Food Web Model Research Articles

Selection of trilateral continuums of life history strategies under food web interactions

The study of life history strategies has a long history in ecology and evolution, but determining the underlying mechanisms driving the evolution of life history variation and its consequences for population regulation remains a major challenge. In this study, a food web model with constant environmental conditions was used to demonstrate how multi-species consumer–resource interactions (food-web interactions) can create variation in the duration of the adult stage, age of maturation, and fecundity among species. The model included three key ecological processes: size-dependent species interactions, energetics, and transition among developmental stages. Resultant patterns of life history variation were consistent with previous empirical observations of the life history strategies of aquatic organisms referred to as periodic, equilibrium, and opportunistic strategies (trilateral continuums of life history strategies). Results from the simulation model suggest that these three life history strategies can emerge from food web interactions even when abiotic environmental conditions are held constant.

Connecting ecology to daily life: how students and teachers relate food webs to the food they eat

This study used sociocultural learning theory to better understand how middle and high school environmental science and biology students and pre- and in-service science teachers connect the daily life activity of eating to the food web model learned in school. We sought to understand how student and teacher perceptions of the environment and their experiences influenced their responses to interview questions regarding this topic. Findings, based on transcribed interviews with 54 study participants, indicate that three quarters of teachers and students were unable to connect the food they eat with ecosystem food webs. Even so, many respondents particularly those from elite public schools, did not demonstrate common food web misconceptions identified by other researchers, instead showing a sophisticated understanding of food web interactions. These findings indicate that even though participants were proficient in their school science understanding of food web interactions, they did not readily think about how their everyday out of school activities, like eating, relate to those interactions. This may be representative of a more general disconnect between formal ecology instruction and daily life activities. We provide several recommendations for how this disconnect can be remedied in our classrooms.

Quantifying effects of hydrological and water quality disturbances on fish with food-web modeling

Accurately delineating the effects of hydrological and water quality habitat factors on the aquatic biota will significantly assist the management of water resources and restoration of river ecosystems. However, current models fail to comprehensively consider the effects of multiple habitat factors on the development of fish species. In this study, a dynamic framework for river ecosystems was set up to explore the effects of multiple habitat factors in terms of hydrology and water quality on the fish community in rivers. To achieve this the biomechanical forms of the relationships between hydrology, water quality, and aquatic organisms were determined. The developing processes of the food web without external disturbance were simulated by 208 models, constructed using Ecopath With Ecosim (EWE). These models were then used to analyze changes in biomass (ΔB) of two representative fish species, Opsariichthys bidens and Carassius auratus, which are widely distributed in Asia, and thus have attracted the attention of scholars and stakeholders, due to the consequence of habitat alteration. Results showed that the relationship between the changes in fish biomass and key habitat factors can be expressed in a unified form. T-tests for the unified form revealed that the means of the two data sets of simulated and observed ΔB for these two fish species (O. bidens and C. auratus) were equal at the significance level of 5%. Compared with other ecological dynamic models, our framework includes theories that are easy to understand and has modest requirements for assembly and scientific expertise. Moreover, this framework can objectively assess the influence of hydrological and water quality variance on aquatic biota with simpler theory and little expertise. Therefore, it is easy to be put into practice and can provide a scientific support for decisions in ecological restoration made by river administrators and stakeholders across the world.

Ecosystem modeling as a framework to convert a multi-disciplinary research approach into a useful model for the Araçá Bay (Brazil)

A model-oriented research project can organize and systematize high quality sampling information and convert observed values into information needed to parameterize ecological models. In this paper, we describe the value parameterization process from an interdisciplinary project to the development of a food web model (using Ecopath) in order to comprehend the Araçá Bay ecosystem structure (Brazil) and to forecast the impact of a port expansion (São Sebastião Port) over a bay environment. Araçá Bay Ecopath model has 34 compartments, Phytoplankton, Phytobenthos, Mangrove, Zooplankton, 10 groups of benthos, 13 fish groups, Shrimp, Crabs, Turtles, two bird groups and two groups of Detritus. The model outputs showed that Araçá Bay is a mature and detritus-based ecosystem. It is highly influenced by the role of benthos groups which are responsible for a huge amount of detritus recycling due to their large biomass value. The simulation of port expansion (primary producers' reduction), using the Ecosim module, indicated negative impacts on almost all living groups and an increase in detritus accumulation, leading the entire bay ecosystem towards its collapse (in the short term). The interdisciplinary organized sampling process presented here is an example of how objectively planned sample design and modeling may guide scientists, local people and stakeholders' decisions with valuable integrated information and overall predictions in order to consider the sustainable use of natural areas and resources.

Impact of stochastic migration on species diversity in meta-food webs consisting of several patches

The structure of space has an appreciable influence on the diversity and stability of ecosystems. So far, there are only few theoretical studies investigating the population dynamics of food webs consisting of many species that can migrate between several patches, and in most of these models migration is a continuous, deterministic process. However, when migration events are rare (for instance because the patches are far apart), migration is a stochastic process and should be modeled accordingly. We present computer simulations of a food web model of many species on a spatial network of several patches, combining deterministic local population dynamics with stochastic migration. We evaluate the influence of the migration rate and other model parameters on local and regional species diversity and on stability. We find that migration increases the number of surviving and coexisting populations by two effects. These are the rescue effect, which restores local populations that have gone extinct, and dynamical coexistence, which sustains local populations that could not persist in the absence of immigration. Both effects occur even when migration events are rare. Species diversity increases on local and regional scales with the frequency of migration events. Furthermore, we investigate the adiabatic limit in which population dynamics always reaches an equilibrium before the next migration event, and we investigate the possible long-term scenarios. While the final state often contains the same food web on all patches, we also find instances where two slightly different food webs coexist on different patches, even when initially each patch contained the same food web.

Sources of variation in stomach contents of predators of Atlantic herring in the Northwest Atlantic during 1973–2014

AbstractSpatial and temporal variation in stomach-contents data is often unquantified or combined in such a way (e.g. averaged among years) that true signal in diets may be lost. Using a delta approach, this paper fits generalized additive mixed models (GAMMs) to the amount of Atlantic herring (Clupea harengus) identified in predator stomachs using only data from stomachs in which herring occurred, and to the probability that a stomach contained herring. Both the amount of herring in stomachs and the probability of a stomach containing herring varied seasonally, spatially, and among years. Of the random effects in each GAMM, the effect of predator species had the largest variance. An index of herring abundance derived from the stomach-contents data was generally consistent with recent herring stock assessments. The temporal and spatial variation in the stomach-contents data suggested that the effect of averaging or combining stomach-contents data among years, seasons, or areas may lead to falsely precise or biased estimates from multispecies assessments or in estimates of consumption, and may restrain the relevance of static foodweb models.

Seasonal dynamics and functioning of the Sylt-Rømø Bight, northern Wadden Sea

The Wadden Sea undergoes large seasonal changes in species abundance and biomass comprising its complex food web. This study examined four carbon food web models of the Sylt-Rømø Bight, one for each season. Each flow model consisted of 66 compartments depicting the respective biomass and energy budget of each ecosystem component and the flows between them. Ecological network analysis (ENA), a set of algorithms to evaluate the functioning of ecological networks, was used to assess the seasonal variability in the system properties of the Sylt-Rømø Bight food webs. We used an uncertainty analysis to quantitatively evaluate the significance of inter-seasonal differences. Clear seasonal variation was observed in most of the whole system indicators such as the flow diversity, the effective link density and the relative redundancy which varied by 12.8%, 17.3% and 10.3% respectively between the highest in summer and the lowest during fall and winter, whereas the relevant ascendency ratio was the highest in winter during the least active months. Other indices such as the average mutual information index, which fluctuated between 1.73 in fall and 1.79 in spring, showed no significant variation between seasons. Results from ENA have great potential for ecosystem management, as it provides a holistic assessment of the functioning of ecosystems.

Microbial parasites make cyanobacteria blooms less of a trophic dead end than commonly assumed.

Parasites exist in every ecosystem and can have large influence on food web structure and function, yet, we know little about parasites' effect on food web dynamics. Here we investigate the role of microbial parasitism (viruses of bacteria, phytoplankton and cyanobacteria, and parasitic chytrids on cyanobacteria) on the dynamics of trophic pathways and food web functioning during a cyanobacteria bloom, using linear inverse food web modeling parameterized with a 2-month long data set (biomasses, infection parameters, etc.). We show the importance of grazing on heterotrophic bacteria (the microbial pathway: DOC → bacteria → consumer) and how consumers depended on bacteria during peak-cyanobacteria bloom, which abundance was partly driven by the viral activity. As bacteria become the main energy pathway to the consumers, the system takes a more web-like structure through increased omnivory, and may thereby facilitate the system's persistence to the cyanobacteria outbreak. We also showed how the killing of cyanobacteria host cells by chytrids had important impact on the food web dynamics by facilitating grazing on the cyanobacteria, and by offering alternative pathways to the consumers. This seemed to increase the system's ability to return to a mix of trophic pathways, which theoretically increases the stability of the system.

Variability and stability in predation landscapes: A cross‐ecosystem comparison on the potential for predator control in temperate marine ecosystems

AbstractPredation can play an important role in population, community and ecosystem processes. When predator guilds are diverse, fluctuations in individual predator populations may have small influences on the guild at large, suggesting that predator diversity stabilizes the amount of predation prey experience. The strength of this phenomenon depends on how synchronously populations within predator assemblages vary and whether all predators are equally important consumers. We utilized a database of biomasses of fish species across ten temperate marine ecosystems paired with diet composition and feeding rate information from mass‐balance food web models to develop a predation index that weights the importance of predators on a prey based on predator feeding habits. Using the index, we asked how diverse sources of predation in these ecosystems are and how much diversity stabilizes variability in predation. Predator assemblages displayed a wide range of diversity; in one‐third of assemblages, a single predator group accounted for over half of all predation. Abundances of predator populations within assemblages generally varied independently of one another (neither synchronously nor asynchronously), implying an intermediate stabilization on predation intensity by predator diversity. Accounting for interaction strength (versus mere presence) is critical for interpreting the predator landscape; doing so led to a wider range in predator assemblage diversity and less synchronous assemblages. This work challenges conventional notions that marine food webs are diverse and therefore less susceptible to predator control. Future work should consider the temporal variability of the predator assemblages and account for differences in mortality induced by each predator population.

Lake Champlain offshore benthic invertebrate community before and after zebra mussel invasion

Abstract Benthic invertebrates are important bio-indicators of water quality and play a significant role in aquatic systems. Lake Champlain has limited benthic invertebrate data which hinders development of food web models, assessment of invasive species impacts, and evaluation of management actions. In June 2016, we assessed benthic invertebrates along three transects in the main basin of Lake Champlain ranging from 5 to 100 m, and then compared results to densities from a limited survey in 1991 prior to the zebra mussel (Dreissena polymorpha) invasion. In 2016, total biomass and density were 1–2 orders of magnitude greater at 5 m than at 20–100 m. Zebra mussels, chironomids, oligochaetes, and gastropods were dominant at 5 m, and oligochaetes and sphaeriids were dominant at 20–100 m. Total density at the 5-m site was 94% lower in 2016 compared to 1991, but similar at the 100-m site. Diporeia, while abundant in many freshwater bodies, is historically rare in Lake Champlain and was not detected in our sampling. Because Lake Champlain benthic invertebrate densities are low and display dissimilar distributions to the Great Lakes, we hypothesize the offshore fish community is likely much more reliant on pelagic rather than benthic production. Although the current composition and biomass suggest the benthic community in Lake Champlain may not be greatly impacted by an invasion of quagga mussel (D. rostriformis bugensis), the potential for quaggas to re-route energy from pelagic to benthic habitats, as it has in the Great Lakes, could limit the Lake Champlain offshore fish community.

Holistic assessment of Chwaka Bay's multi-gear fishery – Using a trophic modeling approach

Abstract East African coastal communities highly depend on marine resources for not just income but also protein supply. The multi-species, multi-gear nature of East African fisheries makes this type of fishery particularly difficult to manage, as there is a trade-off between maximizing total catch from all gears and species and minimizing overfishing of target species and the disintegration of the ecosystem. The use and spatio-temporal overlap of multiple gears in Chwaka Bay (Zanzibar) has led to severe conflicts between fishermen. There is a general concern of overfishing in the bay because of the widespread use of small mesh sizes and destructive gears such as dragnets and spear guns. We constructed an Ecopath food web model to describe the current trophic flow structure and fishing pattern of the bay. Based on this model, we explored the impact of different gears on the ecosystem and the fishing community in order to give advice for gear based management in the bay. Results indicate that Chwaka bay is a productive, shallow water system, with biomass concentrations around the first and second trophic level. The system is greatly bottom-up driven and dominated by primary producers and invertebrates. The trophic and network indicators as well as the community energetics characterize Chwaka Bay as relatively mature. Traps and dragnets have the strongest impact on the ecosystem and on the catches obtained by other gears. Both gears potentially destabilize the ecosystem by reducing the biomass of top-down controlling key species (including important herbivores of macroalgae). The dragnet fishery is the least profitable, but provides most jobs for the fishing community. Thus, a complete ban of dragnets in the bay would require the provision of alternative livelihoods. Due to the low resource biomass of fish in the bay and the indication of a loss of structural control of certain fish groups, Chwaka Bay does not seem to provide scope for further expansion of the fishery. Instead, we recommend an effort control of traps and a reduction in the use of dragnets, partially by redistributing them to the more profitable and less impacting gears (e.g. longlines, gillnets, handlines).

Quantifying carbon fluxes from primary production to mesopelagic fish using a simple food web model

Abstract An ecosystem-based flow analysis model was used to study carbon transfer from primary production (PP) to mesopelagic fish via three groups of copepods: detritivores that access sinking particles, vertical migrators, and species that reside in the surface ocean. The model was parameterized for 40°S to 40°N in the world ocean such that results can be compared with recent estimates of mesopelagic fish biomass in this latitudinal range, based on field studies using acoustic technologies, of ∼13 Gt (wet weight). Mesopelagic fish production was predicted to be 0.32% of PP which, assuming fish longevity of 1.5 years, gives rise to predicted mesopelagic fish biomass of 2.4 Gt. Model ensembles were run to analyse the uncertainty of this estimate, with results showing predicted biomass >10 Gt in only 8% of the simulations. The work emphasizes the importance of migrating animals in transferring carbon from the surface ocean to the mesopelagic zone. It also highlights how little is known about the physiological ecology of mesopelagic fish, trophic pathways within the mesopelagic food web, and how these link to PP in the surface ocean. A deeper understanding of these interacting factors is required before the potential for utilizing mesopelagic fish as a harvestable resource can be robustly assessed.

Unraveling the intricate dynamics of planktonic Arctic marine food webs. A sensitivity analysis of a well-documented food web model

The extreme and variable environment shapes the functioning of Arctic ecosystems and the life cycles of its species. This delicate balance is now threatened by the unprecedented pace and magnitude of global climate change and anthropogenic pressure. Understanding the long-term consequences of these changes remains an elusive, yet pressing, goal. Our work was specifically aimed at identifying which biological processes impact Arctic planktonic ecosystem functioning, and how. Ecological Network Analysis (ENA) indices reveal emergent ecosystem properties that are not accessible through simple in situ observation. These indices are based on the architecture of carbon flows within food webs. But, despite the recent increase in in situ measurements from Arctic seas, many flow values remain unknown. Linear inverse modeling (LIM) allows missing flow values to be estimated from existing flow observations and, subsequent reconstruction of ecosystem food webs. Through a sensitivity analysis on a LIM model of the Amundsen Gulf in the Canadian Arctic, we were able to determine which processes affected the emergent properties of the planktonic ecosystem. The analysis highlighted the importance of an accurate knowledge of the various processes controlling bacterial production (e.g. bacterial growth efficiency and viral lysis). More importantly, a change in the fate of the microzooplankton within the food web can be monitored through the trophic level of mesozooplankton. It can be used as a “canary in the coal mine” signal, a forewarner of larger ecosystem change.

Disentangling food webs interactions in the littoral of temperate shallow lakes

Elucidation of food web interactions provides better understanding of ecosystem functioning, indicates anthropogenic impacts which often cause alterations in environmental conditions and detoriorations in feeding network in terrestrial and aquatic ecosystems. The importance of microfauna and macroinvertebrates in littoral zones of shallow waters has poorly been studied regarding their trophic interactions. This study compares invertebrate assemblage and food web interactions among epiphyton, zooplankton and benthic macroinvertebrates across structural heterogeneity in the littoral zone of three temperate shallow water bodies. Submerged and structurally-complex stands of Ceratophyllum demersum inhabited higher abundance of cladocerans and supported higher diversity and biomass of epiphytic protozoans and invertebrates in comparison to floating-leaved Nuphar lutea stands. Analysis of the ecosystem functioning illustrated the increased biomass of macroinvertebrate predators and a preference of predation over planktonic crustaceans in more complex macrophyte stands. Food webs displayed higher complexity and size with habitat heterogeneity, while epiphyton and zooplankton shared the important fraction in food resources among the invertebrate trophic network. Results of food web modelling indicated zooplankton and epiphyton as more vulnerable to invertebrate predation in the complex submerged macrophytes than in floating-leaved macrophyte stands. Integrated community, ecosystem and food web approaches in explanation of complex trophic interactions in the littoral zones justified an increase in diversity and food-web functional complexity with structural heterogeneity of microhabitats.

Ecological effects of sex differ with trophic positions in a simple food web.

Sexual differences in parental investment, predation pressure, and foraging efforts are common in nature and affect the trophic flow in food webs. Specifically, the sexual differences in predator and prey behavior change in trophic inflow and outflow, respectively, while those in parental investment alter the reproductive allocation of acquired resources in the population. Consequently, these factors may play an important role in determining the system structure and persistence. However, few studies have examined how sexual differences in trophic flow affect food web dynamics. In this study, I show the ecological role of sex by explicitly incorporating sexual differences in trophic flow into a three‐species food web model. The results demonstrated that the ecological waste of males, that is, the amount of trophic inflow into males with less parental investment, plays an important role in system persistence and structure. In particular, the synergy between sexual differences in parental investment and trophic inflows and outflows is important in determining web persistence: Significant impacts of male‐biased trophic flows require the condition of anisogamy. In addition, the dynamic effects of the ecological waste of males differ with trophic level: The coexistence of a food web occurs more frequently with biased inflows into predator males, but occurs less frequently with biased inflows into consumer males. The model analysis indicates that investigating the pattern of sexual differences among trophic positions can enrich our understanding of food web persistence and structure in the real world.

Climate-mediated changes in marine ecosystem regulation during El Niño.

The degree to which ecosystems are regulated through bottom-up, top-down, or direct physical processes represents a long-standing issue in ecology, with important consequences for resource management and conservation. In marine ecosystems, the role of bottom-up and top-down forcing has been shown to vary over spatio-temporal scales, often linked to highly variable and heterogeneously distributed environmental conditions. Ecosystem dynamics in the Northeast Pacific have been suggested to be predominately bottom-up regulated. However, it remains unknown to what extent top-down regulation occurs, or whether the relative importance of bottom-up and top-down forcing may shift in response to climate change. In this study, we investigate the effects and relative importance of bottom-up, top-down, and physical forcing during changing climate conditions on ecosystem regulation in the Southern California Current System (SCCS) using a generalized food web model. This statistical approach is based on nonlinear threshold models and a long-term data set (~60years) covering multiple trophic levels from phytoplankton to predatory fish. We found bottom-up control to be the primary mode of ecosystem regulation. However, our results also demonstrate an alternative mode of regulation represented by interacting bottom-up and top-down forcing, analogous to wasp-waist dynamics, but occurring across multiple trophic levels and only during periods of reduced bottom-up forcing (i.e., weak upwelling, low nutrient concentrations, and primary production). The shifts in ecosystem regulation are caused by changes in ocean-atmosphere forcing and triggered by highly variable climate conditions associated with El Niño. Furthermore, we show that biota respond differently to major El Niño events during positive or negative phases of the Pacific Decadal Oscillation (PDO), as well as highlight potential concerns for marine and fisheries management by demonstrating increased sensitivity of pelagic fish to exploitation during El Niño.

The origin of motif families in food webs

Food webs have been found to exhibit remarkable “motif profiles”, patterns in the relative prevalences of all possible three-species subgraphs, and this has been related to ecosystem properties such as stability and robustness. Analysing 46 food webs of various kinds, we find that most food webs fall into one of two distinct motif families. The separation between the families is well predicted by a global measure of hierarchical order in directed networks—trophic coherence. We find that trophic coherence is also a good predictor for the extent of omnivory, defined as the tendency of species to feed on multiple trophic levels. We compare our results to a network assembly model that admits tunable trophic coherence via a single free parameter. The model is able to generate food webs in either of the two families by varying this parameter, and correctly classifies almost all the food webs in our database. This is in contrast with the two most popular food web models, the generalized cascade and niche models, which can only generate food webs within a single motif family. Our findings suggest the importance of trophic coherence in modelling local preying patterns in food webs.

Food-web dynamics under climate change.

Climate change affects ecological communities through its impact on the physiological performance of individuals. However, the population dynamic of species well inside their thermal niche is also determined by competitors, prey and predators, in addition to being influenced by temperature changes. We use a trait-based food-web model to examine how the interplay between the direct physiological effects from temperature and the indirect effects due to changing interactions between populations shapes the ecological consequences of climate change for populations and for entire communities. Our simulations illustrate how isolated communities deteriorate as populations go extinct when the environment moves outside the species' thermal niches. High-trophic-level species are most vulnerable, while the ecosystem function of lower trophic levels is less impacted. Open communities can compensate for the loss of ecosystem function by invasions of new species. Individual populations show complex responses largely uncorrelated with the direct impact of temperature change on physiology. Such complex responses are particularly evident during extinction and invasion events of other species, where climatically well-adapted species may be brought to extinction by the changed food-web topology. Our results highlight that the impact of climate change on specific populations is largely unpredictable, and apparently well-adapted species may be severely impacted.

Evolution of Future Black Sea Fish Stocks under Changing Environmental and Climatic Conditions

Here we present a case study towards producing quantitative scientific advice on the application of the EU Common Fisheries Policy (CFP) in the Black Sea. We provide estimates of fishing mortality rates at levels which will lead to rebuilding and maintaining stocks above biomass levels that could produce maximum sustainable yield (MSY) under the IPCC RCP4.5 future climate scenario together with the business as usual (BAU) river discharge scenario. In this study, we have implemented a coupled, basin-scale circulation-biogeochemical model and used its output to feed a food web model to test near-future changes that may be observed in the Black Sea ecosystem under the influence of contemporary fisheries exploitation conditions. In order to test model response to changes in climate and related drivers, the future climate scenario (2015-2020) simulation was compared to the present day (2000-2014) simulation. Likewise, to test the sensitivity of the higher trophic level food web model to changes in fishing pressure, a future estimate of fishing pressure was projected based on its respective contemporary value and applied to each fish stock. Using these models, fishing mortality rates that could produce the maximum sustainable yield (FMSY) in future years 2015-2020 and ensure the long-term recovery of the predatory fish stocks of the Black Sea are predicted. Future projections suggest that all fish stock will decrease in all the regions of the Black Sea except for sprat. Anchovy is expected to show the highest decrease in biomass. Analyses on FMSY estimates show that a significant reduction in fisheries exploitation is required for the sustainable management of the Black Sea ecosystems and the related services. This study, for the first time, presents future stock size, FMSY and MSY estimates for the Black Sea for 11 fish species. FMSY values are generally lower than estimates of the scientific, technical and economic committee for fisheries (STECF), mainly because of the explicit food web interactions that the modeling system allows to be considered.

Predicting impacts of lionfish (Pterois volitans) invasion in a coastal ecosystem of southern Brazil

The opportunistic feeder Pterois volitans is a voracious invader, causing large impacts in marine food-webs. We have used a Ecopath-with-Ecosim model to hypothesize an invasion by lionfish and to predict the likely impact of this potential generalist mesopredator in a subtropical food-web model. With thirty-three functional groups, the initial Ecopath model was balanced with a low biomass of lionfish (0.07 t/km2). In Ecosim, three scenarios of different vulnerability settings for the linkages between the introduced fish and its prey were tested, representing the default setting, a top-down control, and an extreme top-down control. The scenarios were tested using different assumptions on the ability of the invasive fish to change the proportions of prey consumed according to prey availability. Our model predicted that the hypothesized lionfish invasion would have a strong impact on this subtropical marine food web: (1) by reducing prey populations and, consequently, food for native predators, and; (2) by predating on key species, causing direct impacts and possibly cascading trophic effects. Reef fish were the most affected, including some groups ecologically and economically important, like lutjanids and groupers. However, some adaptations in the fishing strategy of fishermen are expected which may affect other fish groups. Stakeholders should be warned of the potential ecological and socio-economic impacts that may arise from a lionfish invasion and various strategies and policy options should be immediately developed and applied (1) to prevent the arrival and establishment of the lionfish, and; (2) to make the ecosystem more resilient to this and other possible exotic species.