Simulation modeling of material and energy flow through an ecosystem: Methods and documentation

This paper discusses the process of development of large-scale ecological simulation models as it proceeds through steps divided between two major phases — conceptualization and implementation. The activities in these phases are strongly influenced by model objectives. Activities transpiring in each step are discussed and documentation methods both for the modelling process and its product are introduced for each of the steps. Examples are given from CONIFER, a large-scale coniferous forest model developed by the author in cooperation with another investigator. The role of the model documentation, both in aiding model development and in clearly communicating and allowing criticism of the model, is emphasized. Model evaluation, checking that the model satisfies its objectives, is seen as an area of model development which indicates which parts of the model need reworking.     

荒漠河岸生态系统退化机理的定量分析

通过野外监测资料,采用聚类分析与主成分分析法,分析了荒漠河岸生态系统退化的内外因素与驱动力、植被退化的环境主导因子与生态退化程度,结果表明：（1）地质、地貌与气候条件等因素的影响,塔里木河下游生态系统存在脆弱和不稳定性是其退化的内因,外界干扰则是其退化的外在因素。人为干扰则是其退化发生的驱动力,起因于人口的增加、需求的增长。（2）在此过程中导致植被受损的环境主导因子是水,植被退化是人类干扰作用于植被赖以生存的环境主导因子所致。（3）其不同河段的生态退化可以归为三类：潜在沙漠化类（轻度退化）,轻度沙漠化类（中度退化）,中度或重度沙漠化类（重度退化）,其中第三类型,土壤条件与其它两类差异较大,表现出土壤退化的滞后现象。     

荒漠河岸生态系统退化机理的定量分析

通过野外监测资料,采用聚类分析与主成分分析法,分析了荒漠河岸生态系统退化的内外因素与驱动力、植被退化的环境主导因子与生态退化程度,结果表明:(1)地质、地貌与气候条件等因素的影响,塔里木河下游生态系统存在脆弱和不稳定性是其退化的内因,外界干扰则是其退化的外在因素。人为干扰则是其退化发生的驱动力,起因于人口的增加、需求的增长。(2)在此过程中导致植被受损的环境主导因子是水,植被退化是人类干扰作用于植被赖以生存的环境主导因子所致。(3)其不同河段的生态退化可以归为三类:潜在沙漠化类(轻度退化),轻度沙漠化类(中度退化),中度或重度沙漠化类(重度退化),其中第三类型,土壤条件与其它两类差异较大,表现出土壤退化的滞后现象。     

An invasive riparian tree reduces stream ecosystem efficiency via a recalcitrant organic matter subsidy

A disturbance, such as species invasion, can alter the exchange of materials and organisms between ecosystems, with potential consequences for the function of both ecosystems. Russian olive (Elaeagnus angustifolia) is an exotic tree invading riparian corridors in the western United States, and may alter stream organic matter budgets by increasing allochthonous litter and by reducing light via shading, in turn decreasing in-stream primary production. We used a before-after invasion comparison spanning 35 years to show that Russian olive invasion increased allochthonous litter nearly 25-fold to an invaded vs. a control reach of a stream, and we found that this litter decayed more slowly than native willow. Despite a mean 50% increase in canopy cover by Russian olive and associated shading, there were no significant changes in gross primary production. Benthic organic matter storage increased fourfold after Russian olive invasion compared to pre-invasion conditions, but there were no associated changes in stream ecosystem respiration or organic matter export. Thus, estimated stream ecosystem efficiency (ratio of ecosystem respiration to organic matter input) decreased 14%. These findings show that invasions of nonnative plant species in terrestrial habitats can alter resource fluxes to streams with consequences for whole-ecosystem functions.     

Benthic energy dynamics in a southern Baltic ecosystem

Benthic components and metabolic activity at two stations in the Darss-Zingst estuary (eastern German Baltic coast) were investigated over a seasonal cycle from April 1985 to August 1986. As has been established for temperate and boreal ecosystems, peaks in the biomass of benthic microphytes occurred in the spring and late autumn to winter, presumably caused by settling phytoplankton blooms. Metabolic activity of the benthos did not increase with rising ambient temperatures. Rather, the highest values of oxygen consumption were recorded during the cooler months (spring and winter), when increased numbers of organisms were also observed. This may be a response to a greater food supply to the sediment in the form of settling phytoplankton during these times of year.     

The effects of parasitism on energy flow through laboratory shrimp populations

I quantified the effects of parasitism by the isopod Probopyrus pandalicola on energy flow through the host Palaemonetes pugio by comparing secondary production, metabolism, ingestion, and egestion by unparasitized laboratory shrimp populations to the same parameters for parasitized groups during 10 months. The effects of parasitism on host growth and metabolism vary from month to month. Temperature, season, host age, sex, and reproductive condition affect energetics for host-parasite systems. Probopyrus pandalicola has little effect on host assimilation efficiency. However, tissue growth efficiences during most study months were higher for control shrimp than parasitized shrimp. These differences between groups were of lesser magnitude when parasite production was considered in the calculations. Trophic level energy intake efficiency for parasites was of the order of 6 to 10% throughout much of the study—the highest values were calculated during the parasites' reproductive months. Through parasitic castration, P. pandalicola significantly affects host energetics. Significantly, parasite reproduction was often of the same magnitude as reproduction by unparasitized hosts, although parasite biomass accounts for only about 4% of the total host-parasite system biomass.     

A Markovian model for ecosystem flow analysis

Nutrient flow through ecosystems is modeled as a discrete Markov chain whose transition probabilities are stationary or time inhomogenous according to whether a steady state or dynamic ecosystem is modeled, respectively. Expected residence time and number of intercompartmental transfers for a nutrient within a set of compartments are derived. Variances of these random variables are also considered. A measure for ecosystem resource recycling is given as a weighted sum of probabilities.     

Simulation model of energy flow through an animal organism: General form and applications

This paper presents a general model for energy flow through an animal organism whose consumption and assimilation processes can be described by continuous functions. Two versions of the model are discussed: with continuous and with discrete reproduction. Also, their applicability in analyzing the properties of an actual modelled system is discussed. The results of sensitivity analysis and optimization considerations are presented.     

A protocol for eliciting nonmaterial values through a cultural ecosystem services frame

Stakeholders’ nonmaterial desires, needs, and values often critically influence the success of conservation projects. These considerations are challenging to articulate and characterize, resulting in their limited uptake in management and policy. We devised an interview protocol designed to enhance understanding of cultural ecosystem services (CES). The protocol begins with discussion of ecosystem‐related activities (e.g., recreation, hunting) and management and then addresses CES, prompting for values encompassing concepts identified in the Millennium Ecosystem Assessment (2005) and explored in other CES research. We piloted the protocol in Hawaii and British Columbia. In each location, we interviewed 30 individuals from diverse backgrounds. We analyzed results from the 2 locations to determine the effectiveness of the interview protocol in elucidating nonmaterial values. The qualitative and spatial components of the protocol helped characterize cultural, social, and ethical values associated with ecosystems in multiple ways. Maps and situational, or vignette‐like, questions helped respondents articulate difficult‐to‐discuss values. Open‐ended prompts allowed respondents to express a diversity of ecosystem‐related values and proved sufficiently flexible for interviewees to communicate values for which the protocol did not explicitly probe. Finally, the results suggest that certain values, those mentioned frequently throughout the interview, are particularly salient for particular populations. The protocol can provide efficient, contextual, and place‐based data on the importance of particular ecosystem attributes for human well‐being. Qualitative data are complementary to quantitative and spatial assessments in the comprehensive representation of people's values pertaining to ecosystems, and this protocol may assist in incorporating values frequently overlooked in decision making processes.     

Stream ecosystem functioning under reduced flow conditions.

Assessments of flow reduction in streams often focus on changes to biological communities and in-stream physical characteristics, with little consideration for changes in ecosystem functioning. It is unclear whether functional indicators of ecosystem condition may be useful for assessing the impacts of reduced discharge on small streams. Using weirs and diversions to reduce stream discharge during summer baseflow conditions, we tested the response of leaf breakdown, coarse particulate organic matter (CPOM) retention, and primary production to one month of water abstraction in before-after, control-impact (BACI) designed experiments. Discharge at impact (downstream) reaches decreased by over 85% in each of three small New Zealand streams compared to controls (upstream). There also were decreases in velocity, depth, and wetted width. Sediment cover increased at impact reaches, but there were only small changes to conductivity, pH, and surface water temperature. We installed mesh bags filled with willow leaves in-stream for one month to measure leaf breakdown. Reduced discharge had little influence on leaf breakdown rate in these streams. Travel distances and retention structures for CPOM were evaluated using releases of paper strips and wooden dowelling over a range of discharges. The distance traveled by released CPOM increased with increasing discharge, and the importance of riffles as retention structures increased at lower discharges. We measured the accumulation of chlorophyll a after one month on artificial substrates as an estimate of the relative primary production of control and impact reaches. The differences in chlorophyll a concentrations between control and impact reaches were inconsistent among streams. These ecosystem functions have responded inconsistently to water removal in these streams. However, the strong response of CPOM retention to reduced discharge could complement measures of biological community structure when the influence of reduced discharge is assessed. We recommend further investigation in a wide range of streams to assess the utility of these processes as functional indicators of reduced discharge.     

A retrospective Markovian model for ecosystem resource flow

Historic ecosystem resource flow is modeled as a retrospective discrete Markov chain to obtain the expected values and variances of compartmental residence times and numbers of intercompartmental transfers that compartmental standing crops have experienced since their latest entry into the ecosystem. The transition probabilities of the proposed retrospective Markovian model are either stationary or time inhomogeneous according to whether a steady state or non-steady state ecosystem, respectively, is analyzed.     

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.     

Exploitation and destabilization of a warm, freshwater ecosystem through engineered hydrological change

Exploitation of freshwater resources is having catastrophic effects on the ecological dynamics, stability, and quality of those water resources on a global scale, especially in arid and semiarid regions. Lake Kinneret, Israel (the Biblical Sea of Galilee), the only major natural freshwater lake in the Middle East, has been transformed functionally into a reservoir over the course of approximately 70 years of hydrological alterations aimed mostly at producing electrical power and increasing domestic and agricultural water supply. Historical changes in lake chemistry and biology were reconstructed using analysis of sedimentary nutrient content, stable and radioisotope composition, biochemical and morphological fossils from algae, remains of aquatic invertebrates, and chemical indices of past light regimes. Together, these paleolimnological analyses of the lake's bottom sediments revealed that this transformation has been accompanied by acceleration in the rate of eutrophication, as indicated by increased accumulation rates of phosphorus, nitrogen, organic matter, phytoplankton and bacterial pigments, and remains of phytoplankton and zooplankton. Substantial increases in these indices of eutrophication coincide with periods of increased water-level fluctuations and drainage of a major upstream wetland in the early to middle 20th century and suggest that management of the lake for increased water supply has degraded water quality to the point that ecosystem stability and sustainability are threatened. Such destabilization may be a model for eutrophication of freshwater lakes in other arid regions of the world in which management emphasizes water quantity over quality.     

Effects of land use,cover, and protection on stream and riparian ecosystem services and biodiversity

Protected areas are an important part of broader landscapes that are often used to preserve biodiversity or natural features. Some argue that protected areas may also help ensure provision of ecosystem services. However, the effect of protection on ecosystem services and whether protection affects the provision of ecosystem services is known only for a few services in a few types of landscapes. We sought to fill this gap by investigating the effect of watershed protection status and land use and land cover on biodiversity and the provision of ecosystem services. We compared the ecosystem services provided in and around streams in 4 watershed types: International Union for Conservation of Nature category II protected forests, unprotected forests, unprotected forests with recent timber harvesting, and unprotected areas with agriculture. We surveyed 28 streams distributed across these watershed types in Quebec, Canada, to quantify provisioning of clean water, carbon storage, recreation, wild foods, habitat quality, and terrestrial and aquatic biodiversity richness and abundance. The quantity and quality of ecosystem services and biodiversity were generally higher in sites with intact forest—whether protected or not—relative to those embedded in production landscapes with forestry or agriculture. Clean-water provision, carbon storage, habitat quality, and tree diversity were significantly higher in and around streams surrounded by forest. Recreation, wild foods, and aquatic biodiversity did not vary among watershed types. Although some services can be provided by both protected and unprotected areas, protection status may help secure the continued supply of services sensitive to changes in land use or land cover. Our findings provide needed information about the ecosystem service and biodiversity trade-offs and synergies that result from developing a watershed or from protecting it.     

Framing the ecosystem concept through a longitudinal study of developments in science and policy

This paper examines how scientific literature and policy documents frame the ecosystem concept and how these frames have shaped scientific dialogue and policy making over time. This was achieved by developing a frame typology, as a basis for organizing relevant value expressions, to assess how different frames have altered perspectives of the ecosystem concept. The frame typology and analysis is based on a semi‐grounded and longitudinal document analysis of scientific literature and policy documents using the ecosystem concept. Despite changing discourses and public priorities (e.g., cultural constructs of biodiversity) both science and policy documents are characterized by stable value systems that have not changed substantially since the 1930s. These value systems were defined based on ethical principles that delineate 6 core frames: humans first, dual systems, eco‐science, eco‐holism, animals first, and multicentrism. Specific crises (e.g., climate change) and cross‐disciplinary uptake and re‐uptake of, for example, the ecosystem services concept, have brought new perspectives to the forefront of public discourse. These developments triggered changes in the core frames that, rather than being value based, are based on how the ecosystem is conceptualized under fixed value systems and over time. Fourteen subframes were developed to reflect these longitudinal changes. There are as such clear framing effects in both scientific literature and in policy. Ecosystem research is for instance often characterized by unstated value judgments even though the scientific community does not make these explicit. In contrast, policy documents are characterized by clear value expressions but are principally management driven and human centered.     

Identifying forest ecosystem services through socio-ecological bundles: a case study from northern Jordan

This study aims to identify services provided by forest ecosystems based on locals’ perceptions in the northern part of Jordan. By better understanding preferences of locals and by understanding why they value certain services more heavily, policy-makers and planning managers can make more effective decisions regarding development and conservation. Three hundred respondents were interviewed in order to collect information about forest ecosystem services (ES). Data collection was conducted using a structured questionnaire regarding ES provided by three forest types situated in northern Jordan. ES trade-offs and socio-ecological bundles were identified by analyzing respondents’ socioeconomic demographics and preferences of forest ES through multivariate canonical corresponding analysis (CCA). The statistical analysis indicated that the socioeconomic factors and forest type have an effect on social preferences toward ES. Results displayed a clear trade-off between provisioning services and regulating and cultural services. CCA demonstrated that 73% of the variation in ES value is explained by social factors (i.e. education level, income level, and gender), while 26% of perception variation was attributable to categories of ES supplied by each forest ecosystem. These findings imply that involving people in the place-specific management of public forests using the ES approach gives managers a clearer understanding of the benefits people recognize and value, as well as those they either are not aware of or do not value. Such information is useful in forest management and in public outreach. Although direct policy applications are limited by the research’s nature, the paper provides a starting point for incorporating forest users’ voices into policy discussions and management design.     

Energy flow through the benthos in a Scottish Sea Loch

Analysis of material collected in settlement traps at regular intervals over a period of a year gave an estimate of the annual organic input to the bottom sediment of about 28 gC/m² year, which is a little less than one third of the primary production in the overlying water column. The aerobic benthic community metabolism, estimated from in situ respiration measurements, was not significantly different from the carbon input. The rate of release of ammonia from the sediment was also measured in situ and would be sufficient to supply the greater part of the required input of inorganic nitrogen for photosynthesis in the water column.     

A sensitivity analysis of an ecosystem model of estuarine carbon flow

The North Inlet Marsh-Estuarine System Model (NIMES) is a 19-compartment real-time deterministic ecosystem simulation model of intrasystem carbon flow and exchange between an estuary and adjacent coastal water. A complete sensitivity analysis of this model with regard to POM, DOM and nekton annual exchange and annual system net productivity was completed and the functional relationship between these system behaviors and the perturbed parameters were determined by regression techniques. Simulated POM annual exchange between the estuary and the sea was largely controlled by offshore POM concentration, water column respiration and the gross productivity of the marsh and water column flora. Simulated DOM annual estuarine-oceanic exchange was most sensitive to perturbations in the gross productivity and biomass changes in marsh flora and water column microbial DOM uptake. Simulated nekton exchange reflected a sensitivity to migratory behavior and subtidal benthic biomass changes. System annual net productivity as simulated by the model showed a high sensitivity to all model processes which affected component primary production and respiration. From this sensitivity analysis, a scheme is developed to evaluate research needs for further model development for the North Inlet ecosystem.     

Phosphorus flow analysis of the socioeconomic ecosystem of Shucheng County, China

Human activities disturb the long-term phosphorus (P) cycle in nature, whereby the resulting intensive release of P contributes to the eutrophication of surface water. Hence, a detailed understanding of P flow as it relates to socioeconomic systems is essential for effective nutrient management. This study develops a substance-flow-analysis model for P metabolism for the socioeconomic ecosystem of Shucheng County in Anhui Province in central China as a case study. We estimate P flow using data from questionnaires, face-to-face interviews, published literature, and official statistical databases. Our results show that P flow in Shucheng's current socioeconomic system is linear and openly metabolic. The total P input into Shucheng in 2008 reached 12 748 Mg, mainly as P ores and crops. In all, 43.83% of the total P input was exported, and 30.44% was discharged into surface water. More-balanced nutrient management and options for improving nutrient use efficiency are discussed. The quantifiable, science-based methods used in this study may be applied to neighboring regions of central China for sustainable development and water management.     

The Coleroon river flow and its effect on the Pichavaram mangrove ecosystem

The fresh water reaching an estuary fluctuates on many scales and the fluctuation in the flow over a time period is also due to the anthropogenic activities like construction of dams, diversions and upstream withdrawals. These fluctuations may have profound effects on the estuarine ecosystem, which usually has remarkable biological productivity and diversity. A desktop analysis is carried out by compiling historical records of discharge and other hydrological information to study the dynamics of the river Coleroon, a distributary of the Cauvery River, Tamil Nadu, India. Remote sensing analysis was carried out on images ranging from the years 1977 to 2008 and the changes in the Pichavaram mangrove areas adjoining the Coleroon were studied by comparing the results with the freshwater flow into the ecosystem. Environmental flow analysis indicates that the minimum flow needed for a healthy mangrove system in Pichavaram is 750.75 cumecs; however, this is currently achieved only 12 % of the time. The reduction in freshwater flow over the years has resulted in loss of species diversity and degradation of the mangroves; this needs to be corrected immediately to conserve the Pichavaram mangrove ecosystem.