Underappreciated species in ecology: "ugly fish" in the northwest Atlantic Ocean.

Species shifts and replacements are common in ecological studies. Observations thereof serve as the impetus for many ecological endeavors. Many of the species now known to dominate ecosystem functioning were largely ignored until studies of those underappreciated species elucidated their critical roles. Recognizing the potential importance of underappreciated species has implications for functional redundancies in ecosystems and should alter our approach to long-term monitoring. One example of an applied ecological system containing species shifts, underappreciated species, and potential changes in functional redundancies is the topic of fisheries. The demersal component of many fish communities usually consists of high-profile and commercially valuable species that are targets of fisheries, plus a diverse group of lesser known species that have minimal commercial value and focus. Yet ecologically these traditionally nontargeted species are often a major biomass sink in marine ecosystems and can also be critical in the functioning of bentho-demersal food webs. I examined the biomass trajectories of several species of skates, cottids, lophiids, anarhichadids, zooarcids, and similar species in the northeast U.S. Atlantic ecosystem to determine whether their relative abundance has changed across the past four decades. Distribution and stomach contents of these species were also evaluated over time to further elucidate the relative importance of these species. Landings of these underappreciated bentho-demersal fish were also examined in comparison to those species that historically have been commercially targeted. Of particular emphasis was the evaluation of evidence for sequential stock depletion and the ramifications for functional redundancy for this ecosystem. Results indicate that some of these fish species are now the dominant piscivores, benthivores, and scavengers in this ecosystem. These formerly under-studied species generally have either maintained a consistent population size or have increased in abundance (and expanded in distribution) over the past several decades. Nontraditionally targeted fish species are an often overlooked but important component of bentho-demersal fish communities. Implications for the energy flow and resilience specifically for future fisheries and generally for harvesting biological resources are significant, remaining critical issues for the world's ecosystems.     

Establishing IUCN Red List Criteria for Threatened Ecosystems

Abstract: The potential for conservation of individual species has been greatly advanced by the International Union for Conservation of Nature's (IUCN) development of objective, repeatable, and transparent criteria for assessing extinction risk that explicitly separate risk assessment from priority setting. At the IV World Conservation Congress in 2008, the process began to develop and implement comparable global standards for ecosystems. A working group established by the IUCN has begun formulating a system of quantitative categories and criteria, analogous to those used for species, for assigning levels of threat to ecosystems at local, regional, and global levels. A final system will require definitions of ecosystems; quantification of ecosystem status; identification of the stages of degradation and loss of ecosystems; proxy measures of risk (criteria); classification thresholds for these criteria; and standardized methods for performing assessments. The system will need to reflect the degree and rate of change in an ecosystem's extent, composition, structure, and function, and have its conceptual roots in ecological theory and empirical research. On the basis of these requirements and the hypothesis that ecosystem risk is a function of the risk of its component species, we propose a set of four criteria: recent declines in distribution or ecological function, historical total loss in distribution or ecological function, small distribution combined with decline, or very small distribution. Most work has focused on terrestrial ecosystems, but comparable thresholds and criteria for freshwater and marine ecosystems are also needed. These are the first steps in an international consultation process that will lead to a unified proposal to be presented at the next World Conservation Congress in 2012.     

Prioritizing Conservation Effort through the Use of Biological Soil Crusts as Ecosystem Function Indicators in an Arid Region

Abstract: Conservation prioritization usually focuses on conservation of rare species or biodiversity, rather than ecological processes. This is partially due to a lack of informative indicators of ecosystem function. Biological soil crusts (BSCs) trap and retain soil and water resources in arid ecosystems and function as major carbon and nitrogen fixers; thus, they may be informative indicators of ecosystem function. We created spatial models of multiple indicators of the diversity and function of BSCs (species richness, evenness, functional diversity, functional redundancy, number of rare species, number of habitat specialists, nitrogen and carbon fixation indices, soil stabilization, and surface roughening) for the 800,000‐ha Grand Staircase‐Escalante National Monument (Utah, U.S.A.). We then combined the indicators into a single BSC function map and a single BSC biodiversity map (2 alternative types of conservation value) with an unweighted averaging procedure and a weighted procedure derived from validations performance. We also modeled potential degradation with data from a rangeland assessment survey. To determine which areas on the landscape were the highest conservation priorities, we overlaid the function‐ and diversity‐based conservation‐value layers on the potential degradation layer. Different methods for ascribing conservation‐value and conservation‐priority layers all yielded strikingly similar results (r= 0.89–0.99), which suggests that in this case biodiversity and function can be conserved simultaneously. We believe BSCs can be used as indicators of ecosystem function in concert with other indicators (such as plant‐community properties) and that such information can be used to prioritize conservation effort in drylands.     

Developing a Monitoring Program for Invertebrates: Guidelines and a Case Study

Abstract:  Invertebrates provide the majority of ecosystem services; thus, it is important that they be inventoried, monitored, and protected. Nevertheless, inventories, monitoring, and management generally focus on vertebrates and flowering plants. Consequently, there are few guidelines or case studies for invertebrates. We present a procedure for developing a monitoring program for species-rich invertebrates that entails (1) characterizing the community; (2) identifying surrogates for biodiversity; and (3) establishing efficient methods to monitor surrogates and any ecologically important or sensitive taxa. We used these procedures, biodiversity-based statistical advances, and a survey of arthropods to develop a monitoring plan for the forests of Shenandoah National Park, Virginia (U.S.A.). Our case study revealed that mixed hardwood and hemlock forests had significantly different compositions of arthropods in their soil and understory strata. Of the 10 orders tested Coleoptera and Hymenoptera were the only two to pass most of the five surrogate tests, and their combination improved predictions of overall arthropod diversity. Because arthropods represent the majority of macroscopic species in most ecosystems, the ability of this assemblage to predict overall arthropod diversity makes it a powerful surrogate. Of the 11 collecting methods used, the beat-sheet method was the most efficient for monitoring this surrogate assemblage. To complement this coarse-filter approach to monitoring at-risk, invasive, or other important taxa (fine filter), we used ordination analyses to match 66 taxa with the methods that most effectively sampled them. Our methods serve as a model for developing an invertebrate monitoring plan and should facilitate linking such monitoring with ecosystem functions and management.     

流域水生态功能区划及其关键问题

作为流域生态系统管理和水资源保护的重要手段,如何科学合理地开展流域水生态功能区划,已成为世界各国可持续发展所面临的关键挑战之一.本文立足我国流域综合管理的特点和发展趋势,针对我国现行水功能区划的问题,结合国外流域水生态区划的经验,提出了基于流域生态学、地域分异规律、生态系统健康与生态完整性、流域生态系统管理等理论基础的,以恢复流域持续性、完整性生态系统健康为目标,反映流域水陆耦合体在不同时空尺度景观异质性的流域水生态功能区划及其原则,重点分析了流域水生态系统的空间格局、生态过程以及动态演替等3个区划的关键问题,并提出了区划的方法,以期为我国流域水生态功能区划和流域生态系统管理提供战略层次的科学依据.     

依据(代)生物体对化学品的反应预测对生态系统服务功能的影响

保护生态系统服务功能越来越多地被作为风险评估的目标,但是目前生态风险评估的终点和评估生态系统服务功能受到的潜在影响之间有很大的差距。作者提出了一个框架,将常用的生态毒理学终点与对种群和群落的影响以及生态系统的服务功能联系起来。这个框架建立在机制效应模型的长足进步上,这些模型旨在跨越多种生物组织,并解释各种生物相互作用和反馈。为了说明这一点,作者引入了2个研究案例,它们采用了已完善和已验证的机制效应模型：鱼种群的inSTREAM个体模型和AQUATOX生态系统模型。他们还展示了动态能量平衡理论可以为解释组织级毒性提供一种通用货币。他们认为,一个基于机制模型的框架,可以预测化学品暴露对生态系统服务的影响,再结合经济估值,可以为环境管理提供一种有用的方法。作者强调了使用这个框架的潜在好处以及未来工作中需要解决的挑战。
精选自Forbes, V. E., Salice, C. J., Birnir, B., Bruins, R. J.F., Calow, P., Ducrot, V., Galic, N., Garber, K., Harvey, B. C., Jager, H., Kanarek, A., Pastorok, R., Railsback, S. F., Rebarber, R. and Thorbek, P. (2017), A framework for predicting impacts on ecosystem services from (sub)organismal responses to chemicals. Environmental Toxicology and Chemistry, 36: 845–859. doi: 10.1002/etc.3720
详情请见http://onlinelibrary.wiley.com/doi/10.1002/etc.3720/full
     

基于属性理论的长株潭城市群生态系统健康评价

城市生态系统是一种高度人工化的自然-社会-经济复合生态系统,其健康状况直接影响到城市的可持续发展。针对城市生态系统健康评价标准的不确定性问题,在构建评价指标体系的基础上,提出了基于属性理论的城市生态系统健康评价模型及评价方法。以长株潭城市群为研究区域,运用该模型和方法对其生态系统健康进行了综合评价。评价结果表明：长株潭城市群的生态系统现状属于一般健康类,其中自然生态子系统对区域整体健康状况有较大影响,评价结果与实际情况大体吻合。通过对各子系统层的健康度分析,识别了健康限制因素,并提出了相应的调控措施。采用属性理论方法开展城市生态系统健康评价,能较好地识别系统层综合健康状况和子系统层的健康状况,具有一定的实用价值,其评价结果为促进城市生态建设,有效实施生态系统健康管理提供了科学依据。     

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.     

Identifying key species in ecosystems with stochastic sensitivity analysis

The development of approaches to estimate the vulnerability of biological communities and ecosystems to extirpations and reductions of species is a central challenge of conservation biology. One key aim of this challenge is to develop quantitative approaches to estimate and rank interaction strengths and keystoneness of species and functional groups, i.e. to quantify the relative importance of species. Network analysis can be a powerful tool for this because certain structural aspects of ecological networks are good indicators of the mechanisms that maintain co-evolved, biotic interactions. A static view of ecological networks would lead us to focus research on highly-central species in food webs (topological key players in ecosystems). There are a variety of centrality indices, developed for several types of ecological networks (e.g. for weighted and un-weighted webs). However, truly understanding extinction and its community-wide effects requires the use of dynamic models. Deterministic dynamic models are feasible when population sizes are sufficiently large to minimize noise in the overall system. In models with small population sizes, stochasticity can be modelled explicitly. We present a stochastic simulation-based ecosystem model for identification of “dynamic key species” in situations where stochastic models are appropriate. To demonstrate this approach, we simulated ecosystem dynamics and performed sensitivity analysis using data from the Prince William Sound, Alaska ecosystem model. We then compare these results to those of purely topological analyses and deterministic dynamic (Ecosim) studies. We present the relationships between various topological and dynamic indices and discuss their biological relevance. The trophic group with the largest effect on others is nearshore demersals, the species mostly sensitive to others is halibut, and the group of both considerable effect on and sensitivity to others is juvenile herring. The most important trophic groups in our dynamical simulations appear to have intermediate trophic levels.     

Primary production and rain use efficiency across a precipitation gradient on the Mongolia Plateau

Understanding how the aboveground net primary production (ANPP) of arid and semiarid ecosystems of the world responds to variations in precipitation is crucial for assessing the impacts of climate change on terrestrial ecosystems. Rain-use efficiency (RUE) is an important measure for acquiring this understanding. However, little is known about the response pattern of RUE for the largest contiguous natural grassland region of the world, the Eurasian Steppe. Here we investigated the spatial and temporal patterns of ANPP and RUE and their key driving factors based on a long-term data set from 21 natural arid and semiarid ecosystem sites across the Inner Mongolia steppe region in northern China. Our results showed that, with increasing mean annual precipitation (MAP), (1) ANPP increased while the interannual variability of ANPP declined, (2) plant species richness increased and the relative abundance of key functional groups shifted predictably, and (3) RUE increased in space across different ecosystems but decreased with increasing annual precipitation within a given ecosystem. These results clearly indicate that the patterns of both ANPP and RUE are scale dependent, and the seemingly conflicting patterns of RUE in space vs. time suggest distinctive underlying mechanisms, involving interactions among precipitation, soil N, and biotic factors. Also, while our results supported the existence of a common maximum RUE, they also indicated that its value could be substantially increased by altering resource availability, such as adding nitrogen. Our findings have important implications for understanding and predicting ecological impacts of global climate change and for management practices in arid and semiarid ecosystems in the Inner Mongolia steppe region and beyond.     

Evaluation of ecosystem health for regional development in the southwestern Beijing, China

Ecosystem health assessment is one of the most important issues in regional ecological quality and safety studies. It also has a great significance to ecological conservation and regional development. This study focused on assessing the health status of forest, agriculture and urban ecosystems in the southwestern Beijing, China, including the Fangshan and Fengtai Districts. Based on field surveys and data collection, an assessment index system containing the vigor, organization and resilience factors was developed to measure the ecosystem health. Each index was scored from 1 to 5, representing five levels that contribute to the whole ecosystem health. The analytic hierarchy process (AHP) method was used to measure the weights of each index and three factors, and thus an overall score for a certain ecosystem was calculated. The results show that the forest ecosystem in the Fangshan District had higher health value than the Fengtai District, while the urban ecosystem had a slightly lower value in the Fangshan District than the Fengtai District. Both districts show lower resilience values in forest and urban ecosystems. Maintaining the ecosystem health will definitely benefit the long-term development of two districts. This study suggested that an approach for assessing the regional ecosystem health could be further developed to address spatial and synergy relationships between ecosystems and the three health factors.     

生态系统服务内涵、价值评估与GIS表达

生态系统服务价值评估是当前生态经济学领域的研究热点,为解决生态系统之于人类的重要性提供了量化的参考,也是生态补偿机制建立的重要依据.近5年国际上SCI论文发表数量以年均40%速度增长,我国研究起步于上世纪90年代末,十年来发展迅速,从对国外研究的简单模仿逐渐转向对评估模型参数的修正及对技术方法的适应性集成与发展.生态系统服务的内涵研究不断推进,在阐释“自然组分-生态过程-生态功能-生态服务.获得利益”关系上呈现多种观点,相应地提出了不同的价值评估指标体系;生态系统服务价值评估以瞬时静态为主,动态评估略显不足;研究对象从大尺度和单一生态系统逐渐转向中尺度区域,评估结果表现形式从单一数值化向基于GIS的空间表达发展.在回顾全球范围生态系统服务评估研究成果的基础上,从研究区域分布与学科特点、内涵及分类、评估方法和GIS技术应用等四大方面进行归纳分析,并从生态系统管理和决策支持需求的角度,指出生态系统评估研究已取得显著进展,但有关生态系统服务的内涵与分类、生态系统服务价值的动态评估及GIS技术的应用研究仍待深入.     

Targeting Ecosystem Features for Conservation: Standing Crops in the Florida Everglades

Abstract: The Everglades in southern Florida, U.S.A., is a major focus of conservation activities. The freshwater wetlands of the Everglades do not have high species richness, and no species of threatened aquatic animals or plants live there. We have, however, identified a distinctive ecological feature of the Everglades that is threatened by canal construction, draining, and nutrient enrichment from agricultural runoff. Compared to values reported from other freshwater systems, standing stocks of periphyton in relatively undisturbed areas of the Everglades were unusually high, and standing stocks of invertebrates and fish were unusually low. Averaging data gathered from nine sites and five sampling periods spanning 1 year, we found that periphyton standing crop was 88.2 g/m² (ash-free dry mass), invertebrate standing stock was 0.64 g/m² (dry mass), and fish standing stock was 1.2 g/m² (dry mass of large and small species combined). We found that fish standing stocks were much higher in phosphorus-enriched sites than in nearby reference sites but that invertebrate standing stocks were similar in enriched and reference sites. Our results support the notion that oligotrophy is at least partially responsible for the low standing stocks of fish, but they also suggest that species interactions and a paucity of deep-water refugia are important. Anthropogenic eutrophication in Everglades marshes will lead to the loss of distinctive ecosystem features. A focus on species richness and "hot spots" of threatened species provides no basis for conservation of ecosystems like the Everglades. If oligotrophic ecosystems often have low species richness, they will be underrepresented in preservation networks based on some common criteria for establishing conservation priorities.     

Modelling coral reef ecosystems with limited observational data

Ecosystem models represent potentially powerful tools for coral reef ecosystem managers. They can provide insight into ecosystem dynamics not achievable through alternative means allowing coral reef managers to assess the potential outcome of any given management decision. One of the main limitations in the applicability of ecosystem models is that they often require detailed empirical data and this can restrict their applicability to ecosystems that are either currently well studied or have the resources available to collect the required data. This study describes the development of a coral reef ecosystem model that can be calibrated to an ecosystem with limited empirical data. Based on the assumption that coral reef ecological structure is generic across all tropical coral reefs and that the magnitude of the interactions between ecological components is reef specific, the dynamics of the ecosystem can be replicated based on limited empirical data. The model successfully replicated the dynamics of three individual reef systems including an inshore and oceanic reef within the Great Barrier Reef and a Caribbean reef system. It highlighted the importance of understanding the specific dynamics of a given reef and that a positive management intervention in one system may result in a negative outcome for another. The model was also used to assess the importance of various interactions within coral reef ecosystems. It identified the interactions between hard corals and other non-algal benthic components as being an important (but currently understudied) facet of coral reef ecology. The development of this modelling approach provides access to ecosystem modelling tools for coral reef managers previously excluded due to a lack of resources or technical expertise.     

Predator diversity and trophic interactions

The recognition that predators play important roles in ecosystems has prompted research to resolve how combinations of predator species influence ecosystem functions. Interactions among predator species and their prey can lead to a host of linear and nonlinear effects. Understanding the conditions causing these effects is critical for assigning predator species to functional groups in ways that lead to predictive theory of predator diversity effects on trophic interactions. To this end, I provide a synthesis of experiments examining multiple-predator-species effects on mortality of single shared prey. I show how experimental design and experimental venue can determine the conclusion about the importance of predator diversity on trophic interactions. In addition, I link natural history insights on predator species habitat and hunting behavior with linear and nonlinear multiple-predator effects to derive a new concept of predator diversity effects on trophic interactions. This concept holds that the nature of predator diversity effects is contingent upon predator species hunting mode plus predator and prey species habitat domain (defined as the spatial extent to which a microhabitat is used by a species). This concept allows the classification of multiple-predator effects into four broad functional categories: substitutable, nonlinear due to predator species interference, nonlinear due to intraguild predation, and nonlinear due to predator species synergism. Experimental evidence so far provides ample and comparatively equal support for substitutable, interference, and intraguild effects, and equivocal support for nonlinear synergisms. The paper closes by discussing ways to further a research program aimed at using the building blocks presented here to understand predator functional diversity and trophic interactions in complex ecological systems.     

Assessing ecological function in the context of species recovery

Species interactions matter to conservation. Setting an ambitious recovery target for a species requires considering the size, density, and demographic structure of its populations such that they fulfill the interactions, roles, and functions of the species in the ecosystems in which they are embedded. A recently proposed framework for an International Union for Conservation of Nature Green List of Species formalizes this requirement by defining a fully recovered species in terms of representation, viability, and functionality. Defining and quantifying ecological function from the viewpoint of species recovery is challenging in concept and application, but also an opportunity to insert ecological theory into conservation practice. We propose 2 complementary approaches to assessing a species’ ecological functions: confirmation (listing interactions of the species, identifying ecological processes and other species involved in these interactions, and quantifying the extent to which the species contributes to the identified ecological process) and elimination (inferring functionality by ruling out symptoms of reduced functionality, analogous to the red-list approach that focuses on symptoms of reduced viability). Despite the challenges, incorporation of functionality into species recovery planning is possible in most cases and it is essential to a conservation vision that goes beyond preventing extinctions and aims to restore a species to levels beyond what is required for its viability. This vision focuses on conservation and recovery at the species level and sees species as embedded in ecosystems, influencing and being influenced by the processes in those ecosystems. Thus, it connects and integrates conservation at the species and ecosystem levels.     

Incorporating Surrogate Species and Seascape Connectivity to Improve Marine Conservation Outcomes

Conservation focuses on maintaining biodiversity and ecosystem functioning, but gaps in our knowledge of species biology and ecological processes often impede progress. For this reason, focal species and habitats are used as surrogates for multispecies conservation, but species‐based approaches are not widely adopted in marine ecosystems. Reserves in the Solomon Islands were designed on the basis of local ecological knowledge to conserve bumphead parrotfish (Bolbometopon muricatum) and to protect food security and ecosystem functioning. Bumphead parrotfish are an iconic threatened species and may be a useful surrogate for multispecies conservation. They move across tropical seascapes throughout their life history, in a pattern of habitat use that is shared with many other species. We examined their value as a conservation surrogate and assessed the importance of seascape connectivity (i.e., the physical connectedness of patches in the seascape) among reefs, mangroves, and seagrass to marine reserve performance. Reserves were designed for bumphead parrotfish, but also enhanced the abundance of other species. Integration of local ecological knowledge and seascape connectivity enhanced the abundance of 17 other harvested fish species in local reserves. This result has important implications for ecosystem functioning and local villagers because many of these species perform important ecological processes and provide the foundation for extensive subsistence fisheries. Our findings suggest greater success in maintaining and restoring marine ecosystems may be achieved when they are managed to conserve surrogate species and preserve functional seascape connections. Incorporación de Especies Sustitutas y de Conectividad Marina para Mejorar los Resultados de Conservación     

River restoration: the fuzzy logic of repairing reaches to reverse catchment scale degradation

River restoration is an increasingly common approach utilized to reverse past degradation of freshwater ecosystems and to mitigate the anticipated damage to freshwaters from future development and resource-extraction activities. While the practice of river restoration has grown exponentially over the last several decades, there has been little empirical evaluation of whether restoration projects individually or cumulatively achieve the legally mandated goals of improving the structure and function of streams and rivers. New efforts to evaluate river restoration projects that use channel reconfiguration as a methodology for improving stream ecosystem structure and function are finding little evidence for measurable ecological improvement. While designed channels may have less-incised banks and greater sinuousity than the degraded streams they replace, these reach-scale efforts do not appear to be effectively mitigating the physical, hydrological, or chemical alterations that are responsible for the loss of sensitive taxa and the declines in water quality that typically motivate restoration efforts. Here we briefly summarize this new literature, including the collection of papers within this Invited Feature, and provide our perspective on the limitations of current restoration.     

Analysis of ecosystem structure and function: extended path and flow analysis of a steady-state oyster reef model

An analysis of the extended path and flow structure of a six compartment steady-state oyster reef model was conducted. The extended path and flow structure were analyzed in the context of a refined canonical path classification system based on the systems theory methods of environ and network unfolding analyses. A computer implementation of an operational path classification system facilitated investigation of a finite portion (path length ≤17 arcs) of the direct and indirect path structure of the oyster reef model. Important results of the path structure analysis include: (1) few simple paths and large numbers of compound paths enumerated; (2) dominance of path numbers by subsequent passage terminal cycle paths; (3) structural evidence in support of feedback control in ecosystems; (4) results provide evidence by analogy to support the hypothesis of network homogenization first described using the systems analysis methods of environ analysis and network unfolding; (5) constancy of the pattern of origin–destination path counts with increasing path length; (6) importance of nonliving compartments in the extended path structure of ecosystems. Simultaneous path and flow analysis of the oyster reef model assessed the flow contributions of the fundamental path categories for this model using a modification of a path-based network unfolding method. First passage paths contribute most of the flow; however, multiple passage cyclic paths also provide a large (22%) flow contribution. Because of cycling in the system, the numerous long paths in the extended path structure of this ecosystem model are significant in its function as represented by the flows. These results provide microscopic evidence for the macroscopic results of environ analysis that implicate cycling as a key ecosystem attribute in the mechanisms of holistic system determination. The principles enunciated here for a model with a low cycling index (11%) carry over to, and would be even more significant for, models with high cycling indexes. These results also serve to form a link between the extended structure of food webs and their functioning as represented by energy-matter flows. The present analysis demonstrates that extended path structure, and the component articulation from which it is generated, have significant consequences for ecosystem function.     

Conserving Biological Diversity through Ecosystem Resilience

Confusion over the term ecological redundancy (Walker 1992) requires that the concept be clarified in order to advance the developing theory that maintaining ecosystem function conserves biological diversity. The species approach to conserving biological diversity assumes that the species in trouble are already identified. The ecosystem approach attempts to deal with the problem of conserving all the species in an ecosystem, including those not yet known. This is best achieved by ensuring that the ecosystem continues to function approximately as it has by maintaining its essential structure. Ecosystem stability (the probability of all species persisting) is enhanced if each important functional group of organisms (important for maintaining function and structure) comprises several ecologically equivalent species, each with different responses to environmental factors. In this sense ecological redundancy is good because it enhances ecosystem resilience, but functionally important groups (guilds, functional types) that have only one or very few species deserve priority conservation attention because their functions could be quickly lost with species extinctions.