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.     

Feeding in Arctic darkness: mid-winter diet of the pelagic amphipods Themisto abyssorum and T. libellula

The pelagic amphipods Themisto abyssorum and Themisto libellula represent important links between the herbivore zooplankton community and higher trophic levels of the Arctic marine food webs. Large double structured eyes of both of these hyperiid species are assumed to be used for visual prey detection. However, no information is available on the feeding strategies of these visually searching predators for the period of the polar night, a time of year with no or very low levels of daylight. Here, we report on the stomach and gut content of both Themisto species collected during a January expedition around Svalbard (78° to 81°N). Results indicate that T. abyssorum and T. libellula feed actively during the Arctic winter. The major food source of both amphipods consisted of calanoid copepods, most frequently Calanus finmarchicus.     

Terrestrial, benthic, and pelagic resource use in lakes: results from a three-isotope Bayesian mixing model

Fluxes of organic matter across habitat boundaries are common in food webs. These fluxes may strongly influence community dynamics, depending on the extent to which they are used by consumers. Yet understanding of basal resource use by consumers is limited, because describing trophic pathways in complex food webs is difficult. We quantified resource use for zooplankton, zoobenthos, and fishes in four low-productivity lakes, using a Bayesian mixing model and measurements of hydrogen, carbon, and nitrogen stable isotope ratios. Multiple sources of uncertainty were explicitly incorporated into the model. As a result, posterior estimates of resource use were often broad distributions; nevertheless, clear patterns were evident. Zooplankton relied on terrestrial and pelagic primary production, while zoobenthos and fishes relied on terrestrial and benthic primary production. Across all consumer groups terrestrial reliance tended to be higher, and benthic reliance lower, in lakes where light penetration was low due to inputs of terrestrial dissolved organic carbon. These results support and refine an emerging consensus that terrestrial and benthic support of lake food webs can be substantial, and they imply that changes in the relative availability of basal resources drive the strength of cross-habitat trophic connections.     

The effects of aggregation on the performance of the inverse method and indicators of network analysis

Food webs are usually aggregated into a manageable size for their interpretation and analysis. The aggregation of food web components in trophic or other guilds is often at the choice of the modeler as there is little guidance in the literature as to what biases might be introduced by aggregation decisions. We examined the impacts of the choice of the a priori model on the subsequent estimation of missing flows using the inverse method and on the indices derived from ecological network analysis of both inverse method-derived flows and on the actual values of flows, using the fully determined Sylt-Rømø Bight food web model. We used the inverse method, with the least squares minimization goal function, to estimate ‘missing’ values in the food web flows on 14 aggregation schemes varying in number of compartments and in methods of aggregation. The resultant flows were compared to known values; the performance of the inverse method improved with increasing number of compartments and with aggregation based on both habitat and feeding habits rather than diet similarity. Comparison of network analysis indices of inverse method-derived flows with that of actual flows and the original value for the unaggregated food web showed that the use of both the inverse method and the aggregation scheme affected indices derived from ecological network analysis. The inverse method tended to underestimate the size and complexity of food webs, while an aggregation scheme explained as much variability in some network indices as the difference between inverse-derived and actual flows. However, topological network indices tended to be most robust to both the method of determining flows and to the inverse method. These results suggest that a goal function other than minimization of flows should be used when applying the inverse method to food web models. Comparison of food web models should be done with extreme care when different methodologies are used to estimate unknown flows and to aggregate system components. However, we propose that indices such as relative ascendency and relative redundancy are most valuable for comparing ecosystem models constructed using different methodologies for determining missing flows or for aggregating system components.     

Decadal changes in a NW Mediterranean Sea food web in relation to fishing exploitation

We analysed changes in the ecological roles of species, trophic structure and ecosystem functioning using four standardized mass-balance models of the South Catalan Sea (North-western Mediterranean). Models represented the ecosystem during the late 1970s, mid 1990s, early 2000s, and a simulated no-fishing scenario. The underlying hypothesis was that ecosystem models should quantitatively capture the increasing exploitation in the ecosystem from the 1970s to 2000s, as well as differences between the exploited and non-exploited scenarios. Biomass showed a general decrease, while there was an increase in biomass at lower trophic levels (TL) from the 1970s to 2000s. The efficiency of energy transfer (TE) from lower to higher TLs significantly increased with time. The ecosystem during the 1990s showed higher biomass and flows than during the 1970s and 2000s due to an increase in small pelagic fish biomass (especially sardines). Exploited food webs also showed similarities in terms of general structure and functioning due to high intensity of fishing already in the 1970s. This intensity was highlighted with low trophic levels in the catch, high consumption of production by fisheries, medium to high primary production required to sustain the catches and high losses in secondary production due to fishing. Significant differences on ecosystem structure and functioning were highlighted between the exploited and no-fishing scenarios. Biomass of higher TLs increased under the no-fishing scenario and the mean trophic level of the community and the fish/invertebrate biomass ratios were substantially lower in exploited food webs. The efficiency of energy transfer (TE) from lower to higher TLs was lower under the no-fishing scenario, and it showed a continuous decrease with increasing TL. Marine mammals, large hake, anglerfish and large pelagic fish were identified as keystone species of the ecosystem when there was no fishing, while their ecological importance notably decreased under the exploited periods. On the contrary, the importance of small-sized organisms such as benthic invertebrates and small pelagic fish was higher in exploited food webs.     

Are closure terms appropriate or necessary descriptors of zooplankton loss in nutrient–phytoplankton–zooplankton type models?

Our current knowledge of plankton ecology ascribes a large proportion of zooplankton losses to zooplankton cannibalism and carnivory, rather than via the activity of higher trophic levels beyond the plankton. However, planktonic ecosystem models, such as the widely used nutrient–phytoplankton–zooplankton (NPZ) type models, typically represent all zooplankton losses by mathematically (rather than biologically) justified closure functions. Even where it is assumed that these closure functions include zooplanktonic cannibalism and carnivory, these processes are not explicitly implemented within the grazing function of the zooplankton. Here it is argued that this representation of zooplankton losses through “closure” terms within planktonic food web models is neither appropriate nor necessary. The general consequences of implementing a simple function incorporating zooplankton cannibalism and carnivory (intra-guild predation) within a planktonic food web model are compared against models implementing different types of traditional closure functions. While the modelled biomass outputs may appear similar, the fate of annual primary production and f-ratios vary widely. There appears no justification for the continued use of traditional closure term to depict zooplankton loss processes on biological or modelling arguments. To do so can seriously misrepresent the fate of primary production and thence trophic dynamics.     

Intraspecific variation in a predator affects community structure and cascading trophic interactions

Intraspecific phenotypic variation in ecologically important traits is widespread and important for evolutionary processes, but its effects on community and ecosystem processes are poorly understood. We use life history differences among populations of alewives, Alosa pseudoharengus, to test the effects of intraspecific phenotypic variation in a predator on pelagic zooplankton community structure and the strength of cascading trophic interactions. We focus on the effects of differences in (1) the duration of residence in fresh water (either seasonal or year-round) and (2) differences in foraging morphology, both of which may strongly influence interactions between alewives and their prey. We measured zooplankton community structure, algal biomass, and spring total phosphorus in lakes that contained landlocked, anadromous, or no alewives. Both the duration of residence and the intraspecific variation in foraging morphology strongly influenced zooplankton community structure. Lakes with landlocked alewives had small-bodied zooplankton year-round, and lakes with no alewives had large-bodied zooplankton year-round. In contrast, zooplankton communities in lakes with anadromous alewives cycled between large-bodied zooplankton in the winter and spring and small-bodied zooplankton in the summer. In summer, differences in feeding morphology of alewives caused zooplankton biomass to be lower and body size to be smaller in lakes with anadromous alewives than in lakes with landlocked alewives. Furthermore, intraspecific variation altered the strength of the trophic cascade caused by alewives. Our results demonstrate that intraspecific phenotypic variation of predators can regulate community structure and ecosystem processes by modifying the form and strength of complex trophic interactions.     

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

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

Comparative network analysis toward characterization of systemic organization for human–environmental sustainability

A preliminary study in comparative ecological network analysis was conducted to identify key assumptions and methodological challenges, test initial hypotheses and explore systemic and network structural characteristics for environmentally sustainable ecosystems. A nitrogen network for the U.S. beef supply chain – a small sub-network of the industrial food system analyzed as a pilot study – was constructed and compared to four non-human carbon and nitrogen trophic networks for the Chesapeake Bay and the Florida Everglades. These non-human food webs served as sustainable reference systems. Contrary to the main original hypothesis, the “window of vitality” and the number of network roles did not clearly differentiate between a human sub-network and the more complete non-human networks. The effective trophic level of humans (a partial estimate of trophic level based on the single food source of beef) was much higher (8.1) than any non-human species (maximum of 4.88). Network connectance, entropy, total dependency coefficients, trophic efficiencies and the ascendency to capacity ratio also indicated differences that serve as hypotheses for future tests on more comprehensive human food webs. The study elucidated important issues related to (1) the steady state assumption, which is more problematic for industrial human systems, (2) the absence or dearth of data on contributions of dead humans and human wastes to feed other species in an integrated food web, (3) the ambiguity of defining some industrial compartments as living versus non-living, and (4) challenges with constructing compartments and trophic transfers in industrial versus non-human food webs. The two main novel results are (1) the progress made toward adapting ecological network analysis (ENA) methodology for analysis of human food networks in industrial cultures and (2) characterizing the critical aspects of comparative ENA for understanding potential causes of the problems, and providing avenues for solutions, for environmental sustainability. Based on this work, construction and comparative network analysis of a more comprehensive industrial human food network seems warranted and likely to provide valuable insights for modifying structures of industrial food networks to be more like natural networks and more sustainable.     

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.     

Linking trophic positions and flow structure constraints in ecological networks: Energy transfer efficiency or topology effect?

In the present work we investigate whether the distribution of energy flows in ecosystems responds to criteria of trophic organization. We analyzed weighted and unweighted food webs estimating, for each node, trophic position (TP), Shannon's index of inflow diversity (H) and individual contribution to the whole average mutual information (AMI). Finally, we performed the same analysis on simulated webs that were constructed using the following criteria: (a) preserving topology and varying link strength; (b) modifying position of links and their intensities.     

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

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

Restoring piscivorous fish populations in the Laurentian Great Lakes causes seabird dietary change

Ecosystem change often affects the structure of aquatic communities thereby regulating how much and by what pathways energy and critical nutrients flow through food webs. The availability of energy and essential nutrients to top predators such as seabirds that rely on resources near the water's surface will be affected by changes in pelagic prey abundance. Here, we present results from analysis of a 25-year data set documenting dietary change in a predatory seabird from the Laurentian Great Lakes. We reveal significant declines in trophic position and alterations in energy and nutrient flow over time. Temporal changes in seabird diet tracked decreases in pelagic prey fish abundance. As pelagic prey abundance declined, birds consumed less aquatic prey and more terrestrial food. This pattern was consistent across all five large lake ecosystems. Declines in prey fish abundance may have primarily been the result of predation by stocked piscivorous fishes, but other lake-specific factors were likely also important. Natural resource management activities can have unintended consequences for nontarget ecosystem components. Reductions in pelagic prey abundance have reduced the capacity of the Great Lakes to support the energetic requirements of surface-feeding seabirds. In an environment characterized by increasingly limited pelagic fish resources, they are being offered a Hobsonian choice: switch to less nutritious terrestrial prey or go hungry.     

Fish production and the marine ecosystems of the Scotian Shelf,eastern Canada

Fishery production and food webs have been studied on the Scotian Shelf and upper continental slope along a transect running 270 km SE of Halifax, Canada. The area (TCNAF Division 4W) supports a fishery of roughly 0.15x10⁶ metric tons. Overall primary production of the shelf waters is 102 g C m^-2 year^-1 and of the slope wacers about 128 g C m^-2 year^-1. Demersal fish production (average 4.1 kcal m^-2 year^-1) is highest over the shelf and declines in an offshore direction, while pelagic fish production (average 16 kcal m^-2 year^-1) is highest over the slope and declines in an inshore direction. Hypothetical food webs of these two intergrading ecosystems have been constructed, based on data for primary production, fish catches, and the biomasses of zooplankton and macrobenthos. These lead us to suggest that there are basic differences in food chains and efficiencies between the two ecosystems that account for their differences in production. Although primary production is 17% higher on an average on the Nova Scotian transect than in the North Sea, the apparent zooplankton and macrobenthos production is 31% lower and macrobenthos production may also be lower. Overall fish catch from the Scotian Shelf and slope is about 47% lower per unit area than the catch in the North Sea, despite the fact that the demersal catches are identical. This is accounted for by a much lower overall pelagic catch from the Nova Scotian area, centered in a region that is small compared to the total area. Fish production in different regions cannot be predicted merely on the basis of differences in level of primary production, but must take into account differences in the structure of the ecosystems.     

Origin of compartmentalization in food webs

The response of an ecosystem to perturbations is mediated by both antagonistic and facilitative interactions between species. It is thought that a community's resilience depends crucially on the food web--the network of trophic interactions--and on the food web's degree of compartmentalization. Despite its ecological importance, compartmentalization and the mechanisms that give rise to it remain poorly understood. Here we investigate several definitions of compartments, propose ways to understand the ecological meaning of these definitions, and quantify the degree of compartmentalization of empirical food webs. We find that the compartmentalization observed in empirical food webs can be accounted for solely by the niche organization of species and their diets. By uncovering connections between compartmentalization and species' diet contiguity, our findings help us understand which perturbations can result in fragmentation of the food web and which can lead to catastrophic effects. Additionally, we show that the composition of compartments can be used to address the long-standing question of what determines the ecological niche of a species.     

Can stable isotope ratios provide for community-wide measures of trophic structure?

Stable isotope ratios (typically of carbon and nitrogen) provide one representation of an organism's trophic niche and are widely used to examine aspects of food web structure. Yet stable isotopes have not been applied to quantitatively characterize community-wide aspects of trophic structure (i.e., at the level of an entire food web). We propose quantitative metrics that can be used to this end, drawing on similar approaches from ecomorphology research. For example, the convex hull area occupied by species in delta13C-delta15N niche space is a representation of the total extent of trophic diversity within a food web, whereas mean nearest neighbor distance among all species pairs is a measure of species packing within trophic niche space. To facilitate discussion of opportunities and limitations of the metrics, we provide empirical and conceptual examples drawn from Bahamian tidal creek food webs. These examples illustrate how this methodology can be used to quantify trophic diversity and trophic redundancy in food webs, as well as to link individual species to characteristics of the food web in which they are embedded. Building from extensive applications of stable isotope ratios by ecologists, the community-wide metrics may provide a new perspective on food web structure, function, and dynamics.     

Recherches sur la situation trophique d'un groupe d'organismes pélagiques (Euphausiacea). VI. Conclusions sur le rôle des euphausiacés dans les circuits trophiques de l'océan Pacifique intertropical

The role of euphausiids in the food webs of the Intertropical Pacific Ocean is defined through analysis of their nutrition, vertical distributions and migrations, and their utilization by pelagic predators. It is suggested that the abundance of the group, the extensive vertical migrations of many species and the fact that feeding takes place mainly in subsurface layers, result in a leading role of euphausiids in energy transfer between different bathymetric levels. For night-time feeding predators, they represent a noticeable food source only in the 0 to 300 m water layer, as 97% of the euphausiid biomass concentrates in this layer at night. In the daytime, only the smaller specimens (chiefly genus Stylocheiron), accounting for 10 to 15% of the whole biomass of the group, remain available for epipelagic (0 to 400 m) predators, larger individuals dwelling deeper. Euphausiids account for 8 to 10% of the food ingested by micronektonic fishes, but the species are not the same for different categories of fishes. Migrating fishes caught by pelagic trawls, more or less connected with the deep scattering layer, feed on migrating species in subsurface layers at night as well as in deeper layers during the daytime, and on non-migrating species inhabiting shallower and intermediate layers. On the other hand, fishes which comprise the prey of large long-line tunas, which are not caught by trawls because they are fast swimmers, feed almost solely on species which remain above 400 m in the daytime. These results suggest a certain degree of independance between the trophic webs which concern, on the one hand, epipelagic ichthyofauna (including tuna), and, on the other hand, migrating and deep-living faunas. Migrating populations are able to feed at night upon subsurface organisms, a part of this resource being then transmitted during the day to the deep-living fauna; but the epipelagic ichthyofauna, with a feeding activity restricted to light hours, has few possibilities to benefit from the migrating or deepliving biomass. Therefore, energy transfers seem to be intense only from subsurface (0 to 400 m) to deeper layers. From a more general point of view, these investigations suggest that, in the pelagic system, vertical distributions and migrations, and feeding rhythms, are the main factors determining the structure of the food webs.     

Food utilisation of pelagic mysids, Mysis mixta and M. relicta, during their growing season in the northern Baltic Sea

The dietary habits of the pelagic mysid Mysismixta were studied during its growing season at an open sea location in the Gulf of Finland, the northern Baltic Sea. Stomach samples were taken twice a month from June to September 1997. The most abundant phytoplankton taxa in the stomachs were diatoms and dinoflagellates, and copepods and cladocerans were the most abundant zooplankton identified. A clear change was found in the diets during the study period. Small mysids (3 to 6 mm) fed on sedimented phytoplankton in the early summer (90% benthic particles in June) but shifted gradually to a more pelagic and carnivorous diet (>40% pelagic particles, consisting of ca. 60% zooplankton in September). Seasonal changes in mysid capture ability as well as food availability were suggested to affect the diet composition of mysids during their growth. The ratio of pelagic and benthic food particles could – irrespective of the season – be explained by mysid size, whereas the zooplankton:phytoplankton ratio was better explained by season. The stomach analysis suggests that the mysids needed to attain a threshold size of 8 to 11 mm to initiate feeding on the more evasive copepods. Mysids also started to grow faster at the same time as the proportion of copepods increased in the diet, which suggests that copepods are an important energy source for M. mixta in late summer. Finally, a comparison was made between the M. mixta diet and that of the less abundant M. relicta. The diets of the two pelagic mysid species overlapped by 75% (Schoener's index). The main difference was due to M. mixta eating more zooplankton and pelagic material than M. relicta. Received: 15 September 1999 / Accepted: 18 January 2000     

Independent gradients of producer, consumer, and microbial diversity in lake plankton

Interactions between trophic levels during food web assembly can drive positive correlations in diversity between producers, consumers, and decomposers. However, the contribution of trophic interactions relative to local environmental factors in promoting species diversity is poorly understood, with many studies only considering two trophic levels. Here we examine correlations in diversity among zooplankton, phytoplankton, and bacteria in the pelagic zone of 31 lakes in British Columbia, Canada. We sampled species diversity of zooplankton and phytoplankton through morphological identification, and bacterial genetic diversity was estimated by denaturing gradient gel electrophoresis (DGGE) of 16S rDNA polymorphisms. We looked for correlations in diversity that were independent of the abiotic environment by statistically controlling for 18 limnological variables. No significant correlations were found between the diversity of zooplankton, phytoplankton, and bacteria. In addition, the physical factors that were associated with species composition in one trophic level were independent of those that were important for another. Our results provide no support for the importance of direct feedbacks between producers, consumers, and decomposers in maintaining diversity. Zooplankton, phytoplankton, and bacterial diversity and composition are regulated independently from one another and respond to different environmental variables. These results suggest that species of lake plankton show loose trophic associations with one another due to broad diets in consumers and decomposers.