Evaluation of ecological network analysis: Validation of output

Ecological network analysis (ENA) is a modeling approach increasingly being used to evaluate food webs and provide an ecosystem-based approach to resource management. Unfortunately, validation of ENA output is rarely performed. This study represents part of a larger effort to critically evaluate ENA. Here we validate ENA output using stable isotope analysis (SIA), and where validation is not met, determine the effects of modifying trophic networks to reflect validation.     

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.     

Soft intervention technology as a tool for integrated coastal zone management

After introducing soft defence techniques as an alternative to hard defence techniques, the need is emphasized to consider the coastal area as an integral system. By recalling the main driving factors for coastal management: conflict resolution, resilience and sustainability, we logically arrive at the concepts of ecological engineering and ecotechnology, which are increasingly acknowledged as possible solutions to achieve sustainable use of coastal space as a resource. In this context, we refer to the principles of self design and of ecosystem conservation. In order to deal with real situations we are in need of fundamental ‘tools’ for the application of the soft intervention technology approach. We therefore introduce the concept of physiographic units and develop an initial elaboration for a coastal stretch and for coastal wetlands. The latter deserve more attention because of the already established practices of ecotechnology, at least as far as water and soil quality are concerned, but certainly also concerning morphology, especially in the future. We conclude by briefly discussing how activities undertaken in two research projects currently being conducted under the framework of the Marine Science and Technology Program of the Commission of the European Communities are expected to contribute to the concepts introduced here.     

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.     

Do mucilage events influence pico- and nanoplankton size and structure in the Adriatic sea?

Temporal changes in abundance and biomass of picophytoplankton, heterotrophic pico-eukaryotes, and nanoplankton assemblages were investigated along a transect crossing the Adriatic Sea, from the Italian to the Croatian coast. This 15-months (June 1999-August 2000) investigation allowed comparing microbial parameters during summer 1999 (year without mucilage) and summer 2000 when a major mucilage event occurred. Pico- and nanoplankton assemblages displayed significant differences between the 2 summer periods. The main differences can be summarized as: (i) presence of cyanobacterial blooms (up to 10⁸ cells l^-1) in summer 2000, absent in summer 1999; (ii) an increasing fraction of heterotrophic pico-eukaryotes (up to 5.0 × 10⁶ cells l^-1) and heterotrophic nanoplankton (size 2-5 µm) during mucilage event; (iii) a reduced abundance of small-sized (2-3 µm) phototrophic nanoplankton in summer 2000. Changes in community structure were signals of changes in trophic condition of the system, which resulted in a competitive advantage for small sized pico- and nanoheterotrophs. Data presented here indicated that mucilage events are associated with changes in microbial community structure and functioning in ambient water and induced the amplification of 3-step microbial food chain. The potential use of the heterotrophic pico-eukaryotes for describing alterations of the trophic pathways during mucilage events is discussed.     

An interregional ecological approach for modelling sustainability in a globalizing world—Reviewing existing approaches and emerging directions

In recent decades international trade has become a major source of supplying the need and wants of billions of people around the world. Virtually everyone now consumes resource commodities and manufactured products imported from ‘elsewhere’. In effect, globalization and trade enable consuming populations to support themselves on the output of distant ecosystems half a world away. However, while economic integration implies greater ‘connectivity’ within the global village, the spatial separation of material production (including resource extraction) from consumption eliminates some of the signals i.e., the negative feedbacks coming from supporting eco-systems from reaching those who depend on these ecosystems for their sustainability. At present, despite increasing global connectedness, most environmental studies and models apply to a single spatial scale: local, national or global; analysing diverse pressures on human well-being and ecosystems integrity. This paper argues that both economic globalization and global ecological change should force us to add an interregional scale for quantifying and modelling sustainability. Such an approach recognizes that, in a globalizing world, the sustainability of any given region increasingly depends, directly and indirectly, on the sustainability of many other regions. The following pages describe the interregional approach and illustrate some existing and emerging methods for quantifying, analysing and modelling interregional linkages. It then identifies some of what is still missing, and discusses some of the implications in a changing world.     

Cycling in ecosystems: An individual based approach

Cycling index is an important ecological indicator used in ecosystem analysis. The higher the cycling in an ecosystem, the higher the utilization of mass and energy within the system before it is lost due to respiration and other factors. For a stock-flow type ecosystem model at steady state, Finn’s cycling index (FCI) can be computed using simple matrix algebra. However, it is difficult to measure how well this index represents the actual cycling occurring in the system. In this paper, we study cycling in ecological networks using an individual based approach (particle tracking algorithm). This new simulation method provides access to the pathway data of individual particles that flow in the system, therefore one can quantify cycling using this pathway data quite literally. We used particle tracking simulations (PTS) to compute a cycling index using Finn’s idea of flux based cycling. Our simulation based results (using no matrix algebra) agree with Finn’s cycling index, verifying the accuracy of both the PTS, and the original linear algebraic formulation of FCI.     

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.     

多溴联苯醚在典型电子垃圾污染区域水生食物链上的生物富集特征

为调查电子垃圾回收活动对当地水生生物造成的多溴联苯醚(PBDEs)污染,测定了广东某电子垃圾回收地附近水库中水生生物样品(水蛇、鱼类、草虾和田螺)和对照区样品(鲮鱼)中18种PBDEs含量,通过对样品氮同位素的测定,探讨了PBDEs在该地淡水食物链上的生物富集特征.研究结果显示,∑PBDEs(总PBDEs含量)在水蛇、鱼类、草虾和田螺中分别为39.6～186、1.82～75.7、4.60～17.1和7.26～17.0μg·g-1(脂重),比华南地区鱼体中∑PBDEs的平均含量高3个数量级,表明电子垃圾回收活动已对当地水生生物造成了PBDEs的严重污染.除BDE66、BDE99、BDE153、BDE183和BDE209外,其他13种PBDE单体在食物链上均存在着生物放大效应,但生物放大能力(B值)与正辛醇-水分配系数logKOW相关性不明显,可能是由于某些PBDE单体在生物体内存在代谢作用.以上结果表明,由电子垃圾回收活动导致的PBDEs污染存在较大的生态风险,应该引起高度关注.     

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.     

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

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

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.     

Throughflow analysis: A stochastic approach

Ecological network analysis (ENA), predicated on systems theory and Leontiev input–output analysis, is a method widely used in ecology to reveal ecosystem properties. An important ecosystem property computed in ENA is throughflows, the amount of matter/energy leaving each compartment of the ecosystem. Throughflows are analyzed via a matrix representing their relationships to the driving input at the boundary. Network particle tracking (NPT) builds on ENA to offer a Lagrangian particle method that describes the activity of the ecosystem at the microscopic level. This paper introduces a Lagrangian throughflow analysis methodology using NPT and shows that the NPT throughflow matrix, , agrees with the conventional ENA throughflow matrix, , for ecosystems at steady-state with donor-controlled flows. The matrix is computed solely from the pathways (particles’ histories) generated by NPT simulations and its average over multiple runs of the algorithm with longer simulation time agrees with the Eulerian matrix (Law of Large Numbers). While the traditional NEA throughflow analysis is mostly used with steady-state ecosystem models, the Lagrangian throughflow analysis that we propose can be used with non-steady-state models and paves the way for the development of dynamic throughflow analysis.     

How deep is the footprint? A 3D representation

Depletion of natural capital stocks and use of natural capital flows are the central issues in the sustainability debate. Differentiation of these two components, considering natural capital and its limits, is important for planning and management of land use. This paper offers insights into this issue and proposes a new perspective of the Ecological Footprint (EF) in three dimensions, considering not only its size but also its depth: according to this viewpoint the footprint is not an area but a volume, although maintaining the same value as the one in two dimensions. Use of annual flows provided by the Earth is represented by the footprint size, expressed in global hectares (gha) and plotted in the (x,y) plane. Footprint depth represents the demand for extra land to meet human needs through depletion of stocks of natural capital. It is plotted on the z-axis. It can be seen as the number of years necessary to regenerate resources liquidated in 1 year (and to absorb the respective wastes) or as the number of planets necessary to support the inhabitants of the planet Earth. The evolution of these two components in the last five decades is studied and discussed.     

An inexact-stochastic with recourse model for developing regional economic-ecological sustainability under uncertainty

Effective planning of resources management is important for facilitating socio-economic development and eco-environmental sustainability. Such a planning effort is complicated with a variety of uncertain, dynamic and nonlinear factors as well as their interactions. In this study, an inexact-stochastic quadratic programming with recourse (ISQP-R) method is developed for reflecting dynamics of system uncertainties based on a complete set of scenarios as well as tackling nonlinearities in the objective function to reflect the effects of marginal utility on system benefits and costs. Moreover, since penalties are exercised with recourse against any infeasibility, the ISQP-R can support the analysis of various policy scenarios that are associated with different levels of economic consequences when the promised targets are violated. The developed method is applied to a case study of planning resources management and developing regional ecological sustainability. The results have been generated and are helpful for decision makers in not only identifying desired resources-allocation strategies but also gaining insight into the tradeoff between economic objective and eco-environment violation risk.     

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.     

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

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