Creation and preservation of vegetation patterns by grazing

Structural patterns of tall stands (“tussock”) and short stands (“lawn”) are observed in grazed vegetation throughout the world. Such structural vegetation diversity influences plant and animal diversity. A possible mechanism for the creation and preservation of such patterns is a positive feedback between grazing and plant palatability. Although some theoretical studies have addressed this point in a non-spatial setting, the spatial consequences of this feedback mechanism on the stability and spatial characteristics of vegetation structure patterns have not been studied.We addressed this issue by analyzing a spatially explicit individual-based plant-grazer simulation model, based on published empirical relations and the assumption of optimal foraging.In the model, the selection by the grazer of short stands (that have a higher energy content and are more palatable) is affected by traveling costs and the spatial organization of swards. Nevertheless, the most selected biomass in this type of short stands was the optimal biomass predicted by cropping and digestion constraints. As a result of the optimal foraging strategy, the grazers displayed Lévy-flight traveling behavior during the simulations with characteristic exponent μ ≈ 2.Patterns of short and tall stands created by grazing were preserved for at least a decade. Even in seasonal habitat, the spatial organization of the patterns remained relatively constant, despite fluctuations in the area of short stands. Heterogeneity of initial vegetation increased heterogeneity of the grazing-induced pattern, but did not affect its stability.The area of short stands that was preserved by grazing scaled with the herbivore mass to the power 0.4 and with the carrying capacity of the vegetation to the power −0.75.Patterns of tall and short stands can be created and perpetuated by optimally grazing ruminants, irrespective of possible underlying soil patterns. The simulations generate predictions for the stability and spatial characteristics of such structural vegetation patterns.     

Ideal free distributions under predation risk

 We examine the trade-off between gathering food and avoiding predation in the context of patch use by a group of animals. Often a forager will have to choose between feeding sites that differ in both energetic gain rate and predation risk. The ideal site will have a high gain rate and low risk of predation. However, intake rate will often decrease when the patch is shared with other foragers and it may be optimal for some individuals to feed elsewhere. Within the framework of ideal free theory, we investigate the distribution of foragers that will equalise individual fitness gains. We focus on a two-patch environment with continuous inputs of food. With reference to existing experimental studies, we examine the effects of risk dilution, food input rates and an animal’s expectations of the future. We identify the effect of total animal numbers when one patch is subject to predation risk and the other is safe. Conditions under which the difference in intake rate in the two patches is constant are identified, as are conditions in which the ratio of animals in the two patches is constant. If current conditions do not alter future expectations an increase in input rates to the patches promotes increased use of the risky patch. Yet, if conditions are assumed to persist indefinitely the opposite effect is seen. When both patches are subject to predation risk, dilution of risk favours more extreme distributions, and may lead to more than one stable distribution. The results of these models are used to critically analyse previous work on the energetic equivalence of risk. This paper is intended to help guide the development of new experimental studies into the energy-risk trade-off. Received: 10 February 1995/Accepted after revision: 1 October 1995     

Pattern-oriented modeling of bird foraging and pest control in coffee farms

We develop a model of how land use and habitat diversity affect migratory bird populations and their ability to suppress an insect pest on Jamaican coffee farms. Bird foraging—choosing which habitat patch and prey to use as prey abundance changes over space and time—is the key process driving this system. Following the “pattern-oriented” modeling strategy, we identified nine observed patterns that characterize the real system's dynamics. The model was designed so that these patterns could potentially emerge from it. The resulting model is individual-based, has fine spatial and temporal resolutions, represents very simply the supply of the pest insect and other arthropod food in six habitat types, and includes foraging habitat selection as the only adaptive behavior of birds. Although there is an extensive heritage of bird foraging theory in ecology, most of it addresses only the individual level and is too simple for our context. We used pattern-oriented modeling to develop and test foraging theory for this across-scale problem: rules for individual bird foraging that cause the model to reproduce a variety of patterns observed at the system level. Four alternative foraging theories were contrasted by how well they caused the model to reproduce the nine characteristic patterns. Four of these patterns were clearly reproduced with the “null” theory that birds select habitat randomly. A version of classical theory in which birds stay in a patch until food is depleted to some threshold caused the model to reproduce five patterns; this theory caused lower, not higher, use of habitat experiencing an outbreak of prey insects. Assuming that birds select the nearby patch providing highest intake rate caused the model to reproduce all but one pattern, whereas assuming birds select the highest-intake patch over a large radius produced an unrealistic distribution of movement distances. The pattern reproduced under none of the theories, a negative relation between bird density and distance to trees, appears to result from a process not in the model: birds return to trees at night to roost. We conclude that a foraging model for small insectivorous birds in diverse habitat should assume birds can sense higher food supply but over short, not long, distances.     

Metabolism and swimming efficiency of the bonnethead sharkSphyrna tiburo

Routine metabolic rates of bonnethead sharks,Sphyrna tiburo, of 95 to 4 650 g, ranged from 70.4 to 15.0 kcal kg^–1 d^–1. Over the size range 34 to 95 cm total length, shark swimming-velocities varied from about 29 to 67 cm s^–1. Swimming velocities predicted using Weih's cost-optimization model were similar to observed velocities. The total cost of transport (the energetic cost of transporting 1 unit of body mass 1 km distance) for 1 to 8 kg sharks varied from 0.67 to 0.40 cal g^–1 km^–1. The energetic range (an estimation of the distance traveled after a 25% reduction in body weight) indicates that a 1 kg bonnethead shark would travel 500 km distance in 17 d before displaying a 25% reduction in weight. An 8 kg individual would travel 830 km in 23 d. Although the bonnethead shark is a continuously active species, its routine metabolic rate and the efficiency of its locomotory system may be similar to that of typical bony fishes.     

Modeling a wetland system: The case of Keoladeo National Park (KNP), India

A model for the wetland part of KNP is presented and analyzed. Two-dimensional parameter scans suggest that this minimal model possesses dynamical complexities. Per capita availability of water to “bad” biomass (W₁) is one of the most vital parameters. One can ensure good health of the park by restricting the par capita availability of water to low values. Getting the “bad” biomass removed by granting permits to villagers should go hand in hand with water management and conservation activities. The model presented in this paper may be helpful in designing the timing and nature of human interventions in the form of implementation of well worked out policies in future.     

What did Lotka really say? A critical reassessment of the “maximum power principle”

This paper presents a critical discussion of the so-called “maximum power principle”, often quoted in studies about the energy balance of living systems and also known in the emergy literature as “maximum em-power principle”. Several authors consider this principle highly relevant and some even proposed it as a “fourth law of thermodynamics”. A critical analysis of the original source, namely Alfred Lotka's 1921-22 papers, conducted both in an historical perspective (the connection between Lotka's writings and the ongoing debate at his time) and in a more modern context, leads to a more detailed and less biased assessment. It turns out that in spite of Lotka's very anticipatory and incredibly sharp vision of the possible interconnections between the second law of thermodynamics and evolutionism, doubts arise about the general applicability of his “maximum power principle”. From an accurate analysis of his writings, it can be concluded that: (a) Lotka explicitly and consistently addressed the “optimal use” of the flow of exergy (available energy), and therefore the quantity defined as “em-power” is an incorrect interpretation of Lotka's constrained maximum power principle; (b) “Lotka's principle” can be reformulated within Ziegler's “maximum entropy production” or Prigogine “minimum entropy generation” paradigm only under two different respective sets of rather stringent additional conditions which Lotka was probably already aware of but never explicitly stated.     

Why are bumble bees risk-sensitive foragers?

Summary In a controlled laboratory experiment, we re-examined the question of bumble bee risk-sensitivity. Harder and Real's (1987) analysis of previous work on bumble bee risk aversion suggests that risk-sensitivity in these organisms is a result of their maximizing the net rate of energy return (calculated as the average of expected per flower rates). Whether bees are risk-sensitive foragers with respect to minimizing the probability of energetic shortfall is therefore still an open question. We examined how the foraging preferences of bumble bees for nectar reward variation were affected by colony energy reserves, which we manipulated by draining or adding sucrose solution to colony honey pots. Nine workers from four confined colonies of Bombus occidentalis foraged for sucrose solution in two patches of artificial flowers. These patches yielded the same expected rate of net energy intake, but floral volumes were variable in one patch and constant in the other. Our results show that bumble bees can be both risk-averse (preferring constant flowers) and risk-prone (preferring variable flowers), depending on the status of their colony energy reserves. Diet choice in bumble bees appears to be sensitive to the target value a colony-level energetic requirement. Offprint requests to: R.V. Cartar     

A spatial model of phytoplankton patchiness

The one-dimensional theory of critical-length scales of phytoplankton patchiness is developed to include phytoplankton growth and herbivore grazing as functions of time and space. The critical-length scale L _c for the pathch is then determined by the initial spatial distribution and concentration of the limiting nutrient and herbivores in addition to the daily averaged values of the growth and loss processes. The response of an initial phytoplankton patch to the stresses of turbulent diffusion, nutrient depletion, light periodicity, and nocturnal or continuous herbivore grazing is investigated numerically for several oceanic conditions. Nocturnal grazing, while less stressful on primary production than continous grazing, results in lower phytoplankton standing stocks. Increase in biomass of vertically migrating zooplankton results in a net loss of nutrient which might otherwise be egested, recycled, and utilized in the euphotic zone under continuous grazing conditions. The Ivlev constant is shown via sensitivity analysis to be a significant parameter ultimately influencing phytoplankton production. It is demonstrated numerically that diffusion of phytoplankton cells from areas of high concentration to low concentration prevents the local extinction of the standing stock, thereby rendering a positive herbivore grazing-threshold unnecessary for ecosystem stability.     

Are savannas patch-dynamic systems? A landscape model

Savannas are ecosystems characterized by the coexistence of woody species (trees and bushes) and grasses. Given that savanna characteristics are mainly formed from competition, herbivory, fire, woodcutting, and patchy soil and precipitation characteristics, we propose a spatially explicit model to examine the effects of the above-mentioned parameters on savanna vegetation dynamics in space and time. Furthermore, we investigate the effects of the above-mentioned parameters on tree–bush–grass ratios, as well as the degrees of aggregation of tree–bush–grass biomass. We parameterized our model for an arid savanna with shallow soil depth as well as a mesic one with generally deeper and more variable soil depths. Our model was able to reproduce savanna vegetation characteristics for periods of time over 2000 years with daily updated time steps. According to our results, tree biomass was higher than bush biomass in the arid savanna but bush biomass exceeded tree and grass biomass in the simulated mesic savanna. Woody biomass increased in our simulations when the soil's porosity values were increased (mesic savanna), in combination with higher precipitation. Savanna vegetation varied from open savanna to woodland and back to open savanna again. Vegetation cycles varied over ∼300-year cycles in the arid and ∼220-year cycles in the mesic-simulated savanna. Autocorrelation values indicated that there are both temporal and spatial vegetation cycles. Our model indicated cycling savanna vegetation at the landscape scale, cycles in cells, and patchiness, i.e. patch dynamics.     

Methods to empirically estimate direct and indirect rebound effect of energy-saving technological changes in households

Energy efficiency policies have a special importance within carbon emission reduction policies to mitigate the climate change effects. However, potential reductions of energy consumption and, consequently, its resulting emissions, can be offset through the so called “rebound effect”. The concept of “rebound effect” refers to a set of mechanisms whereby the improvement of efficiency reduces the cost of the energy service and this results in the household energy consumption rising and totally or partially negating the reduction achieved by the energy efficiency improvement. This paper provides a methodology to estimate the static direct plus indirect rebound effect of energy efficiency improvements in the use of energy in households. It is based on the combination of econometric estimations of energy demand functions, re-spending modelling and generalised intput-output of energy modelling. It also provides estimations for Catalonia.     

On the integration of individual foraging strategies with colony ergonomics in social insects: nectar-collection in honeybees

Summary We experimentally tested whether foraging strategies of nectar-collecting workers of the honeybee (Apis mellifera) vary with colony state. In particular, we tested the prediction that bees from small, fast growing colonies should adopt higher workloads than those from large, mature colonies. Queenright small colonies were set up by assembling 10 000 worker bees with approximately 4100 brood cells. Queenright large colonies contained 35 000 bees and some 14 500 brood cells. Thus, treatments differed in colony size but not in worker/brood ratios. Differences in workload were tested in the context of single foraging cycles. Individuals could forage on a patch of artificial flowers offering given quantities and qualities of nectar rewards. Workers of small colonies took significantly less nectar in an average foraging excursion (small: 40.1 ± 1.1 SE flowers; large: 44.8 ± 1.1), but spent significantly more time handling a flower (small: 7.3 ± 0.4 s ; large: 5.8 ± 0.4 s). When the energy budgets for an average foraging trip were calculated, individuals from all colonies showed a behavior close to maximization of net energetic efficiency (i.e., the ratio of net energetic gains to energetic costs). However, bees from small colonies, while incurring only marginally smaller costs, gained less net energy per foraging trip than those from large colonies, primarily as a result of prolonged handling times. The differences between treatments were largest during the initial phases of the experimental period when also colony development was maximally different. Our results are at variance with simple models that assume natural selection to have shaped behavior in a single foraging trip only so as to maximize colony growth. Offprint requests to: P. Schmid-Hempel     

Differential use of conspecific-derived information by sexual and asexual parasitic wasps exploiting partially depleted host patches

When foraging partially depleted patches (i.e., a fraction of hosts are already parasitized), female parasitoids must decide: 1—whether to superparasitize, and 2—whether to stay in their current patch (thus missing the opportunity of finding a better patch elsewhere). To make these decisions, parasitoids may rely on different cues, produced both by the environment and by conspecifics. Animals thriving in different environments may differ in cues they use. In the solitary parasitoid Venturia canescens, thelytokous (asexual) and arrhenotokous (sexual) individuals are found in two contrasting environments. Thelytokous females, from anthropogenic conditions, are known to cope with superparasitism in an adaptive way. On the other hand, little is known about superparasitism by arrhenotokous females. We compared the host exploitation strategies of thelytokous and arrhenotokous females in partially depleted patches. Hosts parasitized by thelytokous females were more frequently avoided than those parasitized by arrhenotokous females, suggesting a stronger chemical marking of the former. Only thelytokous females used information from conspecifics for patch-leaving decisions. The conformity of the differences in the behavior of thelytokous and arrhenotokous females with the environmental conditions they experience in their habitat is discussed.     

Effect of temperature on specific dynamic action in the common octopus,<Emphasis Type="Italic"> Octopus vulgaris</Emphasis> (Cephalopoda)

Feeding causes an increase of metabolic rate, which initially escalates rapidly, reaches a peak value and then gradually declines to the pre-feeding rate. This phenomenon, termed specific dynamic action (SDA), reflects the energy requirements of the behavioral, physiological and biochemical processes that constitute feeding. The effect of temperature on SDA of the common octopus, Octopus vulgaris, was evaluated, by measuring the temporal pattern of the oxygen consumption rates of octopuses, after feeding, at two constant temperatures, 20°C and 28°C. At 20°C, the relative increase in the oxygen consumption rate after feeding (relative SDA) was significantly greater than at 28°C. The peak of the relative SDA occurred 1 h after feeding, and it was 64% at 20°C and 42% at 28°C. However, the SDA absolute peak, SDA duration (9.5 h) and SDA magnitude (the integrated postprandial increase in oxygen uptake) did not differ significantly between the two temperatures, indicating that the energetic cost of feeding was the same at both temperatures. The SDA response in O. vulgaris was much faster than it was in polar species, which have extended SDA responses due to low temperatures, and was also relatively fast in relation to the response in other temperate species, which is probably connected to the remarkably high growth rates of the species. A possible explanation of the observed summer migration of large octopuses from shallow to deeper areas is given, based on the effect of temperature on the energetic requirements of octopuses.     

Social foraging by honeybees: how colonies allocate foragers among patches of flowers

Summary To understand how a colony of honeybees keeps its forager force focussed on rich sources of food, and analysis was made of how the individual foragers within a colony decide to abandon or continue working (and perhaps even recruit to) patches of flowers. A nectar forager grades her behavior toward a patch in response to both the nectar intake rate of her colony and the quality of her patch. This results in the threshold in patch quality for acceptance of a patch being higher when the colonial intake rate of nectar is high than when it is low. Thus colonies can adjust their patch selectivity so that they focus on rich sources when forage is abundant, but spread their workers among a wider range of sources when forage is scarce. Foragers assess their colony's rate of nectar intake while in the nest, unloading nectar to receiver bees. The ease of unloading varies inversely with the colonial intake rate of nectar. Foragers assess patch quality while in the field, collecting nectar. By grading their behavior steeply in relation to such patch variables as distance from the nest and nectar sweetness, foragers give their colony high sensitivity to differences in profitability among patches. When a patch's quality declines, its foragers reduce their rate of visits to the patch. This diminishes the flow of nectar from the poor patch which in turn stimulates recruitment to rich patches. Thus a colony can swiftly redistribute its forager force following changes in the spatial distribution of rich food sources. The fundamental currency of nectar patch quality is not net rate of energy intake, (Gain-Cost)/Time, but may be net energy efficiency, (Gain-Cost)/Cost.     

The relative importance of host-plant genetic diversity in structuring the associated herbivore community

Recent studies suggest that intraspecific genetic diversity in one species may leave a substantial imprint on the surrounding community and ecosystem. Here, we test the hypothesis that genetic diversity within host-plant patches translates into consistent and ecologically important changes in the associated herbivore community. More specifically, we use potted, grafted oak saplings to construct 41 patches of four saplings each, with one, two, or four tree genotypes represented among the host plants. These patches were divided among two common gardens. Focusing first at the level of individual trees, we assess how tree-specific genotypic identity, patch-level genetic diversity, garden-level environmental variation, and their interactions affect the structure of the herbivore community. At the level of host-plant patches, we analyze whether the joint responses of herbivore species to environmental variation and genetic diversity result in differences in species diversity among tree quartets. Strikingly, both species-specific abundances and species diversity varied substantially among host-tree genotypes, among common gardens, and among specific locations within individual gardens. In contrast, the genetic diversity of the patch left a detectable imprint on local abundances of only two herbivore taxa. In both cases, the effect of genetic diversity was inconsistent among gardens and among host-plant genotypes. While the insect community differed significantly among individual host-plant genotypes, there were no interactive effects of the number of different genotypes within the patch. Overall, additive effects of intraspecific genetic diversity of the host plant explained a similar or lower proportion (7-10%) of variation in herbivore species diversity than did variation among common gardens. Combined with the few previous studies published to date, our study suggests that the impact of host-plant genetic diversity on the herbivore community can range from none to nonadditive, is generally low, and reaches its most pronounced impact at small spatial scales. Overall, our findings strengthen the emerging view that the impacts of genetic diversity are system, scale, and context dependent. As the next step in community genetics, we should then start asking not only whether genetic diversity matters, but under what circumstances its imprint is accentuated.     

The model of long-term evolution of the carbon cycle

The weathering processes and their role in the formation of the atmospheric carbon and, as a consequence, on the climate are considered. The model operates in the framework of “active planetary cover”, i.e. considering the interactive role of the biosphere, looking at its development as a non-linear evolutionary system of so-called “virtual biospheres”.     

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.     

林下和撂荒地生境紫茎泽兰幼苗的生长特征及其种内竞争形态反应

比较研究了林下和撂荒地生境有无竞争条件下紫茎泽兰（Ageratina adenophora（Sprengel））幼苗的生长特征、生长速率、生物量和种内竞争的形态反应。结果表明：（1）2类生境条件下紫茎泽兰植株幼苗形态特征基本一致,无竞争条件下紫茎泽兰幼苗根长和叶片数明显高于在竞争条件下对应指标值;（2）同类生境中,在无竞争条件下紫茎泽兰幼苗的生长速率、生物量积累显著比在竞争条件下的高;（3）不同生境中,撂荒地生境紫茎泽兰幼苗生物量显著高于林下生境的值,在无竞争条件下,林下生境紫茎泽兰幼苗生长速率比撂荒地生境的高,而在竞争条件下反之;（4）林下和撂荒地生境紫茎泽兰幼苗植株竞争强度差异不显著,但在林下生境其地上部分的竞争强度显著大于其在撂荒地生境的值,而根的竞争强度相反。可见,紫茎泽兰幼苗在种群建立的过程中具有不同的生态适应性和形态反应,这可能是其能成功入侵不同生境的一种重要生理生态策略。     

Assessing the Potential to Restore Historic Grazing Ecosystems with Tortoise Ecological Replacements

The extinction of large herbivores, often keystone species, can dramatically modify plant communities and impose key biotic thresholds that may prevent an ecosystem returning to its previous state and threaten native biodiversity. A potentially innovative, yet controversial, landscape‐based long‐term restoration approach is to replace missing plant‐herbivore interactions with non‐native herbivores. Aldabran giant (Aldabrachelys gigantea) and Madagascan radiated (Astrochelys radiata) tortoises, taxonomically and functionally similar to the extinct Mauritian giant tortoises (Cylindraspis spp.), were introduced to Round Island, Mauritius, in 2007 to control the non‐native plants that were threatening persistence of native species. We monitored the response of the plant community to tortoise grazing for 11 months in enclosures before the tortoises were released and, compared the cost of using tortoises as weeders with the cost of using manual labor. At the end of this period, plant biomass; vegetation height and cover; and adult, seedling, flower, and seed abundance were 3–136 times greater in adjacent control plots than in the tortoise enclosures. After their release, the free‐roaming tortoises grazed on most non‐native plants and significantly reduced vegetation cover, height, and seed production, reflecting findings from the enclosure study. The tortoises generally did not eat native species, although they consumed those native species that increased in abundance following the eradication of mammalian herbivores. Our results suggest that introduced non‐native tortoises are a more cost‐effective approach to control non‐native vegetation than manual weeding. Numerous long‐term outcomes (e.g., change in species composition and soil seed bank) are possible following tortoise releases. Monitoring and adaptive management are needed to ensure that the replacement herbivores promote the recovery of native plants. Estudiando el Potencial para Restaurar Ecosistemas Históricos de Forrajeo con Reemplazos Ecológicos de Tortugas Terrestres     

How might we model an ecosystem?

Predicting ecosystem effects is of crucial importance in a world at threat from natural and human-mediated change. Here we propose an ecologically defensible representation of an ecosystem that facilitates predictive modelling. The representation has its roots in the early trophic and energetic theory of ecosystem dynamics and more recent functional ecology and network theory. Using the arable ecosystem of the UK as an example, we show that the representation allows simplification from the many interacting plant and invertebrate species, typically present in arable fields, to a more tractable number of trophic-functional types. Our compound hypothesis is that “trophic-functional types of plants and invertebrates can be used to explain the structure, diversity and dynamics of arable ecosystems”. The trophic-functional types act as containers for individuals, within an individual-based model, sharing similar trophic behaviour and traits of biomass transformation. Biomass, or energy, flows between the types and this allows the key ecological properties of individual abundance and body mass, at each trophic height, to be followed through simulations. Our preliminary simulation results suggest that the model shows great promise. The simulation output for simple ecosystems, populated with realistic parameter values, is consistent with current laboratory observations and provides exciting indications that it could reproduce field scale phenomena. The model also produces output that links the individual, population and community scales, and may be analysed and tested using community, network (food web) and population dynamic theory. We show that we can include management effects, as perturbations to parameter values, for modelling the effects of change and indicating management responses to change. This model will require robust analysis, testing and validation, and we discuss how we will achieve this in the future.