The stability of trophic mass-balance models of marine ecosystems: a comparative analysis

Dynamic simulations of 18

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mass-balance marine trophic models are used to explore the stability of systems when briefly impacted by a fishery on the key ‘wasp–waist' populations occurring at intermediate trophic levels. The results are related to different ecosystem goal functions previously identified as representative of three attributes of ecosystems development: community complexity, homeostasis and energetics. System recovery time, the time required for all functional groups to returns to baseline level, and here used as a measure of model stability, was inversely correlated to Finn's Cycling Index, i.e. to the fraction of ecosystem throughput that is recycled, and to the mean length of trophic pathways in the systems. Systems with higher capacity to recycle detritus are systems with a higher ability to recover from perturbations. The results are in agreement with the E.P. Odum's theory of ecosystem development, where recycling is interpreted as a chief positive feedback mechanism that contributes to stability in the mature systems by preventing overshoots and destructive oscillations due to external impacts.     

Modelling partitioning and distribution of micropollutants in the lagoon of Venice: a first step towards a comprehensive ecotoxicological model

The model presented in this paper integrates a large amount of recent and ad hoc collected data concerning environmental contamination from micropollutants in the lagoon of Venice. This model represents the first step in setting up of an ecotoxicological model for the Venice lagoon, to simulate fate of contaminants from abiotic matrices to organisms. Distribution and partitioning of organic and inorganic contaminants are modelled by a two-dimensional model, based both on deterministic and empirical submodels and adapted to a large spectrum of different substances (polychlorinated dibenzo-dioxins/polychlorinated dibenzo-furans (PCDD/F), polychlorinated biphenyls (PCBs), heavy metals). The model was successfully calibrated on a wide set of experimental data. Sensitivity analysis showed that the model is generally not very sensible to parameters values but it is sensible to external conditions (e.g., pollutants loads). Distribution of dissolved and total concentrations of contaminants was obtained for a series of PCDD/F and PCBs congeners and for eight heavy metals. These distributions represent integrated information on ecosystem health, complementary to monitoring data and they are useful to be used for comparisons with various water quality criteria. Simulation scenarios under different external conditions are proposed as examples of use of the model for management purposes.     

Bridging biogeochemical and food web models for an End-to-End representation of marine ecosystem dynamics: The Venice lagoon case study

The simultaneous accounting of effects of natural and anthropogenic changes within a common framework calls for the development of new comprehensive tools capable of integrating biological processes that span a huge range of scales, from viruses to fishes, in addition to their interactions with physical–chemical environmental properties, i.e. End-to-End models.     

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.     

Evidence for the dominance of indirect effects in 50 trophic ecosystem networks

Indirect effects are powerful influences in ecosystems that may maintain species diversity and alter apparent relationships between species in surprising ways. Here, we applied network environ analysis to 50 empirically-based trophic ecosystem models to test the hypothesis that indirect flows dominate direct flows in ecosystem networks. Further, we used Monte Carlo based perturbations to investigate the robustness of these results to potential error in the underlying data. To explain our findings, we further investigated the importance of the microbial food web in recycling energy-matter using components of the Finn Cycling Index and analysis of environ centrality. We found that indirect flows dominate direct flows in 37/50 (74.0%) models. This increases to 31/35 (88.5%) models when we consider only models that have cycling structure and a representation of the microbial food web. The uncertainty analysis reveals that there is less error in the I/D values than the ±5% error introduced into the models, suggesting the results are robust to uncertainty. Our results show that the microbial food web mediates a substantial percentage of cycling in some systems (median = 30.2%), but its role is highly variable in these models, in agreement with the literature. Our results, combined with previous work, strongly suggest that indirect effects are dominant components of activity in ecosystems.     

A trophic cascade triggers collapse of a salt-marsh ecosystem with intensive recreational fishing

Overexploitation of predators has been linked to the collapse of a growing number of shallow-water marine ecosystems. However, salt-marsh ecosystems are often viewed and managed as systems controlled by physical processes, despite recent evidence for herbivore-driven die-off of marsh vegetation. Here we use field observations, experiments, and historical records at 14 sites to examine whether the recently reported die-off of northwestern Atlantic salt marshes is associated with the cascading effects of predator dynamics and intensive recreational fishing activity. We found that the localized depletion of top predators at sites accessible to recreational anglers has triggered the proliferation of herbivorous crabs, which in turn results in runaway consumption of marsh vegetation. This suggests that overfishing may be a general mechanism underlying the consumer-driven die-off of salt marshes spreading throughout the western Atlantic. Our findings support the emerging realization that consumers play a dominant role in regulating marine plant communities and can lead to ecosystem collapse when their impacts are amplified by human activities, including recreational fishing.     

Analysis of annual fluctuations of C. nodosa in the Venice lagoon: Modeling approach

A simple simulation model was developed to describe the growth trends of Cymodocea nodosa (Ucria) Ascherson based on data sets from the Venice lagoon. The model reproduces the seasonal fluctuations in the above and belowground biomass and in shoot density. The modeling results are in good agreement with data on net production, growth rates and chemical–physical parameters of water. It was assumed that light and temperature are the most important factors controlling C. nodosa development, and that the growth was not limited by nutrient availability. The aim was to simulate biomass production as a function of external forcing variables (light, water temperature) and internal control (plant density). A series of simulation experiments were performed with the basic model showing that among the most important phenomena affecting C. nodosa growth are: (1) inhibition of production and recruitment of new shoots by high temperature and (2) light attenuation due to seasonal fluctuation.     

Using a fisheries ecosystem model with a water quality model to explore trophic and habitat impacts on a fisheries stock: A case study of the blue crab population in the Chesapeake Bay

Recent calls for the development of ecosystem-based fisheries management compel the development of resource management tools and linkages between existing fisheries management tools and other resource tools to enable assessment and management of multiple impacts on fisheries resources. In this paper, we describe the use of the Chesapeake Bay Fisheries Ecosystem Model (CBFEM), developed using the Ecopath with Ecosim (EwE) software, and the Chesapeake Bay Water Quality Model (WQM) to demonstrate how linkages between available modeling tools can be used to inform ecosystem-based natural resource management. The CBFEM was developed to provide strategic ecosystem information in support of fisheries management. The WQM was developed to assess impacts on water quality. The CBFEM was indirectly coupled with the WQM to assess the effects of water quality and submerged aquatic vegetation (SAV) on blue crabs. The output from two WQM scenarios (1985-1994), a baseline scenario representing actual nutrient inputs and another with reduced inputs based on a tributary management strategy, was incorporated into the CBFEM. The results suggested that blue crab biomass could be enhanced under management strategies (reduced nutrient input) when the effective search rate of blue crab young-of-the-year's (YOY's) predators or the vulnerability of blue crab YOY to its predators was adjusted by SAV. Such model linkages are important for incorporating physical and biological components of ecosystems in order to explore ecosystem-based fisheries management options.     

Disentangling trophic relationships in a High Arctic tundra ecosystem through food web modeling

Determining the manner in which food webs will respond to environmental changes is difficult because the relative importance of top-down vs. bottom-up forces in controlling ecosystems is still debated. This is especially true in the Arctic tundra where, despite relatively simple food webs, it is still unclear which forces dominate in this ecosystem. Our primary goal was to assess the extent to which a tundra food web was dominated by plant-herbivore or predator-prey interactions. Based on a 17-year (1993-2009) study of terrestrial wildlife on Bylot Island, Nunavut, Canada, we developed trophic mass balance models to address this question. Snow Geese were the dominant herbivores in this ecosystem, followed by two sympatric lemming species (brown and collared lemmings). Arctic foxes, weasels, and several species of birds of prey were the dominant predators. Results of our trophic models encompassing 19 functional groups showed that <10% of the annual primary production was consumed by herbivores in most years despite the presence of a large Snow Goose colony, but that 20-100% of the annual herbivore production was consumed by predators. The impact of herbivores on vegetation has also weakened over time, probably due to an increase in primary production. The impact of predators was highest on lemmings, intermediate on passerines, and lowest on geese and shorebirds, but it varied with lemming abundance. Predation of collared lemmings exceeded production in most years and may explain why this species remained at low density. In contrast, the predation rate on brown lemmings varied with prey density and may have contributed to the high-amplitude, periodic fluctuations in the abundance of this species. Our analysis provided little evidence that herbivores are limited by primary production on Bylot Island. In contrast, we measured strong predator-prey interactions, which supports the hypothesis that this food web is primarily controlled by top-down forces. The presence of allochthonous resources subsidizing top predators and the absence of large herbivores may partly explain the predominant role of predation in this low-productivity ecosystem.     

Significance of pelagic bacteria as a trophic resource in a coral reef lagoon,One Tree Island,Great Barrier Reef

Bacterial numbers and frequency of dividing cells (FDC) were estimated in surface waters at seven stations along a transeet across One Tree lagoon and reefs at 2 h intervals over one tidal cycle in May 1984 and once a day on 8 d in December 1984/January 1985. Cell numbers ranged between 1.24 and 13.10x10⁵ ml^-1, with an overall mean value of 4.81x10⁵. There was a progressive depletion in standing stock from windward to leeward stations across the lagoon, also reflected in the FDC, which generally showed similar dynamic patterns to cell numbers throughout the study. Predator-free seawater (filtered through 2 m pore-size filters) and mucus-enriched incubations were also used to establish the relationship between the growth rate () and FDC for coral-reef populations. Average growth rates predicted from FDC values ranged from 0.062 to 0.174 h^-1, which is equivalent to doubling times of 5.74 to 16.00 h, with an overall mean value of 11.5 h. These fast doubling times suggest that bacteria respond rapidly to pulses of enrichment as they float over the lagoon and reefs, characteristically achieving about one doubling per tidal cycle. This probably ensures that the bulk of labile organic matter and dissolved nutrients is conserved within the immediate coral-reef environment. Estimates of bacterial production also suggest that they may contribute up to 40% of total picoplankton production and about 25% of total microplankton production. Pelagic bacteria are therefore potentially a major food resource for benthic filter-feeders, especially for those mainly dependent on the smaller (pico) components of the plankton.     

A preliminary model of nutrient cycling in sediments of a Mediterranean lagoon

A model of nitrogen and phosphorus cycles in the sediment of a lagoon has been developed. This model was applied to the Thau lagoon (southern France). Sediment was sliced in three layers to reproduce the oxygen profile, which is simulated within the model. Following an equilibrium hypothesis, the model was calibrated against field data. State variables and fluxes were estimated in the sediment and across the sediment-water interface. A Monte Carlo sensitivity analysis was performed to determine the most sensitive parameters and sediment state variables. A dynamic simulation with varying oxygen concentrations then showed the influence of anoxia on the phosphorus and nitrogen fluxes between water and sediment.     

Heavy metals in Hediste diversicolor (polychaeta: nereididae) and salt marsh sediments from the lagoon of Venice (Italy)

The nereid polychaete Hediste diversicolor was frequently used as biomonitor of heavy metals contamination in estuaries. In the present work, the bioaccumulation of five heavy metals (Cd, Cr, Cu, Mn, Pb) in tissues of H. diversicolor collected from six salt marshes in the lagoon of Venice is studied by analysing the relationships between concentrations of the operationally-defined reactive fraction of trace metals in sediments and total concentrations in tissues. The characterisation of sediment samples shows a clear distinction between sites near and far from the Industrial Zone; no analogous pattern is observed for tissue concentrations suggesting that metal bioaccumulation is controlled by complex interactions between metal bioavailability and physiological factors. Considering relationships between metals, strong inverse correlations are observed in polychaetes and sediments for Cu and Cr concentrations suggesting a possible interaction between these two metals. The role played by organic matter (OM) in the availability of metal is highlighted considering inverse relationships found between bioaccumulation factors for Cu, Cr, Cd and OM.     

Evaluating weather effects on interannual variation in net ecosystem productivity of a coastal temperate forest landscape: A model intercomparison

Forest productivity is strongly affected by seasonal weather patterns and by natural or anthropogenic disturbances. However weather effects on forest productivity are not currently represented in inventory-based models such as CBM-CFS3 used in national forest C accounting programs. To evaluate different approaches to modelling these effects, a model intercomparison was conducted among CBM-CFS3 and four process models (ecosys, CN-CLASS, Can-IBIS and 3PG) over a 2500 ha landscape in the Oyster River (OR) area of British Columbia, Canada. The process models used local weather data to simulate net primary productivity (NPP), net ecosystem productivity (NEP) and net biome productivity (NBP) from 1920 to 2005. Other inputs used by the process and inventory models were generated from soil, land cover and disturbance records. During a period of intense disturbance from 1928 to 1943, simulated NBP diverged considerably among the models. This divergence was attributed to differences among models in the sizes of detrital and humus C stocks in different soil layers to which a uniform set of soil C transformation coefficients was applied during disturbances. After the disturbance period, divergence in modelled NBP among models was much smaller, and attributed mainly to differences in simulated NPP caused by different approaches to modelling weather effects on productivity. In spite of these differences, age-detrended variation in annual NPP and NEP of closed canopy forest stands was negatively correlated with mean daily maximum air temperature during July-September (T_amax) in all process models (R² = 0.4-0.6), indicating that these correlations were robust. The negative correlation between T_amax and NEP was attributed to different processes in different models, which were tested by comparing CO₂ fluxes from these models with those measured by eddy covariance (EC) under contrasting air temperatures (T_a). The general agreement in sensitivity of annual NPP to T_amax among the process models led to the development of a generalized algorithm for weather effects on NPP of coastal temperate coniferous forests for use in inventory-based models such as CBM-CFS3: NPP′ = NPP − 57.1 (T_amax − 18.6), where NPP and NPP′ are the current and temperature-adjusted annual NPP estimates from the inventory-based model, 18.6 is the long-term mean daily maximum air temperature during July-September, and T_amax is the mean value for the current year. Our analysis indicated that the sensitivity of NPP to T_amax was nonlinear, so that this algorithm should not be extrapolated beyond the conditions of this study. However the process-based methodology to estimate weather effects on NPP and NEP developed in this study is widely applicable to other forest types and may be adopted for other inventory based forest carbon cycle models.     

Application of a Random Forest algorithm to predict spatial distribution of the potential yield of Ruditapes philippinarum in the Venice lagoon, Italy

We present a modelling framework that combines machine learning techniques and Geographic Information Systems to support the management of an important aquaculture species, Manila clam (Ruditapes philippinarum). We use the Venice lagoon (Italy), the first site in Europe for the production of R. philippinarum, to illustrate the potential of this modelling approach. To investigate the relationship between the yield of R. philippinarum and a set of environmental factors, we used a Random Forest (RF) algorithm. The RF model was tuned with a large data set (n = 1698) and validated by an independent data set (n = 841). Overall, the model provided good predictions of site-specific yields and the analysis of marginal effect of predictors showed substantial agreement among the modelled responses and available ecological knowledge for R. philippinarum. The most influent environmental factors for yield estimation were percentage of sand in the sediment, salinity, and water depth. Our results agree with findings from other North Adriatic lagoons. The application of the fitted RF model to continuous maps of all the environmental variables allowed estimates of the potential yield for the whole basin. Such a spatial representation enabled site-specific estimates of yield in different farming areas within the lagoon. We present a possible management application of our model by estimating the potential yield under the current farming distribution and comparing it to a proposed re-organization of the farming areas. Our analysis suggests a reduction of total yield is likely to result from the proposed re-organization.     

Host glutamine synthetase activities in the giant clam—zooxanthellae symbiosis: effects of clam size,elevated ammonia and continuous darkness

Host tissues and zooxanthellae of the giant clam Tridacna gigas contained glutamine synthetase, with the highest transferase activities present in the gill, followed by the kidney, mantle, zooxanthellae, foot, heart and adductor muscle, in that order. Synthetase activities of glutamine synthetase in host tissues and zooxanthellae were in a similar order, but the differences were not so marked. Host tissues also contained hexokinase, glucose-6-phosphate dehydrogenase and malate dehydrogenase. Highest hexokinase activities were present in the heart, followed, in order, by the gill, mantle, adductor muscle and foot. Highest glucose-6-phosphate dehydrogenase activities were present in the gill, followed by the mantle, heart, adductor muscle and foot. All tissues assayed contained high malate dehydrogenase activities. There was no detectable glutamate dehydrogenase activity. Glutamine synthetase activity in gill and mantle tissue decreased by 1.6% with every 1 cm increase in clam size. Host glutamine synthetase activity decreased by 80% in gill tissue and by 45% in mantle tissue in clams which were maintained for 8 d in continuous darkness. Similar effects were found when clams were kept in light in the presence of elevated ammonia concentrations. It is suggested that both host and symbionts are nitrogen-deficient in small clams and that host glutamine synthetase plays a role in ammonia assimilation by the intact association.     

Zooplankton responses to hydrological and trophic variability in a Mediterranean coastal ecosystem (Lesina Lagoon,South Adriatic Sea)

Spatial and temporal variability in zooplankton was studied at eight stations located in the Lesina Lagoon (South Adriatic Sea) Salinity, temperature, dissolved oxygen and chlorophyll a (in the lagoon) at these stations were also assessed. The zooplankton community was characterised by clear seasonal oscillations and mostly represented by copepods and their larvae. The dominant copepod species were Calanipeda aquaedulcis and Acartia tonsa, which exhibited spatial–temporal segregation in the lagoon. C. aquaedulcis copepodites seemed to be better adapted to oligotrophic and oligohaline conditions compared with the A. tonsa population. The invasive species A. tonsa has completely replaced the formerly abundant Acartia margalefi. A positive correlation was found between abundances, total species numbers and trophic state. An increasing abundance trend was shown from the lagoon towards the sea. The highest diversity indices were recorded at the two channel inlets, during high tide due to the absence of a clear dominance of one or more coastal species and the co-occurrence of species of lagoon and marine origin.     

Parameter constraints in a stream ecosystem model: Incorporation of a priori information in Monte Carlo error analysis

Using a model of mass and nutrient dynamics in a stream, we evaluated the use of a priori information (e.g., mass balance constraints, limits for the state variables, etc.) to increase the accuracy and efficiency of Monte Carlo error analysis. The results indicated that a priori information can effectively eliminate physically unrealistic simulations and reduce uncertainty about the predicted values. Some types of a priori information proved more helpful than increased accuracy in measuring model parameters.     

A simulation model for the effects of burning on the phosphorus and nitrogen cycle of a heathland ecosystem

In this paper, a simulation model is presented for nutrient cycling in heathland ecosystems. The results of simulations are compared with field data of phosphorus and nitrogen accumulation in different compartments of the system in the years after burning, given in the literature on this subject (Chapman, 1967; 1970). It is shown that for the cycling of P the system returns slowly to equilibrium after an initial disturbance. The time span of about 30 years that is required to reach equilibrium is in agreement with the field data. However, P accumulation in living above-ground biomass and litter is overestimated by the model in the first years after burning. Nitrogen accumulation in biomass and litter is systematically underestimated by the model.The results are discussed with reference to the processes in nutrient cycling that are insufficiently understood, and with reference to the management of heathlands.     

Indirect effects and traditional trophic cascades: a test involving wolves, coyotes, and pronghorn

The traditional trophic cascades model is based on consumer resource interactions at each link in a food chain. However, trophic-level interactions, such as mesocarnivore release resulting from intraguild predation, may also be important mediators of cascades. From September 2001 to August 2004, we used spatial and seasonal heterogeneity in wolf distribution and abundance in the southern Greater Yellowstone Ecosystem to evaluate whether mesopredator release of coyotes (Canis latrans), resulting from the extirpation of wolves (Canis lupus), accounts for high rates of coyote predation on pronghorn (Antilocapra americana) fawns observed in some areas. Results of this ecological perturbation in wolf densities, coyote densities, and pronghorn neonatal survival at wolf-free and wolf-abundant sites support the existence of a species-level trophic cascade. That wolves precipitated a trophic cascade was evidenced by fawn survival rates that were four-fold higher at sites used by wolves. A negative correlation between coyote and wolf densities supports the hypothesis that interspecific interactions between the two species facilitated the difference in fawn survival. Whereas densities of resident coyotes were similar between wolf-free and wolf-abundant sites, the abundance of transient coyotes was significantly lower in areas used by wolves. Thus, differential effects of wolves on solitary coyotes may be an important mechanism by which wolves limit coyote densities. Our results support the hypothesis that mesopredator release of coyotes contributes to high rates of coyote predation on pronghorn fawns, and demonstrate the importance of alternative food web pathways in structuring the dynamics of terrestrial systems.