Collapse and reorganization of a food web of Mwanza Gulf, Lake Victoria

Lake Victoria in East Africa is the world's second largest freshwater system. Over the past century the ecosystem has undergone drastic changes. Some 30 years after the introduction of Nile perch (Lates niloticus) and Nile tilapia (Oreochromis niloticus) in the 1950s, the highly diverse community of native haplochromines collapsed, leaving a system dominated by only four species: the native cyprinid dagaa (Rastrineobola argentea) and shrimp (Caridina nilotica), as well as the introduced Nile perch and Nile tilapia. More recently, an unexpected resurgence of haplochromines has been reported. To understand these changes in terms of ecosystem functioning and of changes in growth of trophic groups, we created mass balances of the food web near Mwanza, Tanzania, before, during, and after the Nile perch boom (1977, 1987, and 2005), using the application ECOPATH. We connected these mass balances with a dynamic model assuming linear trends in net growth rates of the trophic groups. Our analysis suggests that the Nile perch boom initially altered the biomass distribution over trophic levels. Also, results indicate that not only fishing but also changes at the detritivores' trophic level might have played an important role in driving changes in the system. Both the mass balances and the dynamic model connecting them reveal that, after a major distortion during the Nile perch boom, the biomass distribution over the main trophic levels had largely recovered its original (1977) state by 2005. However, no such return appeared in terms of community structure. Biodiversity in the new state is dramatically lower, consisting of introduced species and a few native surviving species. We conclude that at an aggregate level Lake Victoria's ecosystem has proved to be resilient in the sense that its overall trophic structure has apparently recovered after a major perturbation. By contrast, its intricate functional structure and associated biodiversity have proved to be fragile and seem unlikely to recover.     

The Effects of Predation by Nile Perch, Lates niloticus L., on the Fish of Lake Nabugabo, with Suggestions for Conservation of Endangered Endemic Cichlids

As in Lake Victoria and Lake Kyoga, most of the native fish species of Lake Nabugabo have either been depleted or have disappeared since Lates niloticus , Nile perch, and Oreochromis niloticus , a tilapiine, were introduced. L. niloticus, O. niloticus, Schilbe intermedius, Brycinus jacksonii , and Rastrineobola argentea are the only species still abundant and widespread in the lake. Of the five haplochromine species endemic to Lake Nabugabo, at least one has vanished and the remaining ones are very rare. The prey of the Nile perch in Lake Nabugabo, with the exception of the prawn Caridina nilotica , which is absent from the lake, are similar to those the predator switched to in Lakes Victoria and Kyoga after haplochromines had been depleted. These consist of ephemeropterans—especially Povilla , the anisopteran nymphs R. argentea , tilapiines, and juvenile Nile perch. Three small lakes adjacent to Lake Nabugabo—Lake Kayanja, Lake Kayugi, and Lake Manywa—contain large numbers of Oreochromis esculentus and two haplochromine species previously known only in Lake Nabugabo. These three lakes should be designated conservation areas for the cichlids and other native species that were previously abundant in Lakes Victoria, Kyoga, and Nabugabo. The swamps around these lakes should be protected so that the Nile perch does not spread into them. Further research should be carried out to ensure the preservation of the endemic species.     

Patterns of the Remnant Cichlid Fauna in Southern Lake Victoria

During the years 1984–1987 Lake Victoria in East Africa experienced what is probably the largest mass extinction of contemporary vertebrates. Within a decade about 200 endemic species of haplochromine cichlids disappeared. The extinctions that occurred in the 1980s have been documented predominantly on species of offshore and sub-littoral waters in the Mwanza Gulf of southern Lake Victoria. Although the littoral fauna of southern Lake Victoria had not been examined in detail, their diversity seemed less affected by the changes in the ecosystem. We give results of the first comprehensive inventory of the littoral cichlid fauna in southern Lake Victoria and discuss its conservation status. We also report on new developments in the sub-littoral fauna after 1990. More than 50 littoral and 15 sub-littoral stations were sampled between the years 1991 to 1995. Of the littoral stations, 34 were sampled for the first time. One hundred sixty three species of haplochromines were collected. Of these, 102 species were previously unknown. About two thirds of them live in rocky areas that were sampled for the first time. Littoral rocky habitats harbored the highest diversity. Since 1990, however, 13 more species disappeared from established sampling stations in littoral habitats. Fishing practices, spreading of exotic fishes, water hyacinth, and eutrophication are considered important threats to the littoral fauna. Also in the upper sub-littoral the number of species declined further. On deeper sub-littoral mud bottoms individual and species numbers increased again, although they are nowhere close to those found before the Nile perch (Lates niloticus) upsurge. This fauna differs from the well studied pre-Nile perch fauna. At well-established sampling stations, the sub-littoral zone is dominated by previously unknown species, and some known species have performed dramatic habitat shifts.     

Quantitative approaches to the analysis of stable isotope food web data

Ecologists use stable isotopes (delta13C, delta15N) to better understand food webs and explore trophic interactions in ecosystems. Traditionally, delta13C vs. delta15N bi-plots have been used to describe food web structure for a single time period or ecosystem. Comparisons of food webs across time and space are increasing, but development of statistical approaches for testing hypotheses regarding food web change has lagged behind. Here we present statistical methodologies for quantitatively comparing stable isotope food web data. We demonstrate the utility of circular statistics and hypothesis tests for quantifying directional food web differences using two case studies: an arthropod salt marsh community across a habitat gradient and a freshwater fish community from Lake Tahoe, USA, over a 120-year time period. We calculated magnitude and mean angle of change (theta) for each species in food web space using mean delta13C and delta15N of each species as the x, y coordinates. In the coastal salt marsh, arthropod consumers exhibited a significant shift toward dependence on Spartina, progressing from a habitat invaded by Phragmites to a restored Spartina habitat. In Lake Tahoe, we found that all species from the freshwater fish community shifted in the same direction in food web space toward more pelagic-based production with the introduction of nonnative Mysis relicta and onset of cultural eutrophication. Using circular statistics to quantitatively analyze stable isotope food web data, we were able to gain significant insight into patterns and changes in food web structure that were not evident from qualitative comparisons. As more ecologists incorporate a food web perspective into ecosystem analysis, these statistical tools can provide a basis for quantifying directional food web differences from standard isotope data.     

Grazer traits, competition, and carbon sources to a headwater-stream food web

We investigated the effect of grazing by a dominant invertebrate grazer (the caddisfly Glossosoma penitum) on the energy sources used by other consumers in a headwater-stream food web. Stable isotope studies in small, forested streams in northern California have shown that G. penitum larvae derive most of their carbon from algae, despite low algal standing crops. We hypothesized that the caddisfly competes with other primary consumers (including mayflies) for algal food and increases their reliance on terrestrial detritus. Because Glossosoma are abundant and defended from predators by stone cases, their consumption of algal energy may reduce its transfer up the food chain. We removed Glossosoma (natural densities >1000 caddisflies/m2) from five approximately 4 m2) stream sections during the summer of 2000 and measured responses of algae, invertebrate primary consumers, and invertebrate predators. The treatment reduced Glossosoma biomass by 80-90%. We observed a doubling in chlorophyll a per area in sections with reduced Glossosoma abundance and aggregative increases in the biomass of undefended primary consumers. Heptageniid mayfly larvae consumed more algae (as measured by stable carbon isotope ratios and gut content analysis) in caddisfly removal plots at the end of the 60-day experiment, although not after one month. We did not see isotopic evidence of increased algal carbon in invertebrate predators, however. Patterns of caddisfly and mayfly diets in the surrounding watershed suggested that mayfly diets are variable and include algae and detrital carbon in variable proportions, but scraping caddisflies consume primarily algae. Caddisfly and mayfly diets are more similar in larger, more productive streams where the mayflies assimilate more algae. Isotopic analysis, in combination with measurements of macroinvertebrate abundance and biomass in unmanipulated plots, suggested that a substantial portion of the invertebrate community (>50% of biomass) was supported, at least partially, by local algal carbon during midsummer. These data suggest that algae may be more important to community dynamics in headwater streams than their relatively low productivity would suggest. Through their high densities and relative invulnerability to predation, armored grazers may also affect community structure and flow of algal and detrital carbon in headwater streams.     

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.     

Lake to land subsidies: experimental addition of aquatic insects increases terrestrial arthropod densities

Aquatic insects are a common and important subsidy to terrestrial systems, yet little is known about how these inputs affect terrestrial food webs, especially around lakes. Myvatn, a lake in northern Iceland, has extraordinary midge (Chironomidae) emergences that result in large inputs of biomass and nutrients to terrestrial arthropod communities. We simulated this lake-to-land resource pulse by collecting midges from Myvatn and spreading their dried carcasses on 1-m2 plots at a nearby site that receives very little midge deposition. We hypothesized a positive bottom-up response of detritivores that would be transmitted to their predators and would persist into the following year. We sampled the arthropod community once per month for two consecutive summers. Midge addition resulted in significantly different arthropod communities and increased densities of some taxa in both years. Detritivores, specifically Diptera larvae, Collembola, and Acari increased in midge-addition plots, and so did some predators and parasitoids. Arthropod densities were still elevated a year after midge addition, and two years of midge addition further increased the density of higher-order consumers (e.g., Coleoptera and Hymenoptera). Midge addition increased arthropod biomass by 68% after one year and 108% after two years. By manipulating the nutrient pulse delivered by midges we were able to elucidate food web consequences of midge deposition and spatial and temporal dynamics that are difficult to determine based on comparative approaches alone. Resources cross ecosystem boundaries and are assimilated over time because of life-history strategies that connect aquatic and terrestrial food webs and these systems cannot be fully understood in isolation from each other.     

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

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

Habitat provisioning for aboveground predators decreases detritivores

Although provisioning of habitat by ecosystem engineers is one of the most common biological interactions, previous studies have mostly focused on facilitative or bottom-up processes. Here we show that engineering effects can indirectly strengthen top-down effects mediated by predator abundance. We conducted a small-scale manipulative field experiment and broad-scale field observations of the plant, web spider, and detrital insect system in forest understory habitats. In the field experiment, artificially increasing architectural elements enhanced the abundance of spiders by providing physical support for web building. Moreover, aerial insects derived from the detrital food web decreased in response to increased spider abundance. As artificial architecture per se did not affect aerial detritivores, these results indicate that ecosystem engineering indirectly strengthens top-down effects mediated by predators. In field observations conducted in 12 cedar forests, path analyses supported the importance of an indirect pathway originating from understory vegetation complexity to spider abundance and to aerial detritivores. The effect size of spiders on detrital insects was similar in the field experiment and in the observations. These results indicate that the engineering effects of plants cascade to detrital insects through web spiders across different scales.     

Lake of flies, or lake of fish? A trophic model of Lake Malawi

Ecosystem-focused models have, for the first time, become available for the combined demersal and pelagic components of a large tropical lake ecosystem, Lake Malawi. These provide the opportunity to explore continuing controversies over the production efficiencies and ecological functioning of large tropical lakes. In Lake Malawi these models can provide important insight to the effect of fishing on fish composition, and the potential competition that the lakefly Chaoborus edulis may have with fisheries production. A mass-balanced trophic model developed for the demersal fish community of the southern and western areas of Lake Malawi was integrated with an existing trophic model developed for the open-water pelagic. Input parameters for the demersal model were obtained from a survey of fish distributions, fish food consumption studies, and from additional published quantitative and qualitative information on the various biotic components of the community. The model was constructed using the Ecopath approach and software. The graphically presented demersal food web spanned four trophic levels and was based primarily on consumption of detritus, zooplankton and sedimented diatoms. Zooplankton was imported into the system at trophic levels three and four through fish predation on carnivorous and herbivorous copepods and Chaoborus larvae. It is proposed that the primary consumption of copepods was by fish migrating into the pelagic zone. Chaoborus larvae in the demersal were probably consumed near the lakebed as they conducted a daily migration from the pelagic to seek refuge in the sediments. This evidence for strong benthic-pelagic coupling provided the opportunity for linking the demersal model to the existing model for the pelagic community so producing the first model for the complete ecosystem. Energy fluxes through the resulting combined model demonstrated that the primary import of biomass to the demersal system was detritus of pelagic origin (72.1%) and pelagic zooplankton (10.6%). Only 15.8% of the biomass consumed within the demersal system was of demersal origin. Lakefly production is efficiently utilised by the lake fish community, and any attempt to improve fishery production through introduction of a non-native plantivorous fish species would have a negative impact on the stability and productivity of the lake ecosystem.     

Elucidation of ecosystem attributes of an oligotrophic lake in Hokkaido,Japan, using Ecopath with Ecosim (EwE)

The fishing practices in the oligotrophic Lake Toya, Hokkaido, Japan, have profound implications in the ecosystem sustainability. The status of the sockeye salmon (Oncorhynchus nerka) population has become a serious concern among the lake managers and policy makers during the last decades. While the decline of the sockeye salmon population has been well documented in Lake Toya, there is considerable uncertainty with regards to the impact on the broader system dynamics. In this study, our objective is to address this knowledge gap by undertaking a synthesis of the Lake Toya food web using the mass-balance modeling software Ecopath with Ecosim (EwE). Our primary research question is to examine the repercussions of the declining sockeye salmon population on the trophic dynamics of the lake. Namely, we assess if there are any competing species that might have benefited from the decrease of sockeye salmon standing biomass and to what extent do these changes propagate through the Lake Toya food web? Our analysis pinpoints the critical role of the Japanese smelt (Hypomesus transpacificus nipponensis) in the system, which demonstrates a wide range of effects on several functional groups at both higher and lower trophic levels in Lake Toya. In particular, being a substantial portion of the masu salmon (Oncorhynchus masou) and adult sockeye salmon diets, the Japanese smelt has a positive impact on the top predators of the system. Amphipods, insects, and shrimp strongly benefit from the autochthonous and allochthonous organic matter in the system, while the tight coupling between phytoplankton and zooplankton seems to be particularly critical for the integrity of the Lake Toya food web. Whereas the values of the different ecosystem attributes (e.g., primary production/biomass, biomass/total throughput, system omnivory index, amount of recycled throughput, Finn's cycling index) provide evidence that Lake Toya is an immature system, we note that the internal redundancy and the system overhead estimates suggest that the lake possesses substantial reserves to overcome external perturbations. We also examined the effects of a variety of fishing policies on the biomass of masu salmon and adult sockeye salmon, which verify the belief that the adult sockeye population is quite fragile with high likelihood to collapse. Our analysis also predicts that sockeye will not rebound unless the fishing pressure exerted is substantially reduced (>50% of the reference levels used). Masu salmon seems to benefit under all the scenarios examined indicating that the intensity of the current fishing activities is significantly lower than its biomass accumulation rate in the system.     

Predator diversity and identity drive interaction strength and trophic cascades in a food web

Declining predator diversity may drastically affect the biomass and productivity of herbivores and plants. Understanding how changes in predator diversity can propagate through food webs to alter ecosystem function is one of the most challenging ecological research topics today. We studied the effects of predator removal in a simple natural food web in the Sierra Nevada mountains of California (USA). By excluding the predators of the third trophic level of a food web in a full-factorial design, we monitored cascading effects of varying predator diversity and composition on the herbivorous beetle Chrysomela aeneicollis and the willow Salix orestera, which compose the first and second trophic levels of the food web. Decreasing predator diversity increased herbivore biomass and survivorship, and consequently increased the amount of plant biomass consumed via a trophic cascade. Despite this simple linear mean effect of diversity on the strength of the trophic cascade, we found additivity, compensation, and interference in the effects of multiple predators on herbivores and plants. Herbivore survivorship and predator-prey interaction strengths varied with predator diversity, predator identity, and the identity of coexisting predators. Additive effects of predators on herbivores and plants may have been driven by temporal niche separation, whereas compensatory effects and interference occurred among predators with a similar phenology. Together, these results suggest that while the general trends of diversity effects may appear linear and additive, other information about species identity was required to predict the effects of removing individual predators. In a community that is not temporally well-mixed, predator traits such as phenology may help predict impacts of species loss on other species. Information about predator natural history and food web structure may help explain variation in predator diversity effects on trophic cascades and ecosystem function.     

Succession patterns of phytoplankton blooms: directionality and influence of algal cell size

Using four replicate microcosms in the laboratory, we induced a phytoplankton bloom by enclosing a natural community sampled from Masnou Harbor (N.E. Spain) in November 1987, and examined the pattern of algal succession during the bloom. Good replicability of the temporal patterns of the community biomass and the abundance of most species demonstrated that succession was a directional, non-random process. The successional pathway observed (small flagellates » small centric diatoms » small flagellates) resembled that observed by other authors studying phytoplankton blooms. This pattern differed from previous models of algal succession in that dinoflagellates never comprised a substantial fraction of the community biomass, and in that algal cell size did not tend to increase along the successional sequence. Algal cell size, however, was an important determinant of phytoplankton community structure, since it constrained the density, but not the biomass, achievable by the different species. We suggest that there is not a single, general pattern of phytoplankton succession, but that distinction should be made, at least between seasonal and bloom patterns of phytoplankton succession.     

Type of disturbance and ecological history determine structural stability.

This study aims to reveal whether complexity, namely, community and trophic structure, of chronically stressed soil systems is at increased risk or remains stable when confronted with a subsequent disturbance. Therefore, we focused on a grassland with a history of four centuries of patchy contamination. Nematodes were used as model organisms because they are an abundant and trophically diverse group and representative of the soil food web and ecosystem complexity. In a field survey, a relationship between contaminants and community structures was established. Following, two groups of soil mesocosms from the field that differed in contamination level were exposed to different disturbance regimes, namely, to the contaminant zinc and a heat shock. The zinc treatment revealed that community structure is stable, irrespective of soil contamination levels. This implies that centuries of exposure to contamination led to adaptation of the soil nematode community irrespective of the patchy distribution of contaminants. In contrast, the heat shock had adverse effects on species richness in the highly contaminated soils only. The total nematode biomass was lower in the highly contaminated field samples; however, the biomass was not affected by zinc and heat treatments of the mesocosms. This means that density compensation occurred rapidly, i.e., tolerant species quickly replaced sensitive species. Our results support the hypothesis that the history of contamination and the type of disturbance determine the response of communities. Despite that ecosystems may be exposed for centuries to contamination and communities show adaptation, biodiversity in highly contaminated sites is at increased risk when exposed to a different disturbance regime. We discuss how the loss of higher trophic levels from the entire system, such as represented by carnivorous nematodes after the heat shock, accompanied by local biodiversity loss at highly contaminated sites, may result in detrimental effects on ecosystem functions.     

Mass mortality ofDiadema antillarum

It has been hypothesized that herbivorous fishes and the regular echinoidDiadema antillarum Philippi compete for benthic algae as their major food resource. Mass mortality ofD. antillarum in February 1984 provided the opportunity to test the hypothesis that herbivorous fishes and sea urchins were competing previously. Visual censuses of herbivorous fishes conducted over 4 yr in four reef zones on Tague Bay Reef, St. Croix, U.S. Virgin Islands, before and after the mass mortality indicated that population densities increased approximately three-fold in backreef and shallow (2m) forereef zones and two-fold, and four-fold in mid (5m) and deep (10m) forereef zones, respectively. Juvenile parrotfishes constituted the major component of these increases, except in the shallow forereef where acanthurids became most abundant. Grazing intensity by herbivorous fishes increased in three of the four reef zones immediately following the mass mortality. These data support the hypothesis that exploitative competition for algal resources was occurring prior to the sea urchin mass-mortality, although alternative hypotheses cannot be discounted completely. Despite the increases in the abundances of, and grazing by, herbivorous fishes, the algal community continued to increase in percent cover and biomass, indicating that increased grazing by fishes does not compensate for the loss of grazing byD. antillarum in controlling algal abundance and community structure.     

Evidence for the dominance of indirect effects in 50 trophic ecosystem networks

Indirect effects are powerful influences in ecosystems that may maintain species diversity and alter apparent relationships between species in surprising ways. Here, we applied network environ analysis to 50 empirically-based trophic ecosystem models to test the hypothesis that indirect flows dominate direct flows in ecosystem networks. Further, we used Monte Carlo based perturbations to investigate the robustness of these results to potential error in the underlying data. To explain our findings, we further investigated the importance of the microbial food web in recycling energy-matter using components of the Finn Cycling Index and analysis of environ centrality. We found that indirect flows dominate direct flows in 37/50 (74.0%) models. This increases to 31/35 (88.5%) models when we consider only models that have cycling structure and a representation of the microbial food web. The uncertainty analysis reveals that there is less error in the I/D values than the ±5% error introduced into the models, suggesting the results are robust to uncertainty. Our results show that the microbial food web mediates a substantial percentage of cycling in some systems (median = 30.2%), but its role is highly variable in these models, in agreement with the literature. Our results, combined with previous work, strongly suggest that indirect effects are dominant components of activity in ecosystems.     

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.     

Quantity and quality: unifying food web and ecosystem perspectives on the role of resource subsidies in freshwaters

Although the study of resource subsidies has emerged as a key topic in both ecosystem and food web ecology, the dialogue over their role has been limited by separate approaches that emphasize either subsidy quantity or quality. Considering quantity and quality together may provide a simple, but previously unexplored, framework for identifying the mechanisms that govern the importance of subsidies for recipient food webs and ecosystems. Using a literature review of > 90 studies of open-water metabolism in lakes and streams, we show that high-flux, low-quality subsidies can drive freshwater ecosystem dynamics. Because most of these ecosystems are net heterotrophic, allochthonous inputs must subsidize respiration. Second, using a literature review of subsidy quality and use, we demonstrate that animals select for high-quality food resources in proportions greater than would be predicted based on food quantity, and regardless of allochthonous or autochthonous origin. This finding suggests that low-flux, high-quality subsidies may be selected for by animals, and in turn may disproportionately affect food web and ecosystem processes (e.g., animal production, trophic energy or organic matter flow, trophic cascades). We then synthesize and review approaches that evaluate the role of subsidies and explicitly merge ecosystem and food web perspectives by placing food web measurements in the context of ecosystem budgets, by comparing trophic and ecosystem production and fluxes, and by constructing flow food webs. These tools can and should be used to address future questions about subsidies, such as the relative importance of subsidies to different trophic levels and how subsidies may maintain or disrupt ecosystem stability and food web interactions.     

Structure and function of a warm monomictic lake

Construction and simulation of a model of Lake Conway, Florida, U.S.A., provided a framework for defining major characteristics of this ecosystem. The relationships that are formalized in this model comprise a set of hypotheses about the nature of a warm monomictic lake. The data that were used to parameterize the model came primarily from literature estimates, although approximations of biomass levels were available from associated research conducted on the lake.Submersed macrophytes are a major biomass component in Lake Conway; simulation suggested that their role in nutrient recycling overshadows their importance in the grazing food chain. Phytoplankton biomass and degree of fluctuation are considerably lower than are observed in most cool temperate lakes, although simulated respiration and herbivory rates are closer to temperate values than tropical values. Simulated epipelic algae biomass varies an order of magnitude during the year, and this group appears to be a significant part of the food chain. Simulated zooplankton consumption and turnover rates are very high, in part because of the relatively small biomass per individual. Simulation of the model suggests that slightly more carbon is processed through the grazing food chain in Lake Conway than through the detritus food chain.