Top Trophic Levels Research Articles

Foundation species support fauna across multiple trophic levels via trophic and non‐trophic mechanisms

Abstract Foundation species, such as seagrasses, trees, and corals, form the biotic basis for many ecosystems. They engineer local habitats and can support many faunal species through trophic (e.g. providing feeding grounds) or non‐trophic (e.g. providing predation shelters and spawning grounds) mechanisms. How the supporting effects of foundation species on fauna change across multiple trophic levels remains poorly understood. We investigated how a foundational seagrass supported faunal species at different trophic levels, using a series of comparative surveys, diet analyses, and field experiments. We found that seagrasses substantially enhanced marine fauna across all but one of four trophic levels. Primary, secondary, and top consumers were significantly more abundant, whilst tertiary consumers were similarly or less abundant in seagrass beds than on unvegetated mudflats. Importantly, seagrasses supported fauna of different trophic levels via different trophic or non‐trophic mechanisms; seagrasses provided mainly feeding, refuging and feeding, feeding, and spawning grounds for primary, secondary, tertiary, and top consumers, respectively. Our work reveals that seagrasses support fauna from primary to top trophic levels via different trophic and non‐trophic mechanisms. Our findings highlight that protecting and restoring seagrasses is critical to enhancing fauna across multiple trophic levels and can inform conservation strategies targeted for fauna at different trophic levels. Our study can help stimulate future studies from other ecosystems to test how foundation species affect fauna across different trophic levels. Read the free Plain Language Summary for this article on the Journal blog.

The role of evolutionary processes in determining trophic structure

There are two contrasting hypotheses regarding the drivers of biomass‐distribution among trophic levels within ecosystems. Energetic or bottom–up models propose control by supply of energy, either from autotrophy or from the underlying trophic level. Dynamical or top–down models propose control by predators in the overlying trophic level. Although multiple approaches have been used to reconcile these hypotheses, they have rarely considered the evolutionary pressures created by different distributions of biomass. Here, we study the effects of these evolutionary processes using an agent‐based, spatially‐explicit, eco‐evolutionary model. We demonstrate that, when ecosystems are simple and predator–prey relationships between species are fixed and do not evolve, primary productivity limits the number of trophic levels. In this case, the abundance in the top trophic level is always limited by productivity. However, productivity‐limited trophic levels subject those below them to limitation by predation, and predation‐limited trophic levels allow trophic levels below them to grow until limited by productivity. These results match the expectations of the exploitation ecosystems hypothesis (EEH), which predicts the same pattern of alternating top–down and bottom–up control on trophic levels. When species are able to evolve to adaptively adjust their trophic levels, however, selection is liable to lead to collapse of the trophic pyramid through prey switching. Under these conditions, all trophic levels experience bottom–up control. When the evolution of prey switching is restricted, higher trophic levels are evolutionarily stable and the predictions of the EEH are once again met; the stability of long food chains is thus dependent on the difficulty associated with prey switching. These results suggest that, at least under the combinations of model parameters explored in this study, evolutionary limitation is key to maintaining trophic structure.

Droughts reshape apex predator space use and intraguild overlap.

Droughts are increasing in frequency and severity globally due to climate change, leading to changes in resource availability that may have cascading effects on animal ecology. Resource availability is a key driver of animal space use, which in turn influences interspecific interactions like intraguild competition. Understanding how climate-induced changes in resource availability influence animal space use, and how species-specific responses scale up to affect intraguild dynamics, is necessary for predicting broader community-level responses to climatic changes. Although several studies have demonstrated the ecological impacts of drought, the behavioural responses of individuals that scale up to these broader-scale effects are not well known, particularly among animals in top trophic levels like large carnivores. Furthermore, we currently lack understanding of how the impacts of climate variability on individual carnivore behaviour are linked to intraguild dynamics, in part because multi-species datasets collected at timescales relevant to climatic changes are rare. Using 11 years of GPS data from four sympatric large carnivore species in southern Africa-lions (Panthera leo), leopards (Panthera pardus), African wild dogs (Lycaon pictus) and cheetahs (Acinonyx jubatus)-spanning 4 severe drought events, we test whether drought conditions impact (1) large carnivore space use, (2) broad-scale intraguild spatial overlap and (3) fine-scale intraguild interactions. Drought conditions expanded space use across species, with carnivores increasing their monthly home range sizes by 35% (wild dogs) to 66% (leopards). Drought conditions increased the amount of spatial overlap between lions and subordinate felids (cheetahs and leopards) by up to 119%, but only lion-cheetah encounter rates were affected by these changes, declining in response to drought. Our findings reveal that drought has a clear signature on the space use of multiple sympatric large carnivore species, which can alter spatiotemporal partitioning between competing species. Our study thereby illuminates the links between environmental change, animal behaviour and intraguild dynamics. While fine-scale avoidance strategies may facilitate intraguild coexistence during periodic droughts, large carnivore conservation may require considerable expansion of protected areas or revised human-carnivore coexistence strategies to accommodate the likely long-term increased space demands of large carnivores under projected increases in drought intensity.

Insight into the trophic niche and prey contribution in the diet of Lutjanus griseus (Linnaeus, 1758) captured by artisanal fishing fleet in the central and southwest Gulf of Mexico

Lutjanus griseus is a common species for fisheries of the Gulf of Mexico, and it is an effective vehicle to transport energy from low to top trophic levels in the ecosystem. The management of fish populations is based on the modeling and prediction of abundances. To accomplish this, trophic ecology results allow the understanding of habitat use and population expansions. To increase the knowledge in the trophic ecology of L. griseus, this study described its trophic niche, diet composition, and the prey contribution in the formation of muscle tissue. Results showed that the trophic niche in Veracruz was wider than in Campeche, this related to the consumption of prey in different habitats, coral reefs areas in Veracruz and seagrass beds in Campeche. According to the δ13C y δ15N values, in both regions the crustaceans were the most important group in the contribution of muscle tissue, however, it contrasted with the stomach content results, which suggested that the diet in Campeche was mainly composed of crustaceans, and in Veracruz of fishes. This difference could be related to the snapper’s movements, as they can be found in sea bottoms covered with seagrass, with high abundance of crustaceans, and sea bottoms composed of coral reefs with high diversity of fishes and other preys. Results highlight the generalist and opportunistic feeding strategy of L. griseus, this strategy allowed its population expansion to the north, consuming the available food source in the different habitats it occupied.

Wind-driven upwelling of iron sustains dense blooms and food webs in the eastern Weddell Gyre

The Southern Ocean is a major sink of anthropogenic CO2 and an important foraging area for top trophic level consumers. However, iron limitation sets an upper limit to primary productivity. Here we report on a considerably dense late summer phytoplankton bloom spanning 9000 km2 in the open ocean of the eastern Weddell Gyre. Over its 2.5 months duration, the bloom accumulated up to 20 g C m−2 of organic matter, which is unusually high for Southern Ocean open waters. We show that, over 1997–2019, this open ocean bloom was likely driven by anomalies in easterly winds that push sea ice southwards and favor the upwelling of Warm Deep Water enriched in hydrothermal iron and, possibly, other iron sources. This recurring open ocean bloom likely facilitates enhanced carbon export and sustains high standing stocks of Antarctic krill, supporting feeding hot spots for marine birds and baleen whales.

Temporal and spatial trends of chlorinated paraffins and organophosphate flame retardants in black-tailed gull (Larus crassirostris) eggs

In this study, eggs of black-tailed gull (Larus crassirostris), a top trophic level predator of marine ecosystem were, for the first time, monitored to assess the temporal and spatial trends of emerging pollutants in South Korea. Two Island regions, namely, Baekryeong-do (Site A) and Hong-do (Site B) were investigated from 2012 to 2018, and the total levels of short chain chlorinated paraffins (SCCPs), medium chain CPs (MCCPs), and organophosphate flame retardants (OPFRs) for both Site A and B were 1180–2931 and 694–2023 ng/g lipid weight (lw), 1287–4898 and 1034–3075 ng/g lw, and 203–499 and 233–409 ng/g lw, respectively. The time-trends of the concentration of pollutants showed an increasing tendency from 2012 to 2018, with the levels predicted to be doubled within three years, following the results of regression analysis. A shift in temporal-trends from shorter to longer chain CPs was noted, suggesting the effect of industrial-related contamination. Especially, significantly high levels of CPs and OPFRs were found in the site adjacent to China, which is reasonable as China is the largest producer and consumer of FRs and plasticizers worldwide. This study is valuable to understand the temporal increment of emerging pollutants as the alternatives of phased-out FRs and plasticizers, while raising the need for continuous environmental management.

Body size dependent dispersal influences stability in heterogeneous metacommunities

Body size affects key biological processes across the tree of life, with particular importance for food web dynamics and stability. Traits influencing movement capabilities depend strongly on body size, yet the effects of allometrically-structured dispersal on food web stability are less well understood than other demographic processes. Here we study the stability properties of spatially-arranged model food webs in which larger bodied species occupy higher trophic positions, while species’ body sizes also determine the rates at which they traverse spatial networks of heterogeneous habitat patches. Our analysis shows an apparent stabilizing effect of positive dispersal rate scaling with body size compared to negative scaling relationships or uniform dispersal. However, as the global coupling strength among patches increases, the benefits of positive body size-dispersal scaling disappear. A permutational analysis shows that breaking allometric dispersal hierarchies while preserving dispersal rate distributions rarely alters qualitative aspects of metacommunity stability. Taken together, these results suggest that the oft-predicted stabilizing effects of large mobile predators may, for some dimensions of ecological stability, be attributed to increased patch coupling per se, and not necessarily coupling by top trophic levels in particular.

Top predators govern multitrophic diversity effects in tritrophic food webs.

It is well known that functional diversity strongly affects ecosystem functioning. However, even in rather simple model communities consisting of only two or, at best, three trophic levels, the relationship between multitrophic functional diversity and ecosystem functioning appears difficult to generalize, because of its high contextuality. In this study, we considered several differently structured tritrophic food webs, in which the amount of functional diversity was varied independently on each trophic level. To achieve generalizable results, largely independent of parametrization, we examined the outcomes of 128,000 parameter combinations sampled from ecologically plausible intervals, with each tested for 200 randomly sampled initial conditions. Analysis of our data was done by training a random forest model. This method enables the identification of complex patterns in the data through partial dependence graphs, and the comparison of the relative influence of model parameters, including the degree of diversity, on food-web properties. We found that bottom-up and top-down effects cascade simultaneously throughout the food web, intimately linking the effects of functional diversity of any trophic level to the amount of diversity of other trophic levels, which may explain the difficulty in unifying results from previous studies. Strikingly, only with high diversity throughout the whole food web, different interactions synergize to ensure efficient exploitation of the available nutrients and efficient biomass transfer to higher trophic levels, ultimately leading to a high biomass and production on the top level. The temporal variation of biomass showed a more complex pattern with increasing multitrophic diversity: while the system initially became less variable, eventually the temporal variation rose again because of the increasingly complex dynamical patterns. Importantly, top predator diversity and food-web parameters affecting the top trophic level were of highest importance to determine the biomass and temporal variability of any trophic level. Overall, our study reveals that the mechanisms by which diversity influences ecosystem functioning are affected by every part of the food web, hampering the extrapolation of insights from simple monotrophic or bitrophic systems to complex natural food webs.

Fisheries restoration potential: Optimizing fisheries profits while maintaining food web structure

There is mounting evidence that fishing impacts can fundamentally alter the structure of whole ecosystems, whereby removal of top trophic levels is expected to generally cascade down the food web, allowing prey at lower trophic levels to flourish. These cascades can lead to decreased diversity but increased productivity in ecosystems, which may lead to increased yields in multi-species fisheries. In a world with a growing human population to feed, increasing consumption of fish protein, and also a growing human footprint on our aquatic ecosystems, this sets up an interesting economic dilemma: do we aim to restore fisheries ecosystems at the expense of foregoing yield? Alternatively, is there a possible way to benefit both restoration and people's livelihoods? Here, we discuss how various market and food web characteristics may allow for equally sustainable and profitable fisheries using a simple food chain model with multi-species fishing pressure. We show that fisheries with higher economic value for top predators relative to intermediate consumers, and with high potential for fisheries-induced trophic cascades, meet criteria for high fishery restoration potential. In these situations, managing fisheries for maximizing profits rather than total productivity can lead to increased maintenance of food web structure and function as well as being economically beneficial. We discuss emerging examples that are consistent with this result.

Abundance data for invertebrate assemblages from intertidal mussel beds along the Atlantic Canadian coast.

This data set describes the abundance of 50 invertebrate taxa found in intertidal mussel beds along the Atlantic Canadian coast. This information resulted from a regional-scale study that investigated the effects of wave exposure on the richness and composition of invertebrate assemblages from intertidal mussel beds. Abundance data are provided for taxa representing the Annelida, Arthropoda, Bryozoa, Chordata, Cnidaria, Echinodermata, Mollusca, Nematoda, Nemertea, and Platyhelminthes. The data characterize mussel beds from wave-sheltered and wave-exposed locations spanning 315 km of the coast of Nova Scotia. Univariate and multivariate analyses revealed that the compositional structure of these invertebrate assemblages differed markedly depending on wave exposure. Overall, because of its taxonomic diversity, the inclusion of data for basal, intermediate, and top trophic levels, and the coverage of two extremes of environmental stress, this data set could be useful to test broader aspects of ecological theory. Areas of ecology that could advance using this data set are those concerning environmental stress models of community organization, abundance-occupancy relationships, species co-occurrence, species abundance distributions, dominance and rarity, spatial scales of population and community variation, and distribution of functional and phylogenetic diversity. Use of this data set for academic or educational purposes is allowed as long as the data source is properly cited. When used for academic or educational purposes, this data set should be cited using the title of this Data Paper, the names of the authors, the year of publication, and the corresponding volume and article numbers.

Spatio-temporal partitioning facilitates mesocarnivore sympatry in the Stara Planina Mountains, Bulgaria

The top trophic level in many terrestrial food webs is typically occupied by mammalian carnivores (Order Carnivora) that broadly affect and shape ecosystems through trophic cascades. Their inter-specific interactions can further complicate effects on community dynamics as a consequence of intra-guild competition. The capacity for competitive mammalian carnivores to segregate their hunting and activity regimes is in major part a function of their similarity, in terms of body-size and dietary niche; termed the ‘niche variation hypothesis’. In this study, we used camera-trapping to investigate intra-guild interactions and spatio-temporal partitioning among five mesocarnivores, the golden jackal (Canis aureus), European badger (Meles meles), red fox (Vulpes vulpes), European wildcat (Felis sylvestris) and stone marten (Martes foina), in the Stara Planina Mountains, Bulgaria. We collected a total of 444 images of golden jackals, 236 images of European badgers, 200 images of red foxes, 171 images of stone martens, and 145 images of European wildcats, from 6612 camera-days across fifteen camera-trapping stations. With respect to body size, the three smaller species (fox, wildcat and marten) were active in different time periods than the two larger competitors (jackal and badger) through both the warm and cold season. The more similar the trophic niche between species pairs (particularly relating to rodent consumption), the greater the spatio-temporal partitioning we observed within the pair; however, this adapted to seasonal dietary shifts. In conclusion, spatial and temporal (fine-scale and seasonal) niche partitioning appeared to reduce encounter probabilities and competition and may act to facilitate sympatric coexistence among this regional mesocarnivore guild.

First study of food webs in a large glacial river: the trophic role of invasive trout

Abstract The aim of this study was to determine the food webs structure of a large Patagonian river in two river sections (Upstream and Midstream) and to evaluate isotopic overlap between native and introduced species. We used stable isotope analyses of δ15N and δ13C and stomach content. The Upstream section had a more complex food webs structure with a greater richness of macroinvertebrates and fish species than Midstream. Upstream basal resources were dominated by filamentous algae. Lake Trout were found to have a higher trophic position than all other fish species in that area although, the most abundant fish species, were Rainbow Trout. Depending on the life stage, Rainbow Trout shifted from prey to competitor/predator. In the Midstream section, the base of the food webs was dominated by coarse particulate organic matter, and adult Rainbow Trout had the highest trophic level. Isotopic values changed among macroinvertebrates and fish for both areas. The two most abundant native and invasive species — Puyen and Rainbow Trout — showed an isotopic separation in Midstream but did not in Upstream areas. The presence of invasive fish that occupy top trophic levels can have a significant impact on native fish populations that have great ecological importance in the region.

Nonlinear patterns in mercury bioaccumulation in American alligators are a function of predicted age

Mercury is a widespread, naturally occurring contaminant that biomagnifies in wetlands due to the methylation of this element by sulfate-reducing bacteria. Species that feed at the top trophic level within wetlands are predicted to have higher mercury loads compared to species feeding at lower trophic levels and are therefore often used for mercury biomonitoring. However, mechanisms for mercury bioaccumulation in sentinel species are often poorly understood, due to a lack of long-term studies or an inability to differentiate between confounding variables. We examined mercury bioaccumulation patterns in the whole blood of American alligators (Alligator mississippiensis) from a long-term mark-recapture study (1979–2017) in South Carolina, USA. Using a growth model and auxiliary information on predicted age at first capture, we differentiated between age- and size-related variation in mercury bioaccumulation, which are often confounded in alligators due to their determinate growth pattern. Contrary to predictions that the oldest or largest individuals were likely to have the highest mercury concentrations, our best-supported model indicated a peak in mercury concentration at 30–40 years of age, depending on the sex, and lower concentrations in the youngest and oldest animals. To evaluate the robustness of our findings, we re-analyzed data from a previously published study of mercury in alligators sampled at Merritt Island National Wildlife Refuge in Florida. Unlike the South Carolina data, the data from Florida contained minimal auxiliary information regarding age, yet the best supported model similarly indicated a peaked rather than increasing relationship between mercury and body size, a less-precise indicator of age. These findings highlight how long-term monitoring can differentiate between confounding variables (e.g., age and size) to better elucidate complex relationships between contaminant exposure and demographic factors in sentinel species.

The influence of biotic and abiotic factors on banded common loon (Gavia immer) reproductive success in a remote, mountainous region of the northeastern United States.

Habitat degradation resulting from anthropogenic activities can threaten wildlife populations. Even wildlife existing in seemingly pristine areas are at risk of airborne pollutants and urban development. The common loon (Gavia immer), a top-trophic level predator in freshwater aquatic ecosystems, has previously experienced detrimental changes in reproductive success as a result of anthropogenic activities. However, long-term studies and large sample sizes are necessary to ascertain the impacts of various anthropogenic activities on this long-lived species. Using a multi-year dataset, we investigated the effects of multiple biotic and abiotic factors on the probability of adult male and female common loon hatching and fledging success. From 1998-2017, we banded individual loons, collected blood samples to assess mercury (Hg) exposure of the birds, and monitored their reproductive success. Adult female loon hatching success was negatively associated with the amount of rainfall received in a given year while fledging success was positively associated with the amount of shoreline development. Adult male loon hatching success was positively associated with the amount of shoreline development and fledging success was negatively associated with the number of other loon pairs on a lake.

Critical review of mercury contamination in Sri Lankan fish and aquatic products.

Mercury (Hg) in fish and aquatic products is a potential threat to human health and international trade, and guidelines of international advisory bodies are established for assessing Hg in these foods. As fish are the most consumed animal protein in Sri Lanka, this assessment summarises for the first-time total mercury concentrations in Sri Lankan marine and freshwater fish, invertebrates, aquatic plants, and fisheries products. It reveals that Hg levels in fish and other edible aquatic species are mostly below the published safety limits, except for certain top trophic level fish (swordfish, tuna, marlin). The review also highlights gaps in Sri Lankan Hg assessments such as a total lack of data for methyl‑mercury in these aquatic species and food products. This data compilation and assessment will serve as an initial baseline for comparison with results from future monitoring and research studies in Sri Lanka while adding to the world-wide Hg database.

Effects of Selenium on Mercury Bioaccumulation in a Terrestrial food Chain from an Abandoned Mercury Mining Region.

Few reports of the relationship exist between mercury (Hg) and selenium (Se) from locations of severe Hg contamination in terrestrial environments. Here, we report the concentrations of Hg and Se as well as Se:Hg molar ratios in biotic samples collected from a region with a long history of Hg mining. Nitrogen isotopes (δ15N) were analyzed to confirm the trophic levels. Results showed that bird feathers at the top trophic level exhibited the highest Hg concentrations, while the lowest concentrations were found in herbivorous insects, demonstrating a significant biomagnification across the food chain. In contrast, herbivorous insects of different types (generalists vs. specialized rice pests) exhibited both the highest and the lowest concentrations of Se, indicating a lack of biomagnification. Indeed, Se was correlated positively with Hg when Se:Hg ratios were greater than one, suggesting Se:Hg molar ratios can be a controlling influence on Hg in terrestrial organisms.

Reef fish functional traits evolve fastest at trophic extremes.

Trophic ecology is thought to exert a profound influence on biodiversity, but the specifics of the process are rarely examined at large spatial and evolutionary scales. We investigate how trophic position and diet breadth influence functional trait evolution in one of the most species-rich and complex vertebrate assemblages, coral reef fishes, within a large-scale phylogenetic framework. We show that, in contrast with established theory, functional traits evolve fastest in trophic specialists with narrow diet breadths at both very low and high trophic positions. Top trophic level specialists exhibit the most functional diversity, while omnivorous taxa with intermediate trophic positions and wide diet breadth have the least functional diversity. Our results reveal the importance of trophic position in shaping evolutionary dynamics while simultaneously highlighting the incredible trophic and functional diversity present in coral reef fish assemblages.

Coexistence of species in a tritrophic food chain model with Holling functional response type IV

We determine conditions on the parameters of a tritrophic food chain model, implying the coexistence of the species. We consider that both predators corresponding to middle and top trophic levels have Holling functional responses type IV, and the prey at the lower trophic level has either linear or logistic growth rate. We prove that the differential system has an equilibrium point in which it exhibits a supercritical Hopf bifurcation independently of the growth rate of the prey. In the logistic case, we prove the existence of at least three equilibrium points in the positive octant and one of them exhibits a supercritical Hopf bifurcation.

Eco-evolutionary Feedbacks from Non-target Species Influence Harvest Yield and Sustainability

Evolution in harvested species has become a major concern for its potential to affect yield, sustainability, and recovery. However, the current singular focus on harvest-mediated evolution in target species overlooks the potential for evolution in non-target members of communities. Here we use an individual-based model to explore the scope and pattern of harvest-mediated evolution at non-target trophic levels and its potential feedbacks on abundance and yield of the harvested species. The model reveals an eco-evolutionary trophic cascade, in which harvest at top trophic levels drives evolution of greater defense or competitiveness at subsequently lower trophic levels, resulting in alternating feedbacks on the abundance and yield of the harvested species. The net abundance and yield effects of these feedbacks depends on the intensity of harvest and attributes of non-target species. Our results provide an impetus and framework to evaluate the role of non-target species evolution in determining fisheries yield and sustainability.

Benthic carbon is inefficiently transferred in the food webs of two eutrophic shallow lakes

Abstract The sum of benthic autotrophic and bacterial production often exceeds the sum of pelagic autotrophic and bacterial production, and hence may contribute substantially to whole‐lake carbon fluxes, especially in shallow lakes. Furthermore, both benthic and pelagic autotrophic and bacterial production are highly edible and of sufficient nutritional quality for animal consumers. We thus hypothesised that pelagic and benthic transfer efficiencies (ratios of production at adjacent trophic levels) in shallow lakes should be similar. We performed whole ecosystem studies in two shallow lakes (3.5 ha, mean depth 2 m), one with and one without submerged macrophytes, and quantified pelagic and benthic biomass, production and transfer efficiencies for bacteria, phytoplankton, epipelon, epiphyton, macrophytes, zooplankton, macrozoobenthos and fish. We expected higher transfer efficiencies in the lake with macrophytes, because these provide shelter and food for macrozoobenthos and may thus enable a more efficient conversion of basal production to consumer production. In both lakes, the majority of the whole‐lake autotrophic and bacterial production was provided by benthic organisms, but whole‐lake primary consumer production mostly relied on pelagic autotrophic and bacterial production. Consequently, transfer efficiency of benthic autotrophic and bacterial production to macrozoobenthos production was an order of magnitude lower than the transfer efficiency of pelagic autotrophic and bacterial production to rotifer and crustacean production. Between‐lake differences in transfer efficiencies were minor. We discuss several aspects potentially causing the unexpectedly low benthic transfer efficiencies, such as the food quality of producers, pelagic–benthic links, oxygen concentrations in the deeper lake areas and additional unaccounted consumer production by pelagic and benthic protozoa and meiobenthos at intermediate or top trophic levels. None of these processes convincingly explain the large differences between benthic and pelagic transfer efficiencies. Our data indicate that shallow eutrophic lakes, even with a major share of autotrophic and bacterial production in the benthic zone, can function as pelagic systems with respect to primary consumer production. We suggest that the benthic autotrophic production was mostly transferred to benthic bacterial production, which remained in the sediments, potentially cycling internally in a similar way to what has previously been described for the microbial loop in pelagic habitats. Understanding the energetics of whole‐lake food webs, including the fate of the substantial benthic bacterial production, which is either mineralised at the sediment surface or permanently buried, has important implications for regional and global carbon cycling.