Modeling the role of macroalgae in a shallow sub-estuary of Narragansett Bay, RI (USA)

Proliferation of macroalgal mats is a frequent consequence of nutrient-driven eutrophication in shallow, photic coastal marine ecosystems. These macroalgae have the potential to significantly modify water quality, plankton productivity, nutrient cycling, and dissolved oxygen dynamics. We developed a model for Ulva lactuca and Gracilaria tikvahiae in Greenwich Bay, RI (USA), a shallow sub-estuary of Narragansett Bay, as part of a larger estuarine ecosystem model. The model predicts the biomass of both species in units of carbon, nitrogen, and phosphorus as a function of primary production, respiration, grazing, decay, and physical exchange, with particular attention to the effects of biomass layering on light attenuation and suppression of metabolic rates. The model successfully reproduced the magnitude and seasonal cycle of area-weighted and peak biomass in Greenwich Bay along with tissue C:N ratios, and highlighted the importance of grazing and inclusion of self-limitation primarily in the form of self-shading to overcome an order of magnitude difference in rates of production and respiration. Inclusion of luxury nutrient uptake demonstrated the importance of internal nutrient storage in fueling production when nutrients are limiting. Macroalgae were predicted to contribute a small fraction of total system primary production and their removal had little effect on predicted water quality. Despite a lack of data for calibration and a fair amount of sensitivity to individual parameter values, which highlights the need for further autecological studies to constrain formulations, the model successfully predicted macroalgal biomass dynamics and their role in ecosystem functioning. Our formulations should be exportable to other temperate systems where macroalgae occur in abundance.     

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.     

Ecosystem modeling of waterhyacinth management in Lake Alice,Florida

A mathematical model was developed for Lake Alice, a small, eutrophic, Kars-topography lake in Gainesville, Florida. The management of the floating aquatic plant, the waterhyacinth (Eichhornia crassipes) was considered one of the main objectives of the model. Simulated diversion of the treated sewage inflow reduced waterhyacinths by 50% or more. Dissolved oxygen remained low. Simulated diversion of the diluting effect of a heating plant effluent led to complete hyacinth takeover and much greater seasonal oscillations in the chemical parameters. Simulated hyacinth control caused the greatest dissolved oxygen fluctuations when spraying and periodic harvesting were tested and least with continuous harvesting. It was suggested that the above mentioned diversions were shown to be ecologically unattractive even without considering the economic costs of alternatives to lake disposal.     

Energy transformations in two tropical Oxbow lakes

The flow and pattern of energy utilization in two tropical ecosystems where polyculture of fishes was practiced were studied. The rates of organic production made by the two tropical natural aquatic systems (oxbow lakes) were studied for two annual cycles. The study of utilization of energy in two tropical lakes revealed that herbivores were more efficient in energy utilization than autotrophs. The energy available in these systems was supplied by allochthonous source. Silver carp and grass carp were found to be most efficient in conversion of energy in the grazing food chain of the lake. The efficiency of the system to fish production was found to be 0.06 (OL-I) ?0.05% (OL-II) of the total available solar radiation. Various forms of life are all accompanied by energy changes, in the studied lakes were found to form a food web.     

Cascading Effects of the Introduced Nile Perch on the Detritivorous/Phytoplanktivorous Species in the Sublittoral Areas of Lake Victoria

In the 1980s an explosive increase of the introduced Nile perch ( Lates sp.; Harrison 1991) in Lake Victoria caused the destruction of approximately 65% of the endemic haplochromine cichlids. The eradication of approximately 200 vertebrate species in less than a decade may well represent the largest extinction event among vertebrates during this century. The introduction of the Nile perch can be considered as a large-scale, albeit unintended, experiment. Farreaching changes in the food web are taking place. We present data on the importance of haplochromines in the ecosystem prior to the Nile perch boom. An analysis of the pelagic community in the sublittoral area of the Mwanza Gulf revealed that the phytoplanktivores formed 18% of the biomass of the total haplochromine community. In the benthic community, the detritivores that frequently included phytoplankton in their diet comprised 31% of the biomass. We examine the hypothesis that algal grazing was reduced by the disappearance of haplochromine phytoplanktivores and detritivores. The disappearance of these groups may have contributed to the recent algal blooms. To investigate this hypothesis we consider a second major change in the system. Coinciding with the decrease of the haplochromines, the number of atyid prawns, Caridina nilotica , has strongly increased. We present preliminary evidence that the stock of the detritivorous haplochromines that formerly comprised most of the demersal ichthyomass has (partially?) been replaced by the prawn. We discuss the possible mechanisms underlying this major change in the food web. With respect to the conservation of the Great African Lakes, tbe collapse of Lake Victoria's ecosystem should serve as an example of how easily a complex ecosystem can be irreversibly destroyed.     

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.     

Study of the nutrient and plankton dynamics in Lake Tanganyika using a reduced-gravity model

An eco-hydrodynamic (ECOH) model is proposed for Lake Tanganyika to study the plankton productivity. The hydrodynamic sub-model solves the non-linear, reduced-gravity equations in which wind is the dominant forcing. The ecological sub-model for the epilimnion comprises nutrients, primary production, phytoplankton biomass and zooplankton biomass. In the absence of significant terrestrial input of nutrients, the nutrient loss is compensated for by seasonal, wind-driven, turbulent entrainment of nutrient-rich hypolimnion water into the epilimnion, which gives rise to high plankton productivity twice in the year, during the transition between two seasons. Model simulations predict well the seasonal contrasts of the measured physical and ecological parameters. Numerical tests indicate that the half saturation constant for grazing by zooplankton and the fish predation rate on zooplankton affect the zooplankton biomass measurably more than that of phytoplankton biomass. This work has implications for the application of this model to predict the climatological biological productivity of Lake Tanganyika.     

固城湖生物资源现状及近20年间的变化趋势

1999年对固城湖渔业生物资源进行了周年调查,结合1981年4月至1982年3月、1987年6月至1988年5月的2次调查资料,比较分析了固城湖生物资源的动态。研究结果表明,从1981年至1999年近20a间,固城湖浮游植物数量增加44.8倍,生物量增加40.6倍,生物多样性减少。浮游动物数量较20世纪80年代下降约50%,生物量高于80年代初期,低于80年代后期。底栖动物生物量下降,其中瓣鳃类、腹足类生物量下降明显,而寡毛类和摇蚊幼虫等数量与生物量增长显著。固城湖水生植物演替剧烈,水生植物群落结构发生了重大变化,已由苦草、轮叶黑藻群落演变为微齿眼子菜、菹草群落,渔业利用价值下降。每年大量水草腐烂沉积,使固城湖水质呈现不断下降趋势。指出加强对固城湖生物资源的调控和综合利用,是固城湖生态保护和建设的关键。     

附着生物对沉水植物伊乐藻生长的研究

以沉水植物伊乐藻(Elodea nuttalii(Plant.)ST.John.)为研究对象,室内利用原位湖水进行培养,通过测定伊乐藻的生物量、叶绿素质量分数、光合作用速率等指标,研究了附着生物量的增加对沉水植物伊乐藻的影响。结果表明,随着附着生物处理量的增加,伊乐藻的生物量、叶绿素质量分数以及光合作用速率随之下降,50d后,各处理的生物量与对照相比分别减少了约5%~15%,叶绿素质量分数以及光合作用速率下降了20%~43%、10%~36%。同时也分析了丝状藻对沉水植物伊乐藻的遮阴作用,初步推测附着生物对沉水植物伊乐藻的影响可能是由于附着生物的遮荫作用。研究的结果为揭示富营养化湖泊沉水植被衰退的机理提供了理论依据。     

The role of ciliates,heterotrophic dinoflagellates and copepods in structuring spring plankton communities at Helgoland Roads,North Sea

Mesocosm experiments coupled with dilution grazing experiments were carried out during the phytoplankton spring bloom 2009. The interactions between phytoplankton, microzooplankton and copepods were investigated using natural plankton communities obtained from Helgoland Roads (54°11.3′N; 7°54.0′E), North Sea. In the absence of mesozooplankton grazers, the microzooplankton rapidly responded to different prey availabilities; this was most pronounced for ciliates such as strombidiids and strobilids. The occurrence of ciliates was strongly dependent on specific prey and abrupt losses in their relative importance with the disappearance of their prey were observed. Thecate and athecate dinoflagellates had a broader food spectrum and slower reaction times compared with ciliates. In general, high microzooplankton potential grazing impacts with an average consumption of 120% of the phytoplankton production (P _p ) were measured. Thus, the decline in phytoplankton biomass could be mainly attributed to an intense grazing by microzooplankton. Copepods were less important phytoplankton grazers consuming on average only 47% of P _p . Microzooplankton in turn contributed a substantial part to the copepods’ diets especially with decreasing quality of phytoplankton food due to nutrient limitation over the course of the bloom. Copepod grazing rates exceeded microzooplankton growth, suggesting their strong top-down control potential on microzooplankton in the field. Selective grazing by microzooplankton was an important factor for stabilising a bloom of less-preferred diatom species in our mesocosms with specific species (Thalassiosira spp., Rhizosolenia spp. and Chaetoceros spp.) dominating the bloom. This study demonstrates the importance of microzooplankton grazers for structuring and controlling phytoplankton spring blooms in temperate waters and the important role of copepods as top-down regulators of microzooplankton.     

Effect of sheep grazing and fire on sage grouse populations in southeastern Idaho

This paper describes the development, evaluation, and use of a model that simulates the effect of grazing and fire on temporal and spatial aspects of sagebrush community vegetation and sage grouse population dynamics. The model is represented mathematically as a discrete-time, stochastic compartment model based on difference equations with a time interval of 1 week. In the model, sheep graze through sage grouse breeding habitat during spring and fall, and different portions of the area can burn at different frequencies, creating a habitat mosaic of burned and unburned areas.The model was evaluated by examining predictions of (1) growth of sagebrush canopy cover after fire, (2) seasonal dynamics of grass and forb biomass under historical environmental conditions, and (3) sage grouse population dynamics associated with selected sagebrush canopy covers. Simulated changes in sagebrush canopy cover following fire correspond well with qualitative reports of long-term trends, simulated seasonal dynamics of herbaceous biomass correspond well with field data, and simulated responses of sage grouse population size and age structure to changing sagebrush canopy cover correspond well to qualitative field observations.Simulation results suggest that large fires occurring at high frequencies may lead to the extinction of sage grouse populations, whereas fires occurring at low frequencies may benefit sage grouse if burned areas are small and sheep grazing is absent. Sheep grazing may contribute to sage grouse population decline, but is unlikely to cause extinction under fire regimes that are favorable to sage grouse.     

Adapting empirical equations to Lake Kinneret data by using three calibration methods

This paper examines whether the relationships between a number of characteristic limnological variables (suspended particulate matter, turbidity, Secchi depth, light attenuation, and chlorophyll a) determined for temperate lakes are consistent with the relationships found in Mediterranean lakes such as Lake Kinneret. We found that the use of published relationships between lake variables may lead to erroneous results when applied indiscriminately to other lake types.     

洞庭湖浮游植物增长的限制性营养元素研究

近20年水质监测资料表明,洞庭湖水体富营养化日趋严重。洞庭湖水体主要污染物为氮和磷,而营养盐赋存形态及其含量对浮游植物生长的影响在洞庭湖尚未见报道。2011年9月至2012年8月对洞庭湖浮游植物生物量及主要营养盐赋存形态与含量进行监测,同时利用藻类增长的生物学（NEB）评价方法对限制浮游植物增长的营养盐进行了研究,并分析了浮游植物生物量与各营养元素之间的相关性。结果表明：洞庭湖主要污染物总氮（TN）和总磷（TP）的年平均值分别为1.90 mg·L-1和0.093 mg·L-1,溶解态无机氮（DIN）平均占ρ（TN）比例为87%,溶解态总磷（DTP）平均占ρ（TP）比例为70%。洞庭湖水体中,DIN是TN的主要贡献者,且不同形态DIN的贡献大小依次为ρ（NO3--N）〉ρ（NH4＋-N）〉ρ（NO2--N）;磷形态组成中,TP主要以溶解反应性磷（SRP）存在。春季洞庭湖水体中ρ（TN）、ρ（TP）较高,这一结果可能源于春季面源污染。洞庭湖水体中ρ（Chla）与氮显著正相关,与磷显著负相关。NEB 实验结果表明氮对洞庭湖浮游植物生长有明显的促进作用,其幅度随氮浓度的增加而加强,而磷对浮游植物的生长影响不大,有时出现抑制作用,硝态氮与磷之间不存在交互作用。因此,氮可能是洞庭湖浮游植物增长的主要限制性营养因子,这一研究暗示在洞庭湖富营养化控制过程中应特别注重氮的控制。     

放牧管理对藏北高寒沼泽草甸群落结构的影响

采用样带法研究了冬季放牧、冬春季放牧和全年放牧3种放牧管理模式对西藏高寒沼泽草甸高度、盖度、地上生物量及群落结构的影响.研究表明,3种放牧管理模式下的高寒沼泽草甸植被群落物种数差异不显著（p〉0.05）,高度、盖度和地上生物量差异显著（p〈0.05）。冬季放牧、冬春季放牧和全年放牧沼泽草甸地上生物量依次降低。放牧管理对沼泽草甸群落多样性影响较大,不同放牧管理模式下的沼泽草甸Shannon-Wiener指数差异显著（p〈0.05）,冬春放牧沼泽草甸Shannon-Wiener指数较冬季放牧和全年放牧沼泽草甸低。冬季放牧和全年放牧沼泽草甸Margalef指数和Pielou指数差异不显著,显著（p〈0.05）高于冬春季放牧沼泽草甸。     

The annual cycle of protozooplankton in the Kiel Bight

Protozooplankton (heterotrophic dinoflagellates and ciliates) composition and biomass was studied in a 20-m water column in the Kiel Bight on 44 occasions between January 1973 and April 1974. Both groups attained comparable biomass maxima during spring and autumn (0.3 to 0.7 g C m^-2 in the 20-m water column) and biomass levels were much lower in summer and lowest in winter. The spring protozooplankton maximum coincided with that of phytoplankton and during the rest of the year, protozooplankton stocks did not appear to be food limited as phytoplankton stocks were large throughout; many protozoans with ingested microplankton cells were observed, indicating that their potential food supply is not restricted to nanoplankton. Non-loricate organisms dominated biomass of the ciliates and tintinnids were of little importance. Tintinnids predominated in plankton samples concentrated by 20 m gauze indicating that most non-loricate ciliates, irrespective of size, were not retained. When phytoplankton sotcks were large (>3 g C m^-2) but those of metazooplankton small, as in spring and autumn, protozooplankton were the major herbivores with biomass levels comparable to those attained in summer by metazooplankton ( 0.5 g C m^-2). A highly significant negative correlation was found between protozooplankton and metazooplankton during the plankton growth season. Predation by the latter is thus an important factor regulating size of the protozooplankton population, although other factors also appear to be in operation. Loss rates of the pelagic system through sedimentation are highest in spring and autumn when protozooplankton dominate the grazing community and loss rates are much lower in summer when metazooplankton are the dominant herbivores. Apparently, the impact of protozooplankton grazing on the pelagic system is quite different to that of the metazooplankton.Publication No. 268 of the Joint Research Programme (SFB 95), Kiel University     

Dynamic model of Lake Chozas (León, NW Spain)—Decrease in eco-exergy from clear to turbid phase due to introduction of exotic crayfish

We developed a dynamic model of the phosphorus cycle in Lake Chozas, a small shallow water body in León (NW Spain). The calibrated model simulated seasonal dynamics of phosphorus concentrations in major components of the lake's ecological network before and after 1997, the year when an invasive allochthonous crustacean, the Louisiana red swamp crayfish (Procambarus clarkii), was introduced into the lake. The shift from clean to turbid phase, due to grazing by crayfish on submerged vegetation, caused a gradual decrease in eco-exergy, reflecting an increase in entropy, related to breakdown of ecosystem internal equilibria. This case study verifies the hypothesis of Marchi et al. (2010) that, after an initial relatively stable state, the allochthonous species may cause an increase in entropy indicating perturbation of the ecosystem.     

Lake phosphorus dynamics and climate warming: A mechanistic model approach

Model results in this work indicate that lakes may respond very differently to climate change depending on their physical character. A physical lake model and a mechanistic phosphorus model are combined with two temperature scenarios generated by a regional climate model (RCM) in three sites in central Sweden—Lake Erken and two basins of Lake Mälaren (Galten and Ekoln). In the phosphorus model water mixing, mineralization, diffusion and biouptake are temperature dependent. In the simulations, Lake Erken is much more sensitive to climate warming than the two basins of Lake Mälaren, and the reason is shown to be the much longer water residence time in Lake Erken (7 years), stressing the importance of internal lake processes. In Galten and Ekoln the water residence times are less than 1 year, and the effects of water temperature changes are small. In Lake Erken the concentration of epilimnetic-dissolved phosphorus is almost doubled in spring and autumn in the warmest climate scenario. Since the lake is mostly phosphorus limited, this means that the potential for phytoplankton production is almost doubled. The implication would be that in Lake Erken, and in other eutrophic lakes with long water residence times, eutrophication problems may become serious in the future, and that managers may need to take action today in order to maintain good water quality in these lakes.     

Structural changes in lake functioning induced from nutrient loading and climate variability

Climate variability is increasingly recognized as an important regulatory factor, capable of influencing the structural properties of aquatic ecosystems. Lakes appear to be particularly sensitive to the ecological impacts of climate variability, and several long time series have shown a close coupling between climate, lake thermal properties and individual organism physiology, population abundance, community structure, and food web dynamics. Thus, understanding the complex interplay among meteorological forcing, hydrological variability, and ecosystem functioning is essential for improving the credibility of model-based water resources/fisheries management. Our objective herein is to examine the relative importance of the ecological mechanisms underlying plankton seasonal variability in Lake Washington, Washington State (USA), over a 35-year period (1964–1998). Our analysis is founded upon an intermediate complexity plankton model that is used to reproduce the limiting nutrient (phosphate)–phytoplankton–zooplankton–detritus (particulate phosphorus) dynamics in the lake. Model parameterization is based on a Bayesian calibration scheme that offers insights into the degree of information the data contain about model inputs and allows obtaining predictions along with uncertainty bounds for modeled output variables. The model accurately reproduces the key seasonal planktonic patterns in Lake Washington and provides realistic estimates of predictive uncertainty for water quality variables of environmental management interest. A principal component analysis of the annual estimates of the underlying ecological processes highlighted the significant role of the phosphorus recycling stemming from the zooplankton excretion on the planktonic food web variability. We also identified a moderately significant signature of the local climatic conditions (air temperature) on phytoplankton growth (r = 0.41), herbivorous grazing (r = 0.38), and detritus mineralization (r = 0.39). Our study seeks linkages with the conceptual food web model proposed by Hampton et al. [Hampton, S.E., Scheuerell, M.D., Schindler, D.E., 2006b. Coalescence in the Lake Washington story: interaction strengths in a planktonic food web. Limnol. Oceanogr. 51, 2042–2051.] to emphasize the “bottom-up” control of the Lake Washington plankton phenology. The posterior predictive distributions of the plankton model are also used to assess the exceedance frequency and confidence of compliance with total phosphorus (15 μg L⁻¹) and chlorophyll a (4 μg L⁻¹) threshold levels during the summer-stratified period in Lake Washington. Finally, we conclude by underscoring the importance of explicitly acknowledging the uncertainty in ecological forecasts to the management of freshwater ecosystems under a changing global environment.     

Carbon-based balanced trophic structure and flows in the offshore Lake Ontario food web before (1987-1991) and after (2001-2005) invasion-induced ecosystem change

Replicate mass-balanced solutions to Ecopath models describing carbon-based trophic structures and flows were developed for the Lake Ontario offshore food web before and after invasion-induced disruption. The food webs link two pathways of energy and matter flow: the grazing chain (phytoplankton-zooplankton-fish) and the microbial loop (bacteria-protozoans) and include 19 species-groups and three detrital groups. Mass-balance was achieved by using constrained optimization techniques to randomly vary initial estimates of biomass and diet composition. After the invasion, production declined for all trophic levels and species-groups except Chinook salmon. The trophic level (TL) increased for smelt, adult sculpin, adult alewife and Chinook salmon. Changes to ecotrophic efficiencies indicate a reduction in phytoplankton grazing, increased predation pressure on Mysis, adult smelt and alewife and decreased predation pressure on protozoans. Specific resource to consumer TTE changed; increasing for protozoans (8.0-11.5%), Mysis (0.6-1.0%), and Chinook salmon (1.0-2.3%) and other salmonines (0.4-0.5%) and decreasing for zooplankton (20.2-15.1%), prey-fish (9.7-8.8%), and benthos (1.7-0.6%). Direct trophic influences of recent invasive species were low. The synchrony of the decline in PP and species-group production indicates strong bottom-up influence. Mass balance required an increase of two to threefold in lower trophic level biomass and production, confirming a previously observed paradoxical deficit in lower trophic level production. Analysis of food web changes suggest hypotheses that may apply to other similar large pelagic systems including, (1) as pelagic primary productivity declines, overgrazing of zooplankton results in an increase in protozoan production and a loss of trophic transfer efficiency, (2) habitat and food web changes increased Mysis predation on Diporeia and contributed to their recent decline, and (3) production of Chinook salmon, the primary piscivore, was uncoupled from pelagic production processes. This study demonstrates the value of food web models to better understand the impact of invasive species and to develop novel hypotheses concerning trophic influences.