Food web structure and trophic interactions of the tropical highland lake Hayq, Ethiopia

We generated a mass-balance model to figure out the food web structure and trophic interactions of the major functional groups of the Ethiopian highland Lake Hayq. Moreover, the study lay down a baseline data for future ecosystem-based investigations and management activities. Extensive data collection has been taken place between October 2007 and May 2009. Ecotrophic efficiency (EE) of several functional groups including phytoplankton (0.8) and detritus (0.85) was high indicating the utilization of the groups within the system. However, the EE of Mesocyclops (0.03) and Thermocyclops (0.30) was very low implying these resources were rather a ‘sink’ in the trophic hierarchy. Flows based on aggregated trophic level sensu Lindeman revealed the importance of both phytoplankton and detritus to higher trophic levels. The computed average transfer efficiency of 11.5% for the first four trophic levels was within the range for highly efficient African lakes. The primary production to respiration (P/R) ratio (1.05) of Lake Hayq indicates the maturity of the ecosystem. We also modeled the food-web by excluding Tilapia and reduced phytoplankton biomass to get insight into the mass balance before Tilapia was introduced. The analysis resulted in a lower system omnivory index (SOI = 0.016) and a reduced P/R ratio (0.13) that described the lake as immature ecosystem, suggesting the introduction of Tilapia might have contributed to the maturity of the lake. Tilapia in Lake Hayq filled an ecological empty niche of pelagic planktivores, and contributed for the better transfer efficiency observed from primary production to fish yield.     

Comparative network analysis toward characterization of systemic organization for human–environmental sustainability

A preliminary study in comparative ecological network analysis was conducted to identify key assumptions and methodological challenges, test initial hypotheses and explore systemic and network structural characteristics for environmentally sustainable ecosystems. A nitrogen network for the U.S. beef supply chain – a small sub-network of the industrial food system analyzed as a pilot study – was constructed and compared to four non-human carbon and nitrogen trophic networks for the Chesapeake Bay and the Florida Everglades. These non-human food webs served as sustainable reference systems. Contrary to the main original hypothesis, the “window of vitality” and the number of network roles did not clearly differentiate between a human sub-network and the more complete non-human networks. The effective trophic level of humans (a partial estimate of trophic level based on the single food source of beef) was much higher (8.1) than any non-human species (maximum of 4.88). Network connectance, entropy, total dependency coefficients, trophic efficiencies and the ascendency to capacity ratio also indicated differences that serve as hypotheses for future tests on more comprehensive human food webs. The study elucidated important issues related to (1) the steady state assumption, which is more problematic for industrial human systems, (2) the absence or dearth of data on contributions of dead humans and human wastes to feed other species in an integrated food web, (3) the ambiguity of defining some industrial compartments as living versus non-living, and (4) challenges with constructing compartments and trophic transfers in industrial versus non-human food webs. The two main novel results are (1) the progress made toward adapting ecological network analysis (ENA) methodology for analysis of human food networks in industrial cultures and (2) characterizing the critical aspects of comparative ENA for understanding potential causes of the problems, and providing avenues for solutions, for environmental sustainability. Based on this work, construction and comparative network analysis of a more comprehensive industrial human food network seems warranted and likely to provide valuable insights for modifying structures of industrial food networks to be more like natural networks and more sustainable.     

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.     

Comparing network analysis methodologies for consumer–resource relations at species and ecosystems scales

This research compares two existing methodologies, mixed trophic impact analysis and utility analysis, which use network analysis to evaluate the direct, pair-wise, and indirect, holistic, ecological relations between ecosystem compartments. The two approaches have many similarities, but differ in some key assumptions which affect both the final results and interpretations. Here, we briefly introduce both methodologies through a series of two simple examples; a 3-compartment competition model and a 3-compartment food chain model, and then apply the methodologies to a 15-compartment ecosystem model of the Chesapeake Bay. This example demonstrates how implementing the various conceptual and methodological assumptions lead to differing results. Notably, the overall number of positive relations is greatly affected by the treatment of the self-interactions and the handling of detritus compartments lead to a distinction between ecological or trophic relations. We recommend slight changes to both methodologies, not necessarily in order to bring them completely together, but because each has some points which are stronger and better defensible.     

Adding rigor to ecological network models by evaluating a set of pre-balance diagnostics: A plea for PREBAL

The widespread use of ecological network models (e.g., Ecopath, Econetwrk, and related energy budget models) has been laudable for several reasons, chief of which is providing an easy-to-use set of modeling tools that can present an ecosystem context for improved understanding and management of living marine resources (LMR). Yet the ease-of-use of these models has led to two challenges. First, the veritable explosion of the use and application of these network models has resulted in recognition that the content and use of such models has spanned a range of quality. Second, as these models and their application have become more widespread, they are increasingly being used in a LMR management context. Thus review panels and other evaluators of these models would benefit from a set of rigorous and standard criteria from which the basis for all network models and related applications for any given system (i.e., the initial, static energy budget) can be evaluated. To this end, as one suggestion for improving network models in general, here I propose a series of pre-balance (PREBAL) diagnostics. These PREBAL diagnostics can be done, now, in simple spreadsheets before any balancing or tuning is executed. Examples of these PREBAL diagnostics include biomasses, biomass ratios, vital rates, vital rate ratios, total production, and total removals (and slopes thereof) across the taxa and trophic levels in any given energy budget. I assert that there are some general ecological and fishery principles that can be used in conjunction with PREBAL diagnostics to identify issues of model structure and data quality before balancing and dynamic applications are executed. I humbly present this PREBAL information as a simple yet general approach that could be easily implemented, could be considered for further incorporation into these model packages, and as such would ultimately result in a straightforward way to evaluate (and perhaps identify areas for improving) initial conditions in food web modeling efforts.     

The effects of aggregation on the performance of the inverse method and indicators of network analysis

Food webs are usually aggregated into a manageable size for their interpretation and analysis. The aggregation of food web components in trophic or other guilds is often at the choice of the modeler as there is little guidance in the literature as to what biases might be introduced by aggregation decisions. We examined the impacts of the choice of the a priori model on the subsequent estimation of missing flows using the inverse method and on the indices derived from ecological network analysis of both inverse method-derived flows and on the actual values of flows, using the fully determined Sylt-Rømø Bight food web model. We used the inverse method, with the least squares minimization goal function, to estimate ‘missing’ values in the food web flows on 14 aggregation schemes varying in number of compartments and in methods of aggregation. The resultant flows were compared to known values; the performance of the inverse method improved with increasing number of compartments and with aggregation based on both habitat and feeding habits rather than diet similarity. Comparison of network analysis indices of inverse method-derived flows with that of actual flows and the original value for the unaggregated food web showed that the use of both the inverse method and the aggregation scheme affected indices derived from ecological network analysis. The inverse method tended to underestimate the size and complexity of food webs, while an aggregation scheme explained as much variability in some network indices as the difference between inverse-derived and actual flows. However, topological network indices tended to be most robust to both the method of determining flows and to the inverse method. These results suggest that a goal function other than minimization of flows should be used when applying the inverse method to food web models. Comparison of food web models should be done with extreme care when different methodologies are used to estimate unknown flows and to aggregate system components. However, we propose that indices such as relative ascendency and relative redundancy are most valuable for comparing ecosystem models constructed using different methodologies for determining missing flows or for aggregating system components.     

Equivalence of the realized input and output oriented indirect effects metrics in Ecological Network Analysis

A new understanding of the consequences of how ecosystem elements are interconnected is emerging from the development and application of Ecological Network Analysis. The relative importance of indirect effects is central to this understanding, and the ratio of indirect flow to direct flow (I/D) is one indicator of their importance. Two methods have been proposed for calculating this indicator. The unit approach shows what would happen if each system member had a unit input or output, while the realized technique determines the ratio using the observed system inputs or outputs. When using the unit method, the input oriented and output oriented ratios can be different, potentially leading to conflicting results. However, we show that the input and output oriented I/D ratios are identical using the realized method when the system is at steady state. This work is a step in the maturation of Ecological Network Analysis that will let it be more readily testable empirically and ultimately more useful for environmental assessment and management.     

On the consequences of aggregation and balancing of networks on system properties derived from ecological network analysis

In the ecological network analysis (ENA) of complex flow food webs the assumption is often made that the models characterizing the flows and stocks of ecosystems occur in a steady state where inflows equals outflows. An assessment of the system indices derived from ENA of six balanced and unbalanced system models, respectively, indicate to differences between indices. The aggregation of highly articulated flow models into models with fewer compartments also has drastic effects on the system metrics, particularly on the information indices.     

Modelling the effects of eutrophication, mitigation measures and an extreme flood event on estuarine benthic food webs

Human-mediated and natural disturbances such as nutrient enrichment, habitat modification, and flood events often result in significant shifts in species composition and abundance that translate into changes in the food web structure. Six mass-balanced models were developed using the “Ecopath with Ecosim” software package to assess changes in benthic food web properties in the Mondego estuarine ecosystem (Portugal). Field, laboratory and literature information were used to construct the models. The main study objective was to assess at 2 sites (a Zostera meadow and a bare sediment area) the effects of: (1) a period of anthropogenic enrichment, which led to excessive production of organic matter in the form of algal blooms (1993/1994); (2) the implementation of mitigation measures, following a long period of eutrophication (1999/2000); and (3) a centenary flood (winter 2000/2001). Different numbers of compartments were identified at each site and in each time period. In general, the Zostera site, due to its complex community, showed a higher number of compartments and a higher level of system activity (i.e. sum of consumptions, respiration, flow to detritus, production, total system throughput, net primary production and system omnivory index). The differences at the two sites in the three time periods in the breakdown of throughput were mainly due to differences in the biomass of the primary producers (higher primary production at the Zostera site). Consumption, respiration and flow to detritus were dominated by the grazers Hydrobia ulvae and Scrobicularia plana at the Zostera and bare sediment sites respectively. At both sites, after recovery measures were implemented there was an increase in S. plana and Hediste diversicolor biomass, consumption, respiration and flows to detritus, and a decrease in H. ulvae biomass and associated flows, which increased again after the flood event. The mass-balanced models showed that the trophic structure of the benthic communities in Mondego estuary was affected differently by each disturbance event. Interestingly, in our study a high system throughput seems to be associated with higher stress levels, which contradicts the idea that higher system activity is always a sign of healthier conditions.     

Ecological network determination of sectoral linkages, utility relations and structural characteristics on urban ecological economic system

Analyzing the structure and functioning of the urban system revealed ways to optimize its structure by adjusting the relationships among compartments, thereby demonstrating how ecological network analysis can be used in urban system research. Based on the account of the extended exergy utilization in the sector of urban socio-economic system, which is considered as the composition of extraction (Ex), conversion (Co), agriculture (Ag), industry (In), transportation (Tr), tertiary (Te) and households (Do) sectors, an urban ecological network model is constructed to gain insights into the economic processes oriented to sustainable urban development. Taking Beijing city as the case, the network accounting and related ecological evaluation of a practical urban economy are carried out in this study in the light of flux, efficiency, utility and structure analysis. The results showed that a large quantity of energy and resources have to be consumed to maintain the structure and function of a city. The thermodynamic efficiencies of individual sector in Beijing remain at a low level. The social system in Beijing is a highly competitive network, and there are 8 competitive relations and only two mutualistic ones. The Domestic and Agricultural sector are the major controlling factors of the system. Moreover, the assessment results of Beijing are compared with the other three socio-economic systems, Norway, UK and Italy, and the ecological network function and structure comparisons are correspondingly illuminated and discussed. The conclusions indicate that the exergy-based network analysis can be refined to become an integrative tool for evaluation, policy-making and regulation for urban socio-economic system management concerning structure and efficiency at urban levels.     

Response of balanced network models to large-scale perturbation: Implications for evaluating the role of small pelagics in the Gulf of Maine

Exploring the response of an ecosystem, and subsequent tradeoffs among its biological community, to human perturbations remains a key challenge for the implementation of an ecosystem approaches to fisheries (EAF). To address this and related issues, we developed two network (or energy budget) models, Ecopath and Econetwrk, for the Gulf of Maine ecosystem. These models included 31 network “nodes” or biomass state variables across a broad range of trophic levels, with the present emphasis to particularly elucidate the role of small pelagics. After initial network balancing, various perturbation scenarios were evaluated to explore how potential changes to different fish, fisheries and lower trophic levels can affect model outputs. Categorically across all scenarios and interpretations thereof, there was minimal change at the second trophic levels and most of the “rebalancing” after a perturbation occurred via alteration of the diet matrix. Yet the model results from perturbations to a balanced energy budget fall into one of three categories. First, some model results were intuitive and in obvious agreement with established ecological and fishing theory. Second, some model results were counter-intuitive upon initial observation, seemingly contradictory to known ecological and fishing theory; but upon further examination the results were explainable given the constraints of an equilibrium energy budget. Finally, some results were counter-intuitive and difficult to reconcile with theory or further examination of equilibrium constraints. A detailed accounting of biomass flows for example scenarios explores some of the non-intuitive results more rigorously. Collectively these results imply a need to carefully track biomass flows and results of any given perturbation and to critically evaluate the conditions under which a new equilibrium is obtained for these types of models, which has implications for dynamic simulations based off of them. Given these caveats, the role of small pelagics as a prominent component of this ecosystem remains a robust conclusion. We discuss how one might use this approach in the context of further developing an EAF, recognizing that a more holistic, integrated perspective will be required as we continue to evaluate tradeoffs among marine biological communities.     

An inverse model for the analysis of the Venice lagoon food web

A steady-state model of the Venice lagoon food web was constructed, based on a comprehensive set of data, which were collected in the years 2001-2005. Energy flows were estimated by means of an inverse methodology of constrained optimization based on the Minimum Norm criterion, i.e. on the minimization of both the sum of squares of the residuals and of the sum of squares of energy flows. The solution was constrained by a set inequalities, which were derived from general eco-physiological knowledge and site specific data on energy flows. The trophic network was represented by thirty-two nodes, including single-species compartments for the species of high economical or ecological relevance. Mass balance equations were weighted, in order to obtain meaningful results in presence of large differences, up to 5 orders of magnitude, among biomasses. A perturbation technique was applied, with the purpose of reducing the risk of finding solutions heavily affected by the set of constraints and of obtaining a more robust representation of the energy flows. The main patterns of energy flow are consistent with those obtained in previous attempts at modelling the Venice lagoon food web. Micro- and macro-phytobenthos account for the largest fraction of the primary production. Energy is then transferred towards higher trophic levels by means of two main pathways: the recycling of dead biomass through the detritus compartment and the direct consumption by grazers. The first pathway is the most important and accounts for approximately two/thirds of the energy transferred to the second trophic level.     

What is hidden behind the concept of ecosystem efficiency in energy transformation?

The number of energy transformation levels in trophic webs is usually below five, but can be extended up to ten when parasites and hyper-parasites are included. Research on the structure and function of food webs is relevant to the complexity–stability–productivity debate. The aim of this theoretical analysis is to link energetic and connectional aspects of ecosystems with information theory. Updating an energetic model reported by Ricklefs [Ecologia, Zanichelli Editore S.p.A., Bologna, Italy, 1993, p. 896], our approach is integrated with a static analysis of food webs. The length of food webs is theoretically associated with the average ecological efficiency which can be empirically correlated with the effective connectance between species. Furthermore, the advantage of greater complexity when applied to a signalling network is qualitatively addressed.The overall efficiency of energy transformation into biomass throughout a trophic web, in an ecosystem with a given number of species, is the resultant of the various ecological efficiencies, η, at the transitions between the trophic levels. However, we propose that an increment in effective connectance and interspecies connectivity based on a superimposed signalling web may increase the η values, despite the fact that signalling per se has an energetic cost. According to this hypothesis, ecosystem stability would not be necessarily reduced by increasing the number of trophic levels, N, whenever stability in terms of persistence is improved by a cost-efficient regulatory network.     

Ecological network and emergy analysis of urban metabolic systems: Model development,and a case study of four Chinese cities

Analysis of the structure and function of urban metabolic systems is an important goal of urban research. We used network pathways and network utility analysis to analyze the basic network structure of the urban metabolic system and the complex ecological relationships within the system, providing a new way to perform such research. Using four Chinese cities as examples, we developed an ecological network model of the urban metabolic system. By using network pathway analysis, we studied the changing relationships between metabolic length and the number of metabolic pathways, and between metabolic length and reachability. Based on the distribution of the number of metabolic pathways, we describe the basic structure and intercompartment relationships of the system. By using the sign distribution in the network utility matrix, we determined the ecological relationships and degree of mutualism between the compartments of the system. The basic components of the system consisted of the internal environment, the external environment, and the agricultural, industrial, and domestic sectors. With increasing metabolic length, the ecological relationships among the components of the system became more diverse, and the numbers of metabolic paths and their reachability improved. Although the basic network structure of the four cities was identical, the mutualism index differed. Beijing's mutualism index was superior to that of Shanghai, and much higher than those of Tianjin and Chongqing. By analyzing the structure and function of the urban metabolic system, we provide suggestions for optimizing the structure and adjusting the relationships, and propose methods for the application of ecological network analysis in future urban system research.     

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.     

Analyzing the tradeoffs among ecological and fishing effects on an example fish community: A multispecies (fisheries) production model

Species interactions such as predation or competition can have notable implications for fish population dynamics. Stock status and associated biological reference points can be misrepresented if these interactions are not considered. Here we present a multispecies, biomass production model (MS-PROD) developed by incorporating modifications to the Schaefer production model. The modifications explicitly emphasize a functional group approach, highlighting the effects of species interactions that include competition between species within a group, competition between groups, and predation. The mathematical formulation also includes constraints on carrying capacity for both the entire system and for individual groups. We applied the model as initialized to the Northeast U.S. Large Marine Ecosystem finfish community. We then executed simulations designed to explore the behavior of the model with respect to fishery exploitation and ecological interactions. Collectively, the results demonstrated the utility, and the need, for incorporating ecological effects into fisheries models.     

The Reynolds transport theorem: Application to ecological compartment modeling and case study of ecosystem energetics

The Reynolds transport theorem (RTT) from mathematics and engineering has a rich history of success in mass transport dynamics and traditional thermodynamics. This paper introduces RTT as a complementary approach to traditional compartmental methods used in ecological modeling and network analysis. A universal system equation for a generic flow quantity is developed into a generic open-system differential expression for conservation of energy. Nonadiabatic systems are defined and incorporated into control volume (CV) and control surface (CS) perspectives of RTT where reductive assumptions in empirical data are then formally introduced, reviewed, and appropriately implemented. Compartment models are abstract, time-dependent systems of simultaneous differential equations describing storage and flow of conservative quantities between interconnected entities (the compartments). As such, they represent a set of flexible and somewhat informal, assumptions, definitions, algebraic manipulations, and graphical depictions subject to influence and selectively parsed expression by the modeler. In comparison, RTT compartment models are more rigorous and formal integro-differential equations and graphics initiated by the RTT universal system equation, forcing an ordered identification of simplifying assumptions, ending with clearly identified depictions of the transfer and transport of conservative substances in physical space and time. They are less abstract in the rigor of their equation development leaving less ambiguity to modeler discretion. They achieve greater consistency with other RTT compartment style models while possibly generating greater conformity with physical reality. Characteristics of the RTT approach are compared with those of a traditional compartment model of energy flow in an intertidal oyster-reef community.     

Weak indirect effects inherent to nitrogen biogeochemical cycling within anthropogenic ecosystems: A network environ analysis

Indirect effects are assumed to be the major causes of the complexity and stability of ecological networks. The complexity of urban-rural complexes (URCs) could also be attributed to the indirect effects associated with human activities. No studies, however, have quantified the strength of indirect effects in relation to urban biogeochemistry. A network environ analysis (NEA) was used for this study to investigate and compare indirect effects in relation to the nitrogen (N) cycling networks of 22 natural ecosystems and five URCs. Results show that indirect effects were proven to be weak for URC N cycling networks (accounting for only ∼2% of the overall effects measured in natural ecosystems). The weak indirect effects found provide a counterexample for the hypothesis that indirect effects are in fact the dominant components of biogeochemical networks. It also implies that human activity in itself does not always raise the complexity of ecological processes as previously suggested. Weak indirect effects also lead to perturbation fragility for URC N cycles (where the decay rate is greater in comparison to natural ecosystems by a factor of 13). In order to improve the robustness and efficiency of URC biogeochemical cycling, a knockout analysis was carried out. By comparing results after removing single interactions between natural ecosystems and URCs it was found that the loss of indirect effects require cooperative strategies to optimize N cycling networks within URCs.