Habitat loss, trophic collapse, and the decline of ecosystem services

The provisioning of sustaining goods and services that we obtain from natural ecosystems is a strong economic justification for the conservation of biological diversity. Understanding the relationship between these goods and services and changes in the size, arrangement, and quality of natural habitats is a fundamental challenge of natural resource management. In this paper, we describe a new approach to assessing the implications of habitat loss for loss of ecosystem services by examining how the provision of different ecosystem services is dominated by species from different trophic levels. We then develop a mathematical model that illustrates how declines in habitat quality and quantity lead to sequential losses of trophic diversity. The model suggests that declines in the provisioning of services will initially be slow but will then accelerate as species from higher trophic levels are lost at faster rates. Comparison of these patterns with empirical examples of ecosystem collapse (and assembly) suggest similar patterns occur in natural systems impacted by anthropogenic change. In general, ecosystem goods and services provided by species in the upper trophic levels will be lost before those provided by species lower in the food chain. The decrease in terrestrial food chain length predicted by the model parallels that observed in the oceans following overexploitation. The large area requirements of higher trophic levels make them as susceptible to extinction as they are in marine systems where they are systematically exploited. Whereas the traditional species-area curve suggests that 50% of species are driven extinct by an order-of-magnitude decline in habitat abundance, this magnitude of loss may represent the loss of an entire trophic level and all the ecosystem services performed by the species on this trophic level.     

Herbivores enable plant survival under nutrient limited conditions in a model grazing system

We make a theoretical study of nitrogen cycling in a model of a grazing system with five compartments. The rates of uptake of nutrient by plants and herbivores are allowed nonlinear forms which involve no a priori assumptions about whether the system is subject to top-down or bottom-up control. We derive a method of piecewise linear approximation which allows analytical study of the system. We then use this method to investigate the properties of the equilibrium states of the system, and in particular whether the system favours donor- or recipient-control, the grazing optimization problem, and the potential benefits of herbivory to plant growth. We are able to generalise our results to all uptake functions of the same qualitative class as those considered, and to show that in general the system will tend to a stable equilibrium state of donor-controlled herbivory. In this model, the presence of the ‘right’ class of herbivore is not only beneficial to plant growth in certain circumstances, but can be essential to their survival, allowing plants to co-exist with herbivores under conditions in which they would be unable to survive alone.     

The impact of detritivorous fishes on a mangrove estuarine system

Sundarban mangrove ecosystem in India is one of the largest detritus based ecosystem of the world and it supplies the detritus and nutrients to the adjacent Hooghly-Matla estuarine complex. In this estuary a group of fish completely detritivorous in nature, belonging to the genus Mugil spp. is present. This group of fish is expected to have important effects on the trophic dynamics of ecosystems, but exact nature of these effects is not known. In order to study the impact that detritivory by fish may have on the estuarine food chain, we developed mathematical formulations. We run two models, one with phytoplankton, zooplankton, carnivorous fish, detritus and nutrient and without this group of fish and a second one after including this fish in the system. In our model this group of fish has no major impact on primary productions of the estuarine system but has extensive role in total fish production. Coexistence of detritivorous fish and carnivorous fish occurs within reasonable parameter range. We have tested different growth rates of phytoplankton, grazing rates and predation rates of zooplankton, detritivorous fish and carnivorous fish for total system equilibrium. Carnivorous fish predation rate on detritivorous fish and detritivorous fish grazing rate on detritus are very important. Different foraging ratios are also tested in this study. Foraging preference of carnivorous fish on detritivorous fish appears significant for the system equilibrium.     

Control targets for the management of biological systems

Attention is focused on biological systems which are describable in terms of ordinary differential equations subject to human control inputs. The concept of an isochronal system is introduced in order to include systems for which the differential equations are valid only over regularly reoccurring time intervals.It is assumed that the control inputs are to be chosen so that an integral cost function of the state of the system, the control used, current time, and the time interval of the control program is minimized. Problems associated with minimizing this cost function over an infinitely long time interval is then considered. Difficulties inherent with minimizing a cost integral on an infinite time interval are shown to be avoided by minimizing an average of the cost function over an unknown but periodic time interval. Under proper circumstance, the optimal control program for the average cost function is either identical to or a good approximation to the optimal control program for the original cost function over an infinitely long time interval.Necessary conditions are obtained for minimizing an average cost function over an unspecified time interval subject to the system equations. For a given problem the necessary conditions will yield but a single system trajectory in the state space. For management purposes this trajectory may be thought of as a target to which the system should be driven and maintained.A number of examples illustrate the use of the necessary conditions to obtain control targets. Certain problems associated with the stability of the target solutions are illustrated with the examples.     

A simple random path method for the analysis of flow networks

The path of a particle through an ecosystem is modelled as a Markov chain. For a given flow network, powers of the transition matrix are used to calculate the distribution of the particles over the network after each transition. The method may be applied for the definition and calculation of trophic levels in food webs. The algorithm yields the trophic level distribution of species, the species composition of trophic levels, and the path length distribution of output flows. In addition, the network can be described as a linear chain, with the throughflows at each step identified. Data from several ecosystems are analyzed by the method, showing that surprising insights may result.     

用排放情景分析系统研究北京市机动车污染问题

针对北京市动车污染非常严重,机动车排放控制涉及的方面较多,解决起来比较复杂的特点,开发了城市机动车排放情况分析系统,以国内外的先进理论和技术为基础,采用系统化的方法研究这个问题,对北京市未来车污染控制的几种情况进行了分析和比较,证明了北京市目前实施新排放标准,采用车辆定期报废制度对控制污染的巨大作用,提出进一步强化I／M制度,完善在用车监测维修体系等控制机动车污染的具体方法。     

Fish wars and cooperation maintenance

In this paper, a discrete-time game model related to a bioresource management problem (fish catching) is considered. We divide a fishery into regions, which are exploited by single players. The center (referee) shares a reservoir between the competitors. The players (countries), which harvest the fish stock are the participants of this game.We assume that there are migratory exchanges between the regions of the reservoir. Therefore, the stock in one region depends not only on the previous stock and catch in the region, but also on the stock and catch in neighboring regions. We derive the Nash and cooperative equilibria for an infinite planning horizon.We consider two ways to maintain the cooperation: incentive equilibrium and time-consistent imputation distribution procedure. We investigate the cooperative incentive equilibrium in the case when the center punishes players for a deviation.Also we consider the case when the center is a player and find the Shapley value and time-consistent imputation distribution procedure. We introduce a new condition which offers an incentive to players to keep cooperating.     

Resources from another place and time: responses to pulses in a spatially subsidized system

As the theoretical bases for the dynamics of spatially subsidized communities emerge, ecologists question whether spatially subsidized communities exhibit similar structure or dynamics to communities that receive strongly pulsed resources. In both cases, communities may be structured by responses to resources that are potentially absent at any given point in time (pulsed communities) or space (subsidized communities), even if pulsed resources are part of the in situ productivity of the system or the subsidies arrive as a relatively constant input from a nearby system. The potential for significant spatial or temporal resource limitation, therefore, may be a key factor influencing in similar ways the persistence of populations, the structure and dynamics of communities, and the evolution of specific life history traits. In most complex systems, however, multiple resources may arrive for various trophic entities at various points in time and from various points in space, and thus it may be difficult to separate or compare the dynamics of spatially subsidized and pulsed systems. In this paper, we explore the effects of interactions between pulses and subsidies in plant and animal populations and communities on highly pulsed and variably subsidized islands in the Gulf of California. While many of the plant and animal communities on the unsubsidized islands in this system respond to pulses of rain in classic ways, responses to these rain pulses on islands subsidized by seabird guano or other marine resources are quite different and variable, and depend on a combination of life history characteristics, physiology, competitive interactions, and trophic relationships. These variable responses to rain pulses then translate into large differences in dynamics and community structure of subsidized vs. unsubsidized islands. Indeed, most systems experience both temporal pulses and spatial subsidies. When considered in tandem, complementary or synergistic effects of the multiple, temporally and spatially variable resources may emerge that help explain complex food web structure and dynamics.     

Habitat Fragmentation Effects on Trophic Processes of Insect-Plant Food Webs

Abstract:  Habitat fragmentation is the transformation of once-extensive landscapes into smaller, isolated remnants surrounded by new types of habitat. There is ample evidence of impoverished biodiversity as a consequence of habitat fragmentation, but its most profound effects may actually result from functional changes in ecological processes such as trophic interactions. We studied the trophic processes of herbivory and parasitism in insect-plant food webs composed of hundreds of species in a fragmented woodland landscape. We recorded all plant species, collected mined leaves, and reared leafminers and parasitoids from 19 woodland remnants. Herbivory and parasitism rates were then analyzed in relation to woodland size and edge or interior location. Herbivory by leaf-mining insects and their overall parasitism rates decreased as woodland remnants became smaller. For each remnant the intensity of both processes differed between edge and interior. Our results provide novel evidence of the magnitude of habitat fragmentation effects, showing they can be so pervasive as to affect trophic processes of highly complex food webs and suggesting a response associated with trophic specialization of the involved organisms as much as with their trophic level.     

Recherches sur la situation trophique d'un groupe d'organismes pélagiques (Euphausiacea). I. Niveaux trophiques des espèces

Trophic relationships are the determining factor of biological equilibrium, as existence and abundance of any population depend on the ratio nutrition-predation. This paper is the first of a series devoted to the study of the trophic position of euphausiid crustaceans in the equatorial and south-tropical Pacific Ocean; it defines the role of a pelagic group in its biotope, suggests methods in trophic studies, and shows that analytical research on limited subjects is a reliable way to reveal more general features applying to the whole pelagic world. Until now, the literature dealing with nutrition of euphausiids in most cases has provided only qualitative lists of items found in the stomachs; the drawbacks of this kind of investigation are that only a small part of the stomach content is recognizable in this way and, moreover, that the remains identified belong only to the food consumed which had the most resistant structures. In order to study the trophic levels of species, it has been necessary to take into account the whole stomach content, to establish if its origin was plant or animal or both. Binocular examination of the dissected stomachs made it possible to discriminate between phyto- and zooplankton. More than 5000 specimens have been examined in this way, each stomach being named V if more than 80% of its content is phytoplankton, A if this same percentage is animal food, VA if phyto- and zooplankton are of comparable importance; the trophic level of each species has been defined as the percentage Σ [A+(VA/2)], which measures the part of animal material among the total food. Although zooplankton appears on the whole to be more important than phytoplankton as food for tropical euphasiids, all the trophic levels are observed among the 16 species studied, from phytophagous to strictly carnivorous.     

Functional taxonomy: An immodest proposal

The thesis is presented that classical taxonomy is of limited value to ecosystem science, and that the further development of ecosystem theory may actually be hindered by a reliance on the biological (phylogenetic) species as the basic functional unit of ecosystems. It is further argued that this situation could be improved if ecologists could agree on a system of functional classification in which ecological taxa would be distinguished solely on the basis of what they do in the context of an ecosystem, and not on their evolutionary relationships.A functional classification system is proposed in which functional taxa for specific ecosystems (ecological sectors) are defined as broad trophic groups of organisms in common vertical habitat zones, and with common inputs and outputs (ecosystem commodities and services). This system is envisioned as potentially useful in the development of comparative ecosystem theory, for constructing simulation models, for ongoing research in economic versus ecological values, and for cataloging new functional information as it becomes available.The proposed system is tentatively applied to the salt marsh estuarine ecosystem in the southeastern U.S.A. and to the swampforest ecosystem in Louisiana (U.S.A.). Twenty-five sectors are identified in the former and twenty in the latter.“It is by endless subdivisions based upon the most inconclusive differences, that some departments of natural history become so repellingly intricate”. Melville (1851), Moby Dick.     

Maximal yields from multispecies fisheries systems: rules for systems with multiple trophic levels.

Increasing centralization of the control of fisheries combined with increased knowledge of food-web relationships is likely to lead to attempts to maximize economic yield from entire food webs. With the exception of predator-prey systems, we lack any analysis of the nature of such yield-maximizing strategies. We use simple food-web models to investigate the nature of yield- or profit-maximizing exploitation of communities including two types of three-species food webs and a variety of six-species systems with as many as five trophic levels. These models show that, for most webs, relatively few species are harvested at equilibrium and that a significant fraction of the species is lost from the web. These extinctions occur for two reasons: (1) indirect effects due to harvesting of species that had positive effects on the extinct species, and (2) intentional eradication of species that are not themselves valuable, but have negative effects on more valuable species. In most cases, the yield-maximizing harvest involves taking only species from one trophic level. In no case was an unharvested top predator part of the yield-maximizing strategy. Analyses reveal that the existence of direct density dependence in consumers has a large effect on the nature of the optimal harvest policy, typically resulting in harvest of a larger number of species. A constraint that all species must be retained in the system (a "constraint of biodiversity conservation") usually increases the number of species and trophic levels harvested at the yield-maximizing policy. The reduction in total yield caused by such a constraint is modest for most food webs but can be over 90% in some cases. Independent harvesting of species within the web can also cause extinctions but is less likely to do so.     

Assessment of impacts of invasive fishes on the food web structure and ecosystem properties of a tropical reservoir in India

A network model of trophic interactions in a tropical reservoir in India was developed with the objective to quantify matter and energy flows between system components and to study the impact of invasive fishes on the ecosystem. Structure of flows and their distribution within and between trophic levels were analysed by aggregating single flows into combined flows for discrete trophic levels. The trophic flows primarily occurred in the first four trophic level (TL) and the food web structure in this reservoir ecosystem was characterized by the dominance of low TL organisms, with the highest TL of only 3.57 for the top predator. Highest system omnivory index (SOI) was observed for indigenous catfishes (0.422), followed by the exotic fish Mozambique Tilapia (0.402). Nile Tilapia and Pearl spots show the highest niche overlap which suggests high competition for similar resources. The mixed trophic impact routine reveals that an increase in the abundance of the African catfish would negatively impact almost all fish groups such as Indian major carps, Pearl spots, indigenous catfishes and Tilapines. The other invasive fish Mozambique Tilapia adversely affects the indigenous catfishes. The most interesting observation in this study is that the most dominant invasive fish in this reservoir, the Nile Tilapia does not negatively impact any of the fish groups. In fact it positively impacts the Indian major carps. The direct and indirect effects of predation between system components (i.e. fish, invertebrates, phytoplankton and detritus) are quantitatively described and the possible influence and role in the ecosystem's functioning of the invasive fish species are discussed.     

Transfer of the chlorinated hydrocarbon PCB in a laboratory marine food chain

The transfer of chlorinated hydrocarbons (CHC) in a laboratory simulation of a three trophic level marine food chain was studied. The food chain consisted of the algal flagellate Dunaliella sp., the rotifer Brachionus plicatilis, and the larva of the Northern anchovy Engraulis mordax. CHC were introduced into the seawater at concentrations representative of near-shore conditions off southern California without the use of dispersing agents. Each trophic level appeared to be in a steady-state at the time of first sampling, 5 days after inoculation. Apparent partition coefficients were calculated for each trophic level. The CHC contamination in the diet of the rotifers and anchovy larvae was also calculated. Unfed anchovy larvae accumulated the same amount of CHC as fed larvae and the final concentration appeared to be dependent on the CHC concentration in the seawater. The data in this report suggest that CHC accumulation is not a food-chain phenomenon but rather the result of direct partitioning of the compounds between the seawater and the test organisms.     

Resource identity modifies the influence of predation risk on ecosystem function

It is well established that predators can scare as well as consume their prey. In many systems, the fear of being eaten causes trait-mediated cascades whose strength can rival or exceed that of more widely recognized density-mediated cascades transmitted by predators that consume their prey. Despite this progress it is only beginning to be understood how the influence of predation risk is shaped by environmental context and whether it can exert an important influence on ecosystem-level processes. This study used a factorial mesocosm experiment that manipulated basal-resource identity (either barnacles, Semibalanus balanoides, or mussels, Mytilus edulis) to determine how resources modify the influence of predation risk, cascade strength, and the efficiency of energy transfer in two, tritrophic, rocky-shore food chains containing the predatory green crab (Carcinus maenas) and an intermediate consumer (the snail, Nucella lapillus). The effect of predation risk and the strength of trait-mediated cascades (both in absolute and relative terms) were much stronger in the barnacle than in the mussel food chain. Moreover, predation risk strongly diminished the efficiency of energy transfer in the barnacle food chain but had no significant effect in the mussel food chain. The influence of resource identity on indirect-effect strength and energy transfer was likely caused by differences in how each resource shapes the degree of risk perceived by prey. We suggest that our understanding of the connection between trophic dynamics and ecosystem functioning will improve considerably once the effects of predation risk on individual behavior and physiology are considered.     

Trophic levels and trophic tangles: the prevalence of omnivory in real food webs

The concept of trophic levels is one of the oldest in ecology and informs our understanding of energy flow and top-down control within food webs, but it has been criticized for ignoring omnivory. We tested whether trophic levels were apparent in 58 real food webs in four habitat types by examining patterns of trophic position. A large proportion of taxa (64.4%) occupied integer trophic positions, suggesting that discrete trophic levels do exist. Importantly however, the majority of those trophic positions were aggregated around integer values of 0 and 1, representing plants and herbivores. For the majority of the real food webs considered here, secondary consumers were no more likely to occupy an integer trophic position than in randomized food webs. This means that, above the herbivore trophic level, food webs are better characterized as a tangled web of omnivores. Omnivory was most common in marine systems, rarest in streams, and intermediate in lakes and terrestrial food webs. Trophic-level-based concepts such as trophic cascades may apply to systems with short food chains, but they become less valid as food chains lengthen.     

博弈论在环境保护中的应用

阐述了博弈论的基本理论在环境保护中的应用,分析结果指出应结合市场机制和政府政策改变博弈方的收益,使博弈达到有利于环境保护的纳什均衡,提高环境保护效率;并探讨了博弈论在环境保护中应用的关键技术与策略。     

紫外线B对海洋生态系统的影响

近年来随着臭氧损失现象日益严重，到达地球表面的紫外线B辐射不断加强，海洋生态系统受到了前所未有的威胁。UV-B的增强改变了生物体赖以生存的水体环境，影响了浮游植物对营养盐的吸收同化，抑制了浮游植物的光合作用，从而减少了海洋对CO2的吸收能力，其结果直接增加了温室效应。同时UV-B也直接影响生态系统食物链的各个营养级，伤害海洋病毒、细菌和浮游植物，降低浮游动物和仔稚鱼的成活率，进而影响到整个海洋生态系统的结构和功能。     

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.     

Simulating forest succession after blowdown events: The crucial role of space for a realistic management

Ecological patterns vary in space and time. Therefore, when using dynamic models in ecology, the spatial aspect should not be neglected prematurely since it could possibly change the model outcomes to a considerable extent. In view of this problem, we describe here a method how to construct a non-spatial version from a spatially explicit simulation model. The principle idea is to suppress the spatial correlations of cells in a grid in time by continuously re-assigning a random neighbourhood for each cell on the grid. Since this procedure actually eliminates the spatial dimensions, it allows to quantify the unadulterated impact of spatial processes on the model results. To illustrate an important application of this approach in the context of forest management we use a grid-based model that simulates succession of Norway spruce (Picea abies (L.) Karst.) at mountainous sites after blowdown events. The output of this model is compared with the results of the deduced non-spatial version of this model regarding the predicted amount of re-growing trees. The non-spatial version dramatically overestimates the number of spruce trees on different microsites. Thus, the uncritical use of the non-spatial model might give reason to wrong management decisions that are based on too optimistic predictions. In practice, this may lead to dangerous situations, especially in mountain forests serving as protection against avalanches and landslides. This example demonstrates the successful applicability of our approach. Our method can be interpreted as a contribution to an extended sensitivity analysis: it analyses the sensitivity of the results due to structural changes of the model. This sensitivity allows one to estimate the redundancy or the necessity of spatially explicit processes in a model with regard to the parsimony principle of modelling. Since our approach is not dependent on special features of the simulation model used here, it is assumed to be applicable for other spatial models, too, and can thus be considered of general interest for a diligent model analysis.