occurring over entire assemblages) and operate over shorter temporal scales (with potentially instantaneous response times) in transmitting top-down effects. A rich body of literature exists on both the direct effects of human removal of predators from ecosystems and predators' effects on prey behavior. Our results draw together these lines of research and provide the first empirical evidence that large-scale human removal of predators from a natural ecosystem indirectly alters prey behavior. These behavioral changes may, in turn, drive previously unsuspected alterations in reef food webs.     

Asymmetry in community regulation: effects of predators and productivity

Eutrophication and predator additions and extinctions are occurring in ecosystems worldwide. Although theory predicts that both will strongly alter the distribution of biomass in whole communities, empirical evidence has not been consolidated to quantitatively determine whether these theoretical predictions are generally borne out in real ecosystems. Here we analyze data from two types of trophic cascade studies, predator removals in factorial combination with fertilization and observed productivity gradients, to assess the role of top-down and bottom-up forces in structuring multi-trophic communities and compare results from these analyses to those from an extensive database of trophic cascade studies. We find that herbivore biomass declines and plant biomass increases in the presence of predators, regardless of system productivity. In contrast, while plants are increased by fertilization, this effect does not significantly increase herbivores in either the presence or absence of predators. These patterns are consistent among marine, freshwater, and terrestrial ecosystems and are largely independent of study size and duration. Thus, top-down effects of predation are transferred through more trophic levels than are bottom-up effects of eutrophication, showing strong asymmetry in the direction of control of biomass distribution in communities.     

Interacting trophic forcing and the population dynamics of herring

Small pelagic fish occupy a central position in marine ecosystems worldwide, largely by determining the energy transfer from lower trophic levels to predators at the top of the food web, including humans. Population dynamics of small pelagic fish may therefore be regulated neither strictly bottom-up nor top-down, but rather through multiple external and internal drivers. While in many studies single drivers have been identified, potential synergies of multiple factors, as well as their relative importance in regulating population dynamics of small pelagic fish, is a largely unresolved issue. Using a statistical, age-structured modeling approach, we demonstrate the relative importance and influence of bottom-up (e.g., climate, zooplankton availability) and top-down (i.e., fishing and predation) factors on the population dynamics of Bothnian Sea herring (Clupea harengus) throughout its life cycle. Our results indicate significant bottom-up effects of zooplankton and interspecific competition from sprat (Sprattus sprattus), particularly on younger age classes of herring. Although top-down forcing through fishing and predation by grey seals (Halichoerus grypus) and Atlantic cod (Gadus morhua) also was evident, these factors were less important than resource availability and interspecific competition. Understanding key ecological processes and interactions is fundamental to ecosystem-based management practices necessary to promote sustainable exploitation of small pelagic fish.     

Invasive species impacts on ecosystem structure and function: A comparison of the Bay of Quinte, Canada, and Oneida Lake, USA, before and after zebra mussel invasion

As invasion rates of exotic species increase, an ecosystem level understanding of their impacts is imperative for predicting future spread and consequences. We have previously shown that network analyses are powerful tools for understanding the effects of exotic species perturbation on ecosystems. We now use the network analysis approach to compare how the same perturbation affects another ecosystem of similar trophic status. We compared food web characteristics of the Bay of Quinte, Lake Ontario (Canada), to previous research on Oneida Lake, New York (USA) before and after zebra mussel (Dreissena polymorpha) invasion. We used ecological network analysis (ENA) to rigorously quantify ecosystem function through an analysis of direct and indirect food web transfers. We used a social network analysis method, cohesion analysis (CA), to assess ecosystem structure by organizing food web members into subgroups of strongly interacting predators and prey. Together, ENA and CA allowed us to understand how food web structure and function respond simultaneously to perturbation. In general, zebra mussel effects on the Bay of Quinte, when compared to Oneida Lake, were similar in direction, but greater in magnitude. Both systems underwent functional changes involving focused flow through a small number of taxa and increased use of benthic sources of production; additionally, both systems structurally changed with subgroup membership changing considerably (33% in Oneida Lake) or being disrupted entirely (in the Bay of Quinte). However, the response of total ecosystem activity (as measured by carbon flow) differed between both systems, with increasing activity in the Bay of Quinte, and decreasing activity in Oneida Lake. Thus, these analyses revealed parallel effects of zebra mussel invasion in ecosystems of similar trophic status, yet they also suggested that important differences may exist. As exotic species continue to disrupt the structure and function of our native ecosystems, food web network analyses will be useful for understanding their far-reaching effects.     

Suppression of soybean aphid by generalist predators results in a trophic cascade in soybeans.

Top-down regulation of herbivores in terrestrial ecosystems is pervasive and can lead to trophic cascades that release plants from herbivory. Due to their relatively simplified food webs, agroecosystems may be particularly prone to trophic cascades, a rationale that underlies biological control. However, theoretical and empirical studies show that, within multiple enemy assemblages, intraguild predation (IGP) may lead to a disruption of top-down control by predators. We conducted a factorial field study to test the separate and combined effects of predators and parasitoids in a system with asymmetric IGP. Specifically we combined ambient levels of generalist predators (mainly Coccinellidae) of the soybean aphid, Aphis glycines Matsumura, with controlled releases of the native parasitoid Lysiphlebus testaceipes (Cresson) and measured their impact on aphid population growth and soybean biomass and yield. We found that generalist predators provided strong, season-long aphid suppression, which resulted in a trophic cascade that doubled soybean biomass and yield. However, contrary to our expectations, L. testaceipes provided minor aphid suppression and only when predators were excluded, which resulted in nonadditive effects when both groups were combined. We found direct and indirect evidence of IGP, but because percentage parasitism did not differ between predator exclusion and ambient predator treatments, we concluded that IGP did not disrupt parasitism during this study. Our results support theoretical predictions that intraguild predators which also provide strong herbivore suppression do not disrupt top-down control of herbivores.     

Two-way coupling versus one-way forcing of plankton and fish models to predict ecosystem changes in the Benguela

‘End-to-end’ models have been adopted in an attempt to capture more of the processes that influence the ecology of marine ecosystems and to make system wide predictions of the effects of fishing and climate change. Here, we develop an end-to-end model by coupling existing models that describe the dynamics of low (ROMS–N₂P₂Z₂D₂) and high trophic levels (OSMOSE). ROMS–N₂P₂Z₂D₂ is a biogeochemical model representing phytoplankton and zooplankton seasonal dynamics forced by hydrodynamics in the Benguela upwelling ecosystem. OSMOSE is an individual-based model representing the dynamics of several species of fish, linked through opportunistic and size-based trophic interactions. The models are coupled through a two-way size-based predation process. Plankton provides prey for fish, and the effects of predation by fish on the plankton are described by a plankton mortality term that is variable in space and time. Using the end-to-end model, we compare the effects of two-way coupling versus one-way forcing of the fish model with the plankton biomass field. The fish-induced mortality on plankton is temporally variable, in part explained by seasonal changes in fish biomass. Inclusion of two-way feedback affects the seasonal dynamics of plankton groups and usually reduces the amplitude of variation in abundance (top-down effect). Forcing and coupling lead to different predicted food web structures owing to changes in the dominant food chain which is supported by plankton (bottom-up effect). Our comparisons of one-way forcing and two-way coupling show how feedbacks may affect abundance, food web structure and food web function and emphasise the need to critically examine the consequences of different model architectures when seeking to predict the effects of fishing and climate change.     

Effects of near‐future ocean acidification,fishing, and marine protection on a temperate coastal ecosystem

Understanding ecosystem responses to global and local anthropogenic impacts is paramount to predicting future ecosystem states. We used an ecosystem modeling approach to investigate the independent and cumulative effects of fishing, marine protection, and ocean acidification on a coastal ecosystem. To quantify the effects of ocean acidification at the ecosystem level, we used information from the peer‐reviewed literature on the effects of ocean acidification. Using an Ecopath with Ecosim ecosystem model for the Wellington south coast, including the Taputeranga Marine Reserve (MR), New Zealand, we predicted ecosystem responses under 4 scenarios: ocean acidification + fishing; ocean acidification + MR (no fishing); no ocean acidification + fishing; no ocean acidification + MR for the year 2050. Fishing had a larger effect on trophic group biomasses and trophic structure than ocean acidification, whereas the effects of ocean acidification were only large in the absence of fishing. Mortality by fishing had large, negative effects on trophic group biomasses. These effects were similar regardless of the presence of ocean acidification. Ocean acidification was predicted to indirectly benefit certain species in the MR scenario. This was because lobster (Jasus edwardsii) only recovered to 58% of the MR biomass in the ocean acidification + MR scenario, a situation that benefited the trophic groups lobsters prey on. Most trophic groups responded antagonistically to the interactive effects of ocean acidification and marine protection (46%; reduced response); however, many groups responded synergistically (33%; amplified response). Conservation and fisheries management strategies need to account for the reduced recovery potential of some exploited species under ocean acidification, nonadditive interactions of multiple factors, and indirect responses of species to ocean acidification caused by declines in calcareous predators.     

Predator diversity and identity drive interaction strength and trophic cascades in a food web

Declining predator diversity may drastically affect the biomass and productivity of herbivores and plants. Understanding how changes in predator diversity can propagate through food webs to alter ecosystem function is one of the most challenging ecological research topics today. We studied the effects of predator removal in a simple natural food web in the Sierra Nevada mountains of California (USA). By excluding the predators of the third trophic level of a food web in a full-factorial design, we monitored cascading effects of varying predator diversity and composition on the herbivorous beetle Chrysomela aeneicollis and the willow Salix orestera, which compose the first and second trophic levels of the food web. Decreasing predator diversity increased herbivore biomass and survivorship, and consequently increased the amount of plant biomass consumed via a trophic cascade. Despite this simple linear mean effect of diversity on the strength of the trophic cascade, we found additivity, compensation, and interference in the effects of multiple predators on herbivores and plants. Herbivore survivorship and predator-prey interaction strengths varied with predator diversity, predator identity, and the identity of coexisting predators. Additive effects of predators on herbivores and plants may have been driven by temporal niche separation, whereas compensatory effects and interference occurred among predators with a similar phenology. Together, these results suggest that while the general trends of diversity effects may appear linear and additive, other information about species identity was required to predict the effects of removing individual predators. In a community that is not temporally well-mixed, predator traits such as phenology may help predict impacts of species loss on other species. Information about predator natural history and food web structure may help explain variation in predator diversity effects on trophic cascades and ecosystem function.     

Long-term trends in the trophic structure of the North Sea fish community: evidence from stable-isotope analysis, size-spectra and community metrics

Fishing has wide-ranging impacts on marine ecosystems. One of the most pervasive signs of intensive fishing is "fishing down the food web", with landings increasingly dominated by smaller species from lower trophic levels. Decreases in the trophic level of landings are assumed to reflect those in fish communities, because size-selective mortality causes decreases in the relative abundance of larger species and in mean body size within species. However, existing analyses of fishing impacts on the trophic level of fish communities have focused on the role of changes in species composition rather than size composition. This will provide a biased assessment of the magnitude of fishing impacts, because fishes feed at different trophic levels as they grow. Here, we combine body size versus trophic level relationships for North Sea fishes (trophic level assessed using nitrogen stable-isotope analysis) with species-size-abundance data from two time-series of trawl-survey data (whole North Sea 1982-2000, central and northern North Sea 1925-1996) to predict long-term trends in the trophic structure of the North Sea fish community. Analyses of the 1982-2000 time-series showed that there was a slow but progressive decline in the trophic level of the demersal community, while there was no trend in the trophic level of the combined pelagic and demersal community. Analyses of the longer time-series suggested that there was no trend in the trophic level of the demersal community. We related temporal changes in trophic level to temporal changes in the slopes of normalised biomass size-spectra (which theoretically represent the trophic structure of the community), mean log₂ body mass and mean log₂ maximum body mass. While the size-based metrics of community structure showed long-term trends that were consistent with the effects of increased fishery exploitation, these trends were only correlated with trophic level for the demersal community. Our analysis suggests that the effects of fishing on the trophic structure of fish communities can be much more complex than previously assumed. This is a consequence of sampled communities not reflecting all the pathways of energy transfer in a marine ecosystem and of the absence of historical data on temporal and spatial changes in the trophic level of individuals. For the North Sea fish community, changes in size structure due to the differential effects of fishing on species and populations with different life histories are a stronger and more universal indicator of fishing effects than changes in mean trophic level.     

Exploring fisheries strategies for the western English Channel using an ecosystem model

An ecosystem model of the western English Channel ecosystem in 1994 was used to explore the effects of the use of a fishing policy optimization routine on profits, number of jobs and ecosystem structure. The optimization for single objective led to the specialization of the fishing fleet, with some fleet types being almost excluded. The profits and mainly the job optimizations led to big changes in the ecosystem structure, with loss of diversity, but the overall biomass of all vertebrate groups represented in the model increased considerably. For the objective focusing on ecosystem structure, there was an increase in biodiversity, with many long-lived groups predicted to increase, although the overall vertebrate biomass suffered just a small increase. An “ideal” mixed policy configuration was found when slightly greater weight was given to ecosystem structure than was given to profits and jobs. This scenario led to an overall reduction in effort but also to increased profits and biodiversity, while keeping the number of jobs at the same level as the baseline estimates. The results of the optimizations showed that the average trophic level of the catches is quite resistant to changes in the underlying system structure. On the other hand, despite the high level of aggregation of the model structure, a biodiversity index estimated by the model presented large changes as a function of the weights placed on the single policy functions, reflecting the changes in the system structure. The output of the application of the fishing optimization presented here should be considered in qualitative rather than in quantitative terms as an aid and part contribution to the complicated discussions on future long term management actions. Nonetheless it points to an overall reduction in fishing capacity, an objective widely accepted within the scientific community, while keeping the fishery in a profitable state.     

Lake of flies, or lake of fish? A trophic model of Lake Malawi

Ecosystem-focused models have, for the first time, become available for the combined demersal and pelagic components of a large tropical lake ecosystem, Lake Malawi. These provide the opportunity to explore continuing controversies over the production efficiencies and ecological functioning of large tropical lakes. In Lake Malawi these models can provide important insight to the effect of fishing on fish composition, and the potential competition that the lakefly Chaoborus edulis may have with fisheries production. A mass-balanced trophic model developed for the demersal fish community of the southern and western areas of Lake Malawi was integrated with an existing trophic model developed for the open-water pelagic. Input parameters for the demersal model were obtained from a survey of fish distributions, fish food consumption studies, and from additional published quantitative and qualitative information on the various biotic components of the community. The model was constructed using the Ecopath approach and software. The graphically presented demersal food web spanned four trophic levels and was based primarily on consumption of detritus, zooplankton and sedimented diatoms. Zooplankton was imported into the system at trophic levels three and four through fish predation on carnivorous and herbivorous copepods and Chaoborus larvae. It is proposed that the primary consumption of copepods was by fish migrating into the pelagic zone. Chaoborus larvae in the demersal were probably consumed near the lakebed as they conducted a daily migration from the pelagic to seek refuge in the sediments. This evidence for strong benthic-pelagic coupling provided the opportunity for linking the demersal model to the existing model for the pelagic community so producing the first model for the complete ecosystem. Energy fluxes through the resulting combined model demonstrated that the primary import of biomass to the demersal system was detritus of pelagic origin (72.1%) and pelagic zooplankton (10.6%). Only 15.8% of the biomass consumed within the demersal system was of demersal origin. Lakefly production is efficiently utilised by the lake fish community, and any attempt to improve fishery production through introduction of a non-native plantivorous fish species would have a negative impact on the stability and productivity of the lake ecosystem.     

Invasive species impacts on ecosystem structure and function: A comparison of Oneida Lake, New York, USA, before and after zebra mussel invasion

Exotic species invasion is widely considered to affect ecosystem structure and function. Yet, few contemporary approaches can assess the effects of exotic species invasion at such an inclusive level. Our research presents one of the first attempts to examine the effects of an exotic species at the ecosystem level in a quantifiable manner. We used ecological network analysis (ENA) and a social network analysis (SNA) method called cohesion analysis to examine the effect of zebra mussel (Dreissena polymorpha) invasion on the Oneida Lake, New York, USA, food web. We used ENA to quantify ecosystem function through an analysis of food web carbon transfer that explicitly incorporated flow over all food web paths (direct and indirect). The cohesion analysis assessed ecosystem structure through an organization of food web members into subgroups of strongly interacting predators and prey. Our analysis detected effects of zebra mussel invasion throughout the entire Oneida Lake food web, including changes in trophic flow efficiency (i.e., carbon flow among trophic levels) and alterations of food web organization (i.e., paths of carbon flow) and ecosystem activity (i.e., total carbon flow). ENA indicated that zebra mussels altered food web function by shunting carbon from pelagic to benthic pathways, increasing dissipative flow loss, and decreasing ecosystem activity. SNA revealed the strength of zebra mussel perturbation as evidenced by a reorganization of food web subgroup structure, with a decrease in importance of pelagic pathways, a concomitant rise of benthic pathways, and a reorganization of interactions between top predator fish. Together, these analyses allowed for a holistic understanding of the effects of zebra mussel invasion on the Oneida Lake food web.     

Reciprocal subsidies between freshwater and terrestrial ecosystems structure consumer resource dynamics

Cross-ecosystem movements of material and energy, particularly reciprocal resource fluxes across the freshwater-land interface, have received major attention. Freshwater ecosystems may receive higher amounts of subsidies (i.e., resources produced outside the focal ecosystem) than terrestrial ecosystems, potentially leading to increased secondary production in freshwaters. Here we used a meta-analytic approach to quantify the magnitude and direction of subsidy inputs across the freshwater-land interface and to determine subsequent responses in recipient animals. Terrestrial and freshwater ecosystems differed in the magnitude of subsidies they received, with aquatic ecosystems generally receiving higher subsidies than terrestrial ecosystems. Surprisingly, and despite the large discrepancy in magnitude, the contribution of these subsidies to animal carbon inferred from stable isotope composition did not differ between freshwater and terrestrial ecosystems, likely due to the differences in subsidy quality. The contribution of allochthonous subsidies was highest to primary consumers and predators, suggesting that bottom-up and top-down effects may be affected considerably by the input of allochthonous resources. Future work on subsidies will profit from a food web dynamic approach including indirect trophic interactions and propagating effects.     

Impact of maximum sustainable yield on independent populations

We investigate a community of independent logistically growing populations under a common harvesting effort which leads to the total maximum sustainable yield (TMSY). It is surprising that in the case of two populations with approximately equal carrying capacities, TMSY is reached while both populations persist, although their biotic potential may differ substantially. In general, however, TMSY with a common harvesting effort implies suboptimal fishing of some populations, overfishing of others and extinction of the rest of the populations. Since extinction of populations is a rule rather than an exception and since a community of independent populations is more robust than an ecosystem with multiple trophic levels, we call for urgent retraction of all legal documents advocating MSY in ecosystems.     

Effects of Fishing on the Ecosystem Structure of Coral Reefs

Overfishing is considered one of the three most significant threats to coral reef ecosystems. Exponentially increasing human populations in the tropics have placed enormous demands upon reefs as a food source. At high intensities, termed ecosystem or Malthusian overfishing, fishing causes major direct and indirect effects on the community structure of fishes and other organisms. It reduces species diversity and leads to local extinctions not only of target species but also of other species not fished directly. Conceivably it could also lead to global extinctions. Loss of keystone species, such as predators of echinoderms, through fishing, can lead to major effects on reef processes, such as accretion of calcium carbonate. Ultimately, sustained heavy fishing may lead to loss of entire functional groups of species, resulting in impairment of the potentially important ecosystem functions provided by those groups. Overfishing has been shown to interact with other agents of disturbance to reduce the ability of reefs to recover from natural occurrences such as hurricanes. Effective management of fishing will require a deeper understanding of the effects of exploitation than we now possess. Research initiatives are underway to examine the responses of fish populations to fishing, generally responses to protection from fishing. There is, however, an urgent need to look beyond fish communities and to consider the entire reef ecosystem. Studies that integrate population and community biology with ecosystem processes will provide a much better understanding of the effects of biodiversity loss on reef function and will improve our ability to manage these complex systems.