Zebra mussels mediate benthic–pelagic coupling by biodeposition and changing detrital stoichiometry

1. The zebra mussel ( Dreissena polymorpha ) is one of the most successful invasive species; it has colonised many aquatic systems in Europe and North America with strong impacts on various ecosystem processes. The effect of D. polymorpha filtration on pelagic seston concentrations has been quantified in several studies, but the magnitude and stoichiometry of the transfer of sestonic biomass into benthic detritus by D. polymorpha and the accompanying enrichment of the benthic habitat is still under-investigated.
2. We studied biodeposition by zebra mussels in two series of laboratory experiments with the food algae Cryptomonas erosa and Scenedesmus obliquus . We also measured the year-round biodeposition rate under natural conditions in the oligotrophic Lake Constance.
3. In all experiments, zebra mussel biodeposition was linearly related to seston concentration. In the field, the relationship changed with a seasonal shift in algal composition and lower biodeposition rates during the spring algal bloom.
4. For both algal species in laboratory experiments, biodeposited material was depleted in phosphorous at an algal concentration ≤0.6 mg ash-free dry mass L⁻¹, but not at higher concentrations. This effect was not observed in the field, probably because of high variation in C : N : P stoichiometry.
5. By mediating the transfer of pelagic resources into the benthos zebra mussels provide a sufficient amount of detritus for benthic invertebrates, especially during summer. Thus, material biodeposited by the mussels might increase benthic secondary production from pelagic resources, and zebra mussels are important mediators of this flux of organic matter from the pelagic zone into the benthos.     

Northern map turtles (Graptemys geographica) derive energy from the pelagic pathway through predation on zebra mussels (Dreissena polymorpha)

1. Zebra mussels ( Dreissena polymorpha ) derive their energy from the pelagic energy pathway by filtering plankton. Because zebra mussels occur in high densities in littoral habitats, they potentially constitute an important trophic link between littoral consumers and pelagic energy sources. Northern map turtles ( Graptemys geographica ) are widespread in North America and consume zebra mussels.
2. We used stable isotopes analyses to quantify the flow of energy from the pelagic pathway to northern map turtles and to infer the contribution of zebra mussels to map turtle biomass. We then built a bioenergetic model to estimate the annual intake of zebra mussels by northern map turtles in Lake Opinicon, Ontario, Canada.
3. Stable isotopes analyses indicated that zebra mussels constitute between 0% and 14% of the diet of males and between 4% and 36% of the diet of females. Assuming that zebra mussels account for all of the pelagic contribution, we estimated that map turtles consume 3200 kg of zebra mussels annually. Because female map turtles are much larger than males and consume more zebra mussels, they are responsible for 95% of the zebra mussel biomass ingested annually.
4. The pelagic pathway supports an important part of the standing crop biomass of map turtles in Lake Opinicon. We highlight the importance of freshwater turtles in lake ecosystems. Unravelling the trophic interactions mediated by freshwater turtles will lead to a more integrated picture of lake ecosystems.     

Bottom‐up and top‐down effects of browning and warming on shallow lake food webs

Productivity and trophic structure of aquatic ecosystems result from a complex interplay of bottom‐up and top‐down forces that operate across benthic and pelagic food web compartments. Projected global changes urge the question how this interplay will be affected by browning (increasing input of terrestrial dissolved organic matter), nutrient enrichment and warming. We explored this with a process‐based model of a shallow lake food web consisting of benthic and pelagic components (abiotic resources, primary producers, grazers, carnivores), and compared model expectations with the results of a browning and warming experiment in nutrient‐poor ponds harboring a boreal lake community. Under low nutrient conditions, the model makes three major predictions. (a) Browning reduces light and increases nutrient supply; this decreases benthic and increases pelagic production, gradually shifting productivity from the benthic to the pelagic habitat. (b) Because of active habitat choice, fish exert top‐down control on grazers and benefit primary producers primarily in the more productive of the two habitats. (c) Warming relaxes top‐down control of grazers by fish and decreases primary producer biomass, but effects of warming are generally small compared to effects of browning and nutrient supply. Experimental results were consistent with most model predictions for browning: light penetration, benthic algal production, and zoobenthos biomass decreased, and pelagic nutrients and pelagic algal production increased with browning. Also consistent with expectations, warming had negative effects on benthic and pelagic algal biomass and weak effects on algal production and zoobenthos and zooplankton biomass. Inconsistent with expectations, browning had no effect on zooplankton and warming effects on fish depended on browning. The model is applicable also to nutrient‐rich systems, and we propose that it is a useful tool for the exploration of the consequences of different climate change scenarios for productivity and food web dynamics in shallow lakes, the worldwide most common lake type.     

Effects of invasive zebra mussels on phytoplankton,turbidity, and dissolved nutrients in reservoirs

Few experiments have quantified the effects of invasive zebra mussels (Dreissena polymorpha) on man-made reservoirs relative to other aquatic habitats. Reservoirs, however, are the dominate water body type in many of the states that are at the current front of the zebra mussel invasion into the western United States. The objective of this research, therefore, was to determine how zebra mussels affected phytoplankton, turbidity, and dissolved nutrients in water that was collected from three Kansas reservoirs that varied in trophic state (mesotrophic to hypereutrophic), but all experienced frequent cyanobacterial blooms. Laboratory mesocosm experiments were conducted to document the effects of zebra mussels on cyanobacteria and general water quality characteristics in the reservoir water. Zebra mussels significantly reduced algal biomass, and the total biovolume of cyanobacteria (communities were dominated by Anabaena) in each reservoir experiment. The effects of zebra mussels on other major algal groups (diatoms, flagellates, and green algae) and algal diversity were less consistent and varied between the three reservoir experiments. Similarly, the effects of zebra mussels on nutrient concentrations varied between experiments. Zebra mussels increased dissolved phosphorus concentrations in two of the reservoir experiments, but there was no effect of zebra mussels on dissolved phosphorus in the mesotrophic reservoir experiment. Combined, our results strongly suggest that zebra mussels have the potential to significantly impact reservoirs as they continue to expand throughout the western United States. Moreover, the magnitude of these effects may be context dependent and vary depending on the trophic state and/or resident phytoplankton communities of individual reservoirs as has similarly been reported for natural lakes.     

Stoichiometry of regenerated nutrients differs between native and invasive freshwater mussels with implications for algal growth

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Foods of <Emphasis Type="Italic">Velella velella</Emphasis> (Cnidaria: Hydrozoa) in algal rafts and its distribution in Irish seas

The pleustonic hydrozoan, Velella velella, occurs throughout tropical to cold-temperate oceans of the world and sometimes are stranded in masses along hundreds of kilometers of beaches. In June 2009, we encountered algal rafts in the Celtic Sea containing many V. velella that we immediately preserved for gut content analysis. Available prey were enumerated from raft-associated fauna and zooplankton sampled nearby. The identifiable prey (331) in V. velella comprised 78% raft-associated prey (primarily harpacticoid copepods, cumaceans, small fish) and 22% pelagic prey (calanoid copepods, barnacle nauplii, fish eggs). Comparison of ingested with available prey showed selection for fish eggs and small fish, among others; therefore, the null hypothesis, that V. velella consumed all available prey equally, was rejected. Transport by wind and water concentrate Velella spp. in convergences with algal rafts, which suggests that they are important predators of raft—as well as pelagic fauna. A visual survey in September 2004 across the Celtic Sea and beach-stranding data show that V. velella is very abundant in Irish waters at times. Its circumpolar abundance, consumption of pelagic prey and production from symbiotic zooxanthellae, and mass deposition on beaches suggest that V. velella is important in open-ocean carbon cycling and in transport of pelagic production to landmasses.     

Effects of zebra mussels on phytoplankton and ciliates: a field mesocosm experiment

Many observational studies in North American lakes have documenteddecreases in phytoplankton abundance after the invasion of thezebra mussel (Dreissena polymorpha). However, few field experimentshave examined in detail the effect of zebra mussels on phytoplanktonabundance and species composition over an extended period. Replicatedin situ mesocosms were used to evaluate the impact of naturaldensities of zebra mussels on phytoplankton and ciliate biovolume,and algal species composition over a 5-week period in a habitatthat lacked extant mussel populations. Mussel biomass used inthe experiment was determined using a regression model basedon a data analysis that predicts zebra mussel biomass from totalphosphorus concentration. Within 1 week, zebra mussels decreasedphytoplankton biovolume by 53% and ciliate biovolume by 71%.The effect of zebra mussels on ciliate biovolume was sustainedthroughout the study. However, the effect of zebra mussels onphytoplankton abundance gradually waned over the remaining 4weeks of the experiment, such that the declining effect of zebramussels could not be explained by a shift towards less edibleand/or faster growing algal species. The mussels’ decliningcondition could help to explain the effect observed over thecourse of the experiment.     

Increased Benthic Algal Primary Production in Response to the Invasive Zebra Mussel (Dreissena polymorpha) in a Productive Ecosystem, Oneida Lake, New York

Increased water clarity associated with zebra mussel (Dreissena polymorpha) populations may favor benthic algal primary production in freshwater systems previously dominated by pelagic phytoplankton production. While zebra mussel-mediated water clarity effects on benthic primary production have been implicated in published reports, few production estimates are available. This study estimates benthic primary production in Oneida Lake, NY before and after zebra mussel invasion (1992), using measured photosynthetic parameters ( , α^B and β) from sampled benthic algal communities. In the summers of 2003 and 2004, primary production was measured as O₂ evolution from algal communities on hard (cobble) and soft (sediment) substrate from several depths. We also backcast estimates of benthic primary production from measurements of light penetration since 1975. Estimates of whole-lake epipelic and epilithic algal primary production showed a significant (4%) increase and exhibited significantly less interannual variability subsequent to the establishment of zebra mussels. We applied our model to two lakes of differing trophic status; the model significantly overestimated benthic primary production in a hypereutrophic lake, but there was no significant difference between the actual and predicted primary production values in the oligotrophic lake. The hypereutrophic lake had higher zebra mussel densities than Oneida (224 vs. 41 per sample respectively). Though total community respiration (measured in total darkness) was factored into our model predictions of production, our model may need modification when heterotrophic respiration is a large portion of total community metabolism.     

Response of Tetraselmis suecica to nutrient and grazer manipulation

Three methods of algal quantification (direct cell counts, chlorophyll a extraction, in vivo fluorescence) were used to evaluate the response of the unicellular green flagellate Tetraselmis suecica to nutrients and grazers. Nutrient enrichment enhanced total cell counts, chlorophyll a concentration and in vivo and DCMU-fluorescence. Photosynthetic efficiency was reduced in the complete F2 medium as indicated by the high level of in vivo fluorescence, whereas photosynthetic efficiency was increased by the introduction of mussels to the F2 medium. The addition of mussels significantly increased the proportion of non-motile cells, but did not reduce the total cell count. The effect of mussel grazing on algae could be underestimated if only total cells were counted or only the chlorophyll a concentration was measured. The results indicate that these three methods measure different properties of an algal culture and are complementary to each other in assessing the quality and quantity of an algal population. Direct algal counting offers a reliable numerical assessment for cell population abundance. Chlorophyll a concentration was closely correlated to the total cell count. In the presence of mussels, in vivo fluorescence did not correlate with either algal cell counts or chlorophyll a concentration, indicating that the measurement of in vivo fluorescence may be misleading for estimating algal abundance under different culture conditions. This revised version was published online in August 2006 with corrections to the Cover Date.     

INVERTEBRATE‐MEDIATED NUTRIENT LOADING INCREASES GROWTH OF AN INTERTIDAL MACROALGA1

Even in nitrogen‐replete ecosystems, microhabitats exist where local‐scale nutrient limitation occurs. For example, coastal waters of the northeastern Pacific Ocean are characterized by high nitrate concentrations associated with upwelling. However, macroalgae living in high‐zone tide pools on adjacent rocky shores are isolated from this upwelled nitrate for extended periods of time, leading to nutrient limitation. When high‐intertidal pools are isolated during low tide, invertebrate‐excreted ammonium accumulates, providing a potential nitrogen source for macroalgae. I quantified the influence of mussels (Mytilus californianus Conrad) on ammonium accumulation rates in tide pools. I then evaluated the effects of ammonium loading by mussels on nitrogen assimilation and growth rates of Odonthalia floccosa (Esp.) Falkenb., a common red algal inhabitant of pools on northeastern Pacific rocky shores. Odonthalia was grown in artificial tide pool mesocosms in the presence and absence of mussels. Mesocosms were subjected to a simulated tidal cycle mimicking emersion and immersion patterns of high‐intertidal pools on the central Oregon coast. In the presence of mussels, ammonium accumulated more quickly in the mesocosms, resulting in increased rates of nitrogen assimilation into algal tissues. These increased nitrogen assimilation rates were primarily associated with higher growth rates. In mesocosms containing mussels, Odonthalia individuals added 41% more biomass than in mesocosms without mussels. This direct positive effect of mussels on macroalgal biomass represents an often overlooked interaction between macroalgae and invertebrates. In nutrient‐limited microhabitats, such as high‐intertidal pools, invertebrate‐excreted ammonium is likely an important local‐scale contributor to macroalgal productivity.     

Indirect trophic interactions with an invasive species affect phenotypic divergence in a top consumer

While phenotypic responses to direct species interactions are well studied, we know little about the consequences of indirect interactions for phenotypic divergence. In this study we used lakes with and without the zebra mussel to investigate effects of indirect trophic interactions on phenotypic divergence between littoral and pelagic perch. We found a greater phenotypic divergence between littoral and pelagic individuals in lakes with zebra mussels and propose a mussel-mediated increase in pelagic and benthic resource availability as a major factor underlying this divergence. Lakes with zebra mussels contained higher densities of large plankton taxa and large invertebrates. We suggest that this augmented resource availability improved perch foraging opportunities in both the littoral and pelagic zones. Perch in both habitats could hence express a more specialized foraging morphology, leading to an increased divergence of perch forms in lakes with zebra mussels. As perch do not prey on mussels directly, we conclude that the increased divergence results from indirect interactions with the mussels. Our results hence suggest that species at lower food web levels can indirectly affect phenotypic divergence in species at the top of the food chain.     

Determining algal assemblages in oligotrophic lakes and streams: comparing information from newly developed pigment/chlorophyll a ratios with direct microscopy

1. Pigment analyses by high performance liquid chromatography (HPLC) are commonly used for determining algal groups in marine and estuarine areas but are underdeveloped in freshwaters. In this study, 15 characteristic pelagic algal species (representing five algal groups) of oligo‐ / mesotrophic lakes were cultured and pigment / Chl a ratios determined at three light intensities. 2. With the exception of cyanophytes, light treatment had little effect on pigment / Chl a ratios. This justifies the use of the same pigment / Chl a ratios during seasonal studies where light conditions may change. 3. The determined pigment / Chl a ratios were tested on seasonal samples from five oligo‐ / mesotrophic lakes and three streams using CHEMTAX software. Pigment ratios of both pelagic and benthic algal communities from the lakes and streams were analysed to determine whether the pelagic algae‐based ratios can be used for benthic algal communities. 4. HPLC combined with CHEMTAX was useful for identifying freshwater phytoplankton classes and for quantifying the abundance of phytoplankton groups. However, although correlations were significant for six of seven phytoplankton classes studied, they were weak and varied with season. 5. HPLC was valid for quantifying benthic diatom groups in stream samples, whereas for lakes more benthic algal groups were recorded with HPLC than with microscopy and correlations between the two methods were not significant. 6. The use of both HPLC and microscopy is recommended as a cost‐efficient method for analysing many samples. It is crucial, however, that the CHEMTAX software is calibrated with the correct information, and the user is aware of the limitations.     

Light and nutrients regulate energy transfer through benthic and pelagic food chains

The amount of energy flowing to top trophic levels depends on primary production and the efficiency at which it is converted to production at each trophic level. In aquatic systems, algal production is often limited by light and nutrients, and the nutritional quality of algae depends on the relative balance of these two resources. In this study, we used a mesocosm experiment to examine how light and nutrient variation affected food chain efficiency (FCE, defined as the proportion of primary production converted to top trophic level production), using a food web with benthic and pelagic food chains. We also related variation in benthic and pelagic efficiencies to the nutritional quality of primary producers, i.e. carbon:nitrogen:phosphorus stoichiometry. As predicted, pelagic and benthic FCEs were highest under low light/high nutrient conditions, the treatment with the best algal food quality, i.e. the lowest C:nutrient ratios. Pelagic FCE and pelagic herbivore efficiency (HE_P) were more responsive than benthic FCE to variation in light and nutrients. Furthermore, pelagic FCE and HE_P were highly correlated with algal C:P, suggesting ‘carryover effects’ of algal food quality on carnivores (larval fish) via effects on herbivore (zooplankton) quality. Benthic (tadpole) production was primarily explained by primary production rate, suggesting food quantity rather than quality drives their production. However, benthic FCE was also highest at low light/high nutrients and was significantly correlated with food quality. The stronger effect of food quality in mediating pelagic compared to benthic efficiencies, is consistent with differences in the stoichiometric mismatches between algae and consumers. Pelagic FCE and HE_P were more likely to be P‐limited, whereas benthic FCE was more likely N‐limited. This study is the first to examine both pelagic and benthic FCE within the same system, and highlights the importance of differential consumer needs in determining how food quality affects energy transfer efficiency.     

Do mussel patches provide a refuge for algae from grazing gastropods?

On rocky shores, cover of macroalgae is often greater growingepibiotically on mussels compared to algae growing directlyattached to rock. A survey of two shores on the east coast ofIreland confirmed that mussel beds contained greater percentagealgal cover and more diverse algal assemblages compared to thoseon rock. The reasons for this difference are not clear. It hasbeen suggested that mussel beds provide a refuge for algae fromgrazing gastropods. Surprisingly, we found no evidence to supportthis. Using wax discs, gastropod grazing patterns were foundto be similar within the mussel beds as on rock. The musselbeds do not appear to provide a refuge for algae from grazingactivity at this scale and we suggest other possible mechanismsfor the prevalence of epibiotic algal cover on mussels. Intertidalgrazers may in fact affect the epibiotic algae on mussels andthereby affect indirectly the persistence of mussel beds. (Received 14 May 2007; accepted 20 October 2007)     

Zebra mussel δ 13C and δ 15N as a proxy for depth-specific pelagic isotope profiles and lake temperature

Zebra mussels (Dreissena polymorpha) can be used to provide a baseline stable isotope signature, time-integrated with primary production. However, since zebra mussels are uncommon in pelagic zones, their potential as reference species in pelagic water columns has not been fully explored. By investigating mussels growing suspended on a single vertical cable in Lake Constance, we were able to document seasonal (April, May, and August) and depth-dependent (0–22 m) variation in mussel δ ¹⁵Ν and δ ¹³C. We found a strong correlation between temperature and mussel δ ¹⁵N from differing depths_; and a strong relationship between temperature and Δ¹⁵Ν (estimated as δ ¹⁵Ν_mussel ? δ ¹⁵Ν_POM) and Δ¹³C (δ ¹³C_mussel ? δ ¹³C_POM). In a pattern that remained consistent over all months, Δ¹⁵Ν decreased with temperature, to the extent that negative values were recorded at temperatures >13°C. Utilizing cable-dwelling pelagic mussels as indicators of variation in isotope and temperature for pelagic water could be used as a novel field approach, comparable to in situ experimentation. We suggest that the pelagic mussel approach can be employed in tandem with benthic or littoral mussel isotope values, to develop mussel-based lake isoscapes. Such isoscapes may be pertinent to the study of seasonal trophic limnoecology and in tracking the movements of animals.     

Effects of the benthic suspension feeder Dreissena polymorpha on zooplankton in a large river

1. We conducted a series of in situ enclosure experiments to assess the impact of zebra mussels ( Dreissena polymorpha ) on the plankton of the Ohio River. Adult mussels were suspended in pelagic enclosures ('potamocorrals') at three densities (0, 1000, 2500 mussels per corral) and incubated for 6 days with daily plankton and physiochemical sampling.
2. The presence of adult zebra mussels was correlated with a shift in composition of the phytoplankton community and a severe reduction in some rotifers. The effects of zebra mussels on the larger zooplankton were taxon-dependent, but bacterial density showed no trend among treatments.
3. Zebra mussels may have significant negative impacts on zooplankton, which may in turn alter riverine food webs.     

Growth in length of Mytilus edulis L. fed on different algal diets

Mytilus edulis L. was fed on different algal diets and the increase in shell length was measured every 12–24 h. The mussels respond within 12 h to major changes in the diet. When pre-starved mussels were fed every 24 h with monocultures of Tetraselmis suecica (Kylin) Butch, there was a pronounced lag period followed by a linear increase in shell growth rate. When both pre-starved and pre-fed mussels were fed on equal rations of T. suecica with concentrations ranging from 2.5 to 10.0 × 10⁷ cells · 1^?1, the growth rate levelled off at about the same rate. Within the same range of concentrations there was a linear correlation between final growth rates and algal cell concentration. Feeding with monocultures of Isochrysis galbana (Parke) or Thalassiosira pseudonana Hasle et Heimdal (Hustedt) gave approximately the same shell growth as with Tetraselmis suecica alone, while combinations of these three algal species produced significant synergistic effects. When filtrate only from T. suecica cultures was supplied to the mussels there was a rapid initial stimulation of the shell growth. Centrifugated cells of T. suecica which were resuspended in filtered sea water, homogenized and sonicated to rupture the cells, gave 13% less growth than with untreated cells (P < 0.05). The use of the accurate laser diffraction method for length growth measurements may greatly reduce the time and effort involved in growth experiments.     

Hard rock versus soft bottom: the fauna associated with intertidal mussel beds on hard bottoms along the coast of Chile,and considerations on the functional role of mussel beds

The fauna associated with hard bottom mussel beds along the exposed Pacific coast of Chile was examined. The abundance of adult (>10 mm body length) purple mussels Perumytilus purpuratus varied between 32 and 75 individuals per 50 cm², and their biomass between 4.8 and 8.6 g AFDW per 50 cm² at eight sampling sites between Arica (18°S) and Chiloé (42°S). At all sampling sites, the associated fauna was dominated by suspension-feeding organisms (cirripeds, spionid and sabellid polychaetes, a small bivalve) followed by grazing peracarids and gastropods. Predators and scavengers also reached high abundances while deposit- and detritus-feeding organisms were of minor importance. The majority of organisms associated with these hard bottom mussel beds feed on resources obtained from the water column or growing on the mussels rather than on materials deposited by the mussels. This is in contrast to the fauna associated with mussel beds on soft bottoms, which comprises many species feeding on material accumulated by mussels (faeces and pseudofaeces) and deposited within the mussel bed. Many of the organisms dwelling between mussels both on hard bottoms and on soft bottoms have direct development, but organisms with pelagic development also occur abundantly within mussel beds. We propose that species with direct development are disproportionately favoured by the structurally complex habitat with diverse interstitial spaces between the mussels, which provides ample shelter for small organisms. We conclude that mussels on hard-bottoms primarily provide substratum for associated fauna while mussels on soft bottoms provide both substratum and food resources. Electronic Publication     

Nutrient loading associated with agriculture land use dampens the importance of consumer‐mediated niche construction

The linkages between biological communities and ecosystem function remain poorly understood along gradients of human‐induced stressors. We examined how resource provisioning (nutrient recycling), mediated by native freshwater mussels, influences the structure and function of benthic communities by combining observational data and a field experiment. We compared the following: (1) elemental and community composition (algal pigments and macroinvertebates) on live mussel shells and on nearby rocks across a gradient of catchment agriculture and (2) experimental colonisation of benthic communities on live vs. sham shells controlling for initial community composition and colonisation duration. We show that in near pristine systems, nutrient heterogeneity mediated by mussels relates to greater biodiversity of communities, which supports the notion that resource heterogeneity can foster biological diversity. However, with increased nutrients from the catchment, the relevance of mussel‐provisioned nutrients was nearly eliminated. While species can persist in disturbed systems, their functional relevance may be diminished or lost.     

Assessment of Top-Down and Bottom-Up Controls on the Collapse of Alewives (<Emphasis Type="Italic">Alosa pseudoharengus</Emphasis>) in Lake Huron

Food web models are powerful tools to inform management of lake ecosystems, where top-down (predation) and bottom-up (resource) controls likely propagate through multiple trophic levels because of strong predator–prey links. We used the Ecopath with Ecosim modeling approach to assess these controls on the Lake Huron main basin food web and the 2003 collapse of an invasive pelagic prey fish, alewife (Alosa pseudoharengus). We parameterized two Ecopath models to characterize food web changes occurring between two study periods of 1981–1985 and 1998–2002. We also built an Ecosim model and simulated food web time-dynamics under scenarios representing different levels of top-down control by Chinook salmon (Oncorhynchus tshawytscha) and of bottom-up control by quagga mussels (Dreissena rostriformis bugensis) and nutrients. Ecopath results showed an increase in the relative importance of bottom-up controls between the two periods, as production decreased across all trophic levels. The production of non-dreissenid benthos decreased most, which could cause decreases in production of pelagic prey fishes feeding on them. Ecosim simulation results indicated that the alewife collapse was caused by a combination of top-down and bottom-up controls. Results showed that while controls by Chinook salmon were relatively constant before alewife collapse, controls by quagga mussels and nutrients increased jointly to unsustainable levels. Under current conditions of low nutrients and high quagga mussel biomass, simulation results showed that recovery of alewives is unlikely regardless of Chinook salmon biomass in Lake Huron, which implies that the shrinking prey base cannot support the same level of salmonine predators as that prevailed during the 1980s.