Shortened food chain length in a fished versus unfished coral reef.

Direct exploitation through fishing is driving dramatic declines of wildlife populations in ocean environments, particularly for predatory and large-bodied taxa. Despite wide recognition of this pattern and well-established consequences of such trophic downgrading on ecosystem function, there have been few empirical studies examining the effects of fishing on whole system trophic architecture. Understanding these kinds of structural impacts is especially important in coral reef ecosystems-often heavily fished and facing multiple stressors. Given the often high dietary flexibility and numerous functional redundancies in diverse ecosystems such as coral reefs, it is important to establish whether web architecture is strongly impacted by fishing pressure or whether it might be resilient, at least to moderate-intensity pressure. To examine this question, we used a combination of bulk and compound-specific stable isotope analyses measured across a range of predatory and low-trophic-level consumers between two coral reef ecosystems that differed with respect to fishing pressure but otherwise remained largely similar. We found that even in a high-diversity system with relatively modest fishing pressure, there were strong reductions in the trophic position (TP) of the three highest TP consumers examined in the fished system but no effects on the TP of lower-level consumers. We saw no evidence that this shortening of the affected food webs was being driven by changes in basal resource consumption, for example, through changes in the spatial location of foraging by consumers. Instead, this likely reflected internal changes in food web architecture, suggesting that even in diverse systems and with relatively modest pressure, human harvest causes significant compressions in food chain length. This observed shortening of these food webs may have many important emergent ecological consequences for the functioning of ecosystems impacted by fishing or hunting. Such important structural shifts may be widespread but unnoticed by traditional surveys. This insight may also be useful for applied ecosystem managers grappling with choices about the relative importance of protection for remote and pristine areas and the value of strict no-take areas to protect not just the raw constituents of systems affected by fishing and hunting but also the health and functionality of whole systems.

Towards a mechanistic understanding of variation in aquatic food chain length.

Ecologists have long sought to understand variation in food chain length (FCL) among natural ecosystems. Various drivers of FCL, including ecosystem size, resource productivity and disturbance, have been hypothesised. However, when results are aggregated across existing empirical studies from aquatic ecosystems, we observe mixed FCL responses to these drivers. To understand this variability, we develop a unified competition-colonisation framework for complex food webs incorporating all of these drivers. With competition-colonisation tradeoffs among basal species, our model predicts that increasing ecosystem size generally results in a monotonic increase in FCL, while FCL displays non-linear, oscillatory responses to resource productivity or disturbance in large ecosystems featuring little disturbance or high productivity. Interestingly, such complex responses mirror patterns in empirical data. Therefore, this study offers a novel mechanistic explanation for observed variations in aquatic FCL driven by multiple environmental factors.

You are what your host eats: The trophic structure and food chain length of a symbiont community are coupled with the plastic diet of the host ant.

Food chain length provides key information on the flow of nutrients and energy in ecosystems. Variation in food chain length has primarily been explained by environmental drivers such as ecosystem size and productivity. Most insights are obtained from theory or aquatic systems, but the importance of these drivers remains largely untested in terrestrial systems. We exploited red wood ant nests markedly differing in size as natural experiments to quantify the drivers of trophic structure and food chain length of their symbiont arthropod communities. Using stable isotopes, we explored the variation in the trophic positions of four symbiont species with the trophic position of the top predator as a proxy for food chain length of the symbiont community. Nest size did not affect food chain length, nor trophic distance between the symbionts. Instead, food chain length and the trophic positions of the symbionts were strongly affected by the host's foraging decisions. When the host diet shifted from predominantly herbivorous to more predacious, the trophic position of the symbionts and food chain length strongly increased. We show for the first time that a food web can be structured by biotic interactions with an engineering species rather than by abiotic environmental variables.

Inquiline predator increases nutrient-cycling efficiency of Nepenthes rafflesiana pitchers.

The modified-leaf pitchers of Nepenthes rafflesiana pitcher plants are aquatic, allochthonous ecosystems that are inhabited by specialist inquilines and sustained by the input of invertebrate prey. Detritivorous inquilines are known to increase the nutrient-cycling efficiency (NCE) of pitchers but it is unclear whether predatory inquilines that prey on these detritivores decrease the NCE of pitchers by reducing detritivore populations or increase the NCE of pitchers by processing nutrients that may otherwise be locked up in detritivore biomass. Nepenthosyrphus is a small and poorly studied genus of hoverflies and the larvae of one such species is a facultatively detritivorous predator that inhabits the pitchers of N. rafflesiana. We fitted a consumer-resource model to experimental data collected from this system. Simulations showed that systems containing the predator at equilibrium almost always had higher NCEs than those containing only prey (detritivore) species. We showed using a combination of simulated predator/prey exclusions that the processing of the resource through multiple pathways and trophic levels in this system is more efficient than that accomplished through fewer pathways and trophic levels. Our results thus support the vertical diversity hypothesis, which predicts that greater diversity across trophic levels results in greater ecosystem functioning.

Longer Food Chains in Pelagic Ecosystems: Trophic Energetics of Animal Body Size and Metabolic Efficiency.

Factors constraining the structure of food webs can be investigated by comparing classes of ecosystems. We find that pelagic ecosystems, those based on one-celled primary producers, have longer food chains than terrestrial ecosystems. Yet pelagic ecosystems have lower primary productivity, contrary to the hypothesis that greater energy flows permit higher trophic levels. We hypothesize that longer food chain length in pelagic ecosystems, compared with terrestrial ecosystems, is associated with smaller pelagic animal body size permitting more rapid trophic energy transfer. Assuming negative allometric dependence of biomass production rate on body mass at each trophic level, the lowest three pelagic animal trophic levels are estimated to add biomass more rapidly than their terrestrial counterparts by factors of 12, 4.8, and 2.6. Pelagic animals consequently transport primary production to a fifth trophic level 50-190 times more rapidly than animals in terrestrial webs. This difference overcomes the approximately fivefold slower pelagic basal productivity, energetically explaining longer pelagic food chains. In addition, ectotherms, dominant at lower pelagic animal trophic levels, have high metabolic efficiency, also favoring higher rates of trophic energy transfer in pelagic ecosystems. These two animal trophic flow mechanisms imply longer pelagic food chains, reestablishing an important role for energetics in food web structure.

Inter-annual changes in detritus-based food chains can enhance plant growth response to elevated atmospheric CO2.

Elevated atmospheric CO2 generally enhances plant growth, but the magnitude of the effects depend, in part, on nutrient availability and plant photosynthetic pathway. Due to their pivotal role in nutrient cycling, changes in abundance of detritivores could influence the effects of elevated atmospheric CO2 on essential ecosystem processes, such as decomposition and primary production. We conducted a field survey and a microcosm experiment to test the influence of changes in detritus-based food chains on litter mass loss and plant growth response to elevated atmospheric CO2 using two wetland plants: a C3 sedge (Scirpus olneyi) and a C4 grass (Spartina patens). Our field study revealed that organism's sensitivity to climate increased with trophic level resulting in strong inter-annual variation in detritus-based food chain length. Our microcosm experiment demonstrated that increased detritivore abundance could not only enhance decomposition rates, but also enhance plant growth of S. olneyi in elevated atmospheric CO2 conditions. In contrast, we found no evidence that changes in the detritus-based food chains influenced the growth of S. patens. Considered together, these results emphasize the importance of approaches that unite traditionally subdivided food web compartments and plant physiological processes to understand inter-annual variation in plant production response to elevated atmospheric CO2.

Limits on ecosystem trophic complexity: insights from ecological network analysis.

Articulating what limits the length of trophic food chains has remained one of the most enduring challenges in ecology. Mere counts of ecosystem species and transfers have not much illumined the issue, in part because magnitudes of trophic transfers vary by orders of magnitude in power-law fashion. We address this issue by creating a suite of measures that extend the basic indexes usually obtained by counting taxa and transfers so as to apply to networks wherein magnitudes vary by orders of magnitude. Application of the extended measures to data on ecosystem trophic networks reveals that the actual complexity of ecosystem webs is far less than usually imagined, because most ecosystem networks consist of a multitude of weak connections dominated by a relatively few strong flows. Although quantitative ecosystem networks may consist of hundreds of nodes and thousands of transfers, they nevertheless behave similarly to simpler representations of systems with fewer than 14 nodes or 40 flows. Both theory and empirical data point to an upper bound on the number of effective trophic levels at about 3-4 links. We suggest that several whole-system processes may be at play in generating these ecosystem limits and regularities.

Prey state shapes the effects of temporal variation in predation risk.

The ecological impacts of predation risk are influenced by how prey allocate foraging effort across periods of safety and danger. Foraging decisions depend on current danger, but also on the larger temporal, spatial or energetic context in which prey manage their risks of predation and starvation. Using a rocky intertidal food chain, we examined the responses of starved and fed prey (Nucella lapillus dogwhelks) to different temporal patterns of risk from predatory crabs (Carcinus maenas). Prey foraging activity declined during periods of danger, but as dangerous periods became longer, prey state altered the magnitude of risk effects on prey foraging and growth, with likely consequences for community structure (trait-mediated indirect effects on basal resources, Mytilus edulis mussels), prey fitness and trophic energy transfer. Because risk is inherently variable over time and space, our results suggest that non-consumptive predator effects may be most pronounced in productive systems where prey can build energy reserves during periods of safety and then burn these reserves as 'trophic heat' during extended periods of danger. Understanding the interaction between behavioural (energy gain) and physiological (energy use) responses to risk may illuminate the context dependency of trait-mediated trophic cascades and help explain variation in food chain length.

Migratory-derived resources induce elongated food chains through middle-up food web effects.

BACKGROUND: Seasonal movements of animals often result in the transfer of large amounts of energy and nutrients across ecosystem boundaries, which may have large consequences on local food webs through various pathways. While this is known for both terrestrial- and aquatic organisms, quantitative estimates on its effects on food web structure and identification of key pathways are scarce, due to the difficulty in obtaining replication on ecosystem level with negative control, i.e. comparable systems without migration. METHODS: In this study, we estimate the impact of Arctic charr (Salvelinus alpinus) migration on riverine ecosystem structure, by comparing multiple streams with strictly resident populations above natural migration barriers with streams below those barriers harboring partially migratory populations. We compared density estimates and size structure between above and below populations. Diet differences were examined through the analysis of stomach contents, changes in trophic position were examined by using stable isotopes. To infer growth rate of resident individuals, back-growth calculation was performed using otoliths. RESULTS: We find higher densities of small juveniles in partially migratory populations, where juvenile Arctic charr show initially lower growth, likely due to higher intraspecific competition. After reaching a size, where they can start feeding on eggs and smaller juveniles, which are both more frequent in partially migratory populations, growth surpasses that of resident populations. Cannibalism induced by high juvenile densities occurred almost exclusively in populations with migration and represents an altered energy pathway to the food web. The presence of large cannibalistic charr feeding on smaller ones that have a similar trophic level as charr from strictly resident populations (based on stomach content) coupled with steeper δ15N-size regression slopes illustrate the general increase of food chain length in systems with migration. CONCLUSIONS: Our results thus suggest that the consumption of migration-derived resources may result in longer food chains through middle-up rather than bottom-up effects. Furthermore, by occupying the apex of the food chain and feeding on juvenile conspecifics, resident individuals experience reduced competition with their young counterparts, which potentially balances their fitness with migratory individuals.

Long-term recovery of benthic food webs after stream restoration.

The assessment of restoration success often neglects trophic interactions within food webs, focusing instead on biodiversity and community structure. Here, we analysed the long-term recovery of food web structure based on stable isotopes (δ13C and δ15N) of benthic invertebrates and quantified responses of food web metrics to time since restoration. The samples derived from twelve restored sites with different restoration ages, sampled annually from 2012 to 2021, and covering an investigation period of up to 28 years after restoration for the whole catchment. Temporal developments of the restored sites were compared to the development of two near-natural sites. The restoration measures consisted of the cessation of sewage inflow and morphological restoration of the channels. As a clear and consistent result over almost all sites, trophic similarity (proportion of co-existing species occupying similar trophic niches) increased with time since restoration, and reached values of near-natural sites, suggesting an increase in the stability and resilience of the food webs. Surprisingly, resource diversity decreased at most restored sites within 10 years after restoration, probably due to the removal of wastewater-derived resources, and a shift towards leaf litter as the dominant resource following the regrowth of the riparian vegetation. Food chain length showed no consistent pattern over time at the different sites both increasing and decreasing with time since restoration. Overall, restoration had clear effects on the food web structure of stream ecosystems. While some effects such as the increase in trophic similarity were consistent at almost all sites, others such as response of the food chain length were context dependent. The study demonstrates the potential of utilizing food web metrics, particularly trophic similarity, in restoration research to achieve a more holistic understanding of ecosystem recovery.

Food webs in isolation: The food-web structure of a freshwater reservoir with armoured shores in a former coastal bay area.

Many shallow coastal bays have been closed off from the sea to mitigate the risk of flooding, resulting in coastal reservoir lakes with artificial armoured shorelines. Often these enclosed ecosystems show a persistent decline in biodiversity and ecosystem services, which is likely reflected in their food-web structure. We therefore hypothesize that the food webs of coastal reservoir lakes with armoured shorelines (1) consist of relatively few species with a low food-web connectance and short food chains, and (2) are mainly fuelled by autochthonous organic matter produced in the pelagic zone. To investigate these two hypotheses, we used stable-isotope analysis to determine the food-web structure of lake Markermeer (The Netherlands), a large reservoir lake with armoured shorelines in a former coastal bay area. Contrary to expectation, connectance of the food web in lake Markermeer was comparable to other lakes, while food-chain length was in the higher range. However, the trophic links revealed that numerous macroinvertebrates and fish species in this constructed lake exhibited omnivorous feeding behaviour. Furthermore, in line with our second hypothesis, primary consumers heavily relied on pelagically derived organic matter, while benthic primary production exerted only a minor and seasonal influence on higher trophic levels. Stable-isotope values and the C:N ratio of sediment organic matter in the lake also aligned more closely with phytoplankton than with benthic primary producers. Moreover, terrestrial subsidies of organic matter were virtually absent in lake Markermeer. These findings support the notion that isolation of the lake through shore armouring and the lack of littoral habitats in combination with persistent resuspension of sediments have affected the food web. We argue that restoration initiatives should prioritize the establishment of land-water transition zones, thereby enhancing habitat diversity, benthic primary production, and the inflow of external organic matter while preserving pelagic primary production.

Assessment of the impact of dams on aquatic food webs using stable isotopes: Current progress and future challenges.

Dams have disrupted natural river systems worldwide and although population and community level effects on aquatic biota have been well documented, food web responses remain poorly understood and difficult to characterize. The application of stable isotope analysis (SIA) provides a means to assess the effect of dams on food webs. Here we review the effect of dams on aquatic food webs using SIA, aiming to detect knowledge gaps in the field of dam impacts on aquatic food webs and propose a conceptual framework to help formulate hypotheses about dam impacts on food webs guided by food web theory. Dams can affect aquatic food webs via two pathways: a bottom-up pathway with altered basal food sources and their transfer to consumers through changes in flow, nutrients, temperature and sediment, and a top-down pathway with consumer species composition altered mainly through habitat fragmentation and related physiochemical changes. Taking these mechanisms into consideration, the impact of dams on food web attributes derived from SIA was evaluated. These studies generally apply mixing models to determine how dams alter the dominant carbon sources supporting food webs, use δ15N to examine how dams alter food-chain length, or use Layman metrics of isotope variability to assess niche changes for invertebrate and fish assemblages. Most studies compare the patterns of SIA metrics spatially (e.g. upstream vs reservoir vs downstream of dams; regulated vs unregulated rivers) and temporally (before vs after dam construction), without explicit hypotheses and/or links to theoretical concepts of food webs. We propose several steps to make SIA studies of dam impacts more rigorous and enhance their potential for producing novel insights. Future studies should quantify the shape and strength of the effect of dams on SIA-measured food web response, be conducted at larger temporal and spatial scales (particularly along the river longitudinal continuum and the lateral connected ecosystems (e.g., floodplains)), and consider effects of dams on food web resilience and tipping points.

Environmental and biological factors are joint drivers of mercury biomagnification in subarctic lake food webs along a climate and productivity gradient.

Subarctic lakes are getting warmer and more productive due to the joint effects of climate change and intensive land-use practices (e.g. forest clear-cutting and peatland ditching), processes that potentially increase leaching of peat- and soil-stored mercury into lake ecosystems. We sampled biotic communities from primary producers (algae) to top consumers (piscivorous fish), in 19 subarctic lakes situated on a latitudinal (69.0-66.5° N), climatic (+3.2 °C temperature and +30% precipitation from north to south) and catchment land-use (pristine to intensive forestry areas) gradient. We first tested how the joint effects of climate and productivity influence mercury biomagnification in food webs focusing on the trophic magnification slope (TMS) and mercury baseline (THg baseline) level, both derived from linear regression between total mercury (log10THg) and organism trophic level (TL). We examined a suite of environmental and biotic variables thought to explain THg baseline and TMS with stepwise generalized multiple regression models. Finally, we assessed how climate and lake productivity affect the THg content of top predators in subarctic lakes. We found biomagnification of mercury in all studied lakes, but with variable TMS and THg baseline values. In stepwise multiple regression models, TMS was best explained by negative relationships with food chain length, climate-productivity gradient, catchment properties, and elemental C:N ratio of the top predator (full model R2 = 0.90, p < 0.001). The model examining variation in THg baseline values included the same variables with positive relationships (R2 = 0.69, p = 0.014). Mass-standardized THg content of a common top predator (1 kg northern pike, Esox lucius) increased towards warmer and more productive lakes. Results indicate that increasing eutrophication via forestry-related land-use activities increase the THg levels at the base of the food web and in top predators, suggesting that the sources of nutrients and mercury should be considered in future bioaccumulation and biomagnification studies.

Multitrophic diversity sustains ecological complexity by dampening top-down control of a shallow marine benthic food web.

Biodiversity is typically considered as a one-dimensional metric (e.g., species richness), yet the consequences of species loss may be different depending on where extinctions occur in the food web. Here, I used a manipulative field experiment in a temperate subtidal marine system to explore the implications of diversity loss at multiple trophic levels for ecosystem functioning and food web structure. The four manipulated predators included the small painted goby and common prawn, which are also fed on by the larger black goby and shore crab. Antagonistic interactions between the manipulated predators (e.g., intraguild predation, intimidation, interference competition) limited their negative effects on the rest of the food web. Top-down control was so suppressed at the highest level of multitrophic diversity that the resulting food webs were as complex and productive as those containing no manipulated predators. Negative interactions between the predators weakened as multitrophic diversity decreased, however, resulting in stronger consumption of lower trophic levels and a simpler food web with lower rates of two key ecosystem processes: primary production and decomposition. These results show how indirect interactions between predators on multiple trophic levels help to promote the complexity and functioning of natural systems.

Environmental correlates of food-chain length, mean trophic level and trophic level variance in invaded riverine fish assemblages.

Examining how the trophic structure of biotic assemblages is affected by human impacts, such as habitat degradation and the introduction of alien species, is important for understanding the consequences of such impacts on ecosystem functioning. We used general linear mixed models and hierarchical partitioning analyses of variance to examine for the first time the applicability of three hypotheses (ecosystem-size, productivity and disturbance) for explaining food-chain length (FCL) in invaded fish assemblages. We used Fishbase trophic level (TL) estimates for 16 native and 18 alien fish species in an extensive riverine system in north-eastern Spain (99,700 km2, 15 catchments, 530 sites). The FCL of assemblages ranged from 2.7 to 4.42. Ecosystem size-related variables (Strahler stream order, physical habitat diversity) and human-disturbance (conductivity) made the largest contribution to the explained variance in the FCL model after accounting for spatial confounding factors and collinearity among predictors. Within-assemblage TL also was positively associated with Strahler stream order, suggesting that large rivers have the highest trophic diversity. High conductivity was negatively associated with FCL, as did with the mean TL of fish assemblages. However, an inverse association was found between mean TL and Strahler stream order, possibly because the presence of fish species of high TL may be offset by larger numbers of alien species of lower TL in large rivers. Given that there may be trophic replacements among native and alien species, this inference needs to be addressed with detailed trophic studies. However, reducing water conductivity by improved wastewater treatment and better agricultural practices probably would help to conserve the fish species on the apices of aquatic food-webs.

Lake size and fish diversity determine resource use and trophic position of a top predator in high-latitude lakes.

Prey preference of top predators and energy flow across habitat boundaries are of fundamental importance for structure and function of aquatic and terrestrial ecosystems, as they may have strong effects on production, species diversity, and food-web stability. In lakes, littoral and pelagic food-web compartments are typically coupled and controlled by generalist fish top predators. However, the extent and determinants of such coupling remains a topical area of ecological research and is largely unknown in oligotrophic high-latitude lakes. We analyzed food-web structure and resource use by a generalist top predator, the Arctic charr Salvelinus alpinus (L.), in 17 oligotrophic subarctic lakes covering a marked gradient in size (0.5-1084 km(2)) and fish species richness (2-13 species). We expected top predators to shift from littoral to pelagic energy sources with increasing lake size, as the availability of pelagic prey resources and the competition for littoral prey are both likely to be higher in large lakes with multispecies fish communities. We also expected top predators to occupy a higher trophic position in lakes with greater fish species richness due to potential substitution of intermediate consumers (prey fish) and increased piscivory by top predators. Based on stable carbon and nitrogen isotope analyses, the mean reliance of Arctic charr on littoral energy sources showed a significant negative relationship with lake surface area, whereas the mean trophic position of Arctic charr, reflecting the lake food-chain length, increased with fish species richness. These results were supported by stomach contents data demonstrating a shift of Arctic charr from an invertebrate-dominated diet to piscivory on pelagic fish. Our study highlights that, because they determine the main energy source (littoral vs. pelagic) and the trophic position of generalist top predators, ecosystem size and fish diversity are particularly important factors influencing function and structure of food webs in high-latitude lakes.

Trophic network of aquatic macroinvertebrates along an altitudinal gradient in a Neotropical mountain river

ABSTRACT Studies of trophic networks and the evaluation of processes that occur along altitudinal gradients in river systems are of great importance because they allow an understanding of energy flow dynamics and provide scientific tools for the planning and management of river ecosystems. This research describes the trophic network of aquatic macroinvertebrates along an altitudinal gradient of the Gaira River, a mountain Neotropical watercourse located in the Sierra Nevada de Santa Marta in northern Colombia. The organisms were collected in the upper, middle and lower reaches of the river during the rainy and dry seasons (between 2010 and 2013). Trophic relationships were evaluated through gut content analysis. The contents were determined and quantified using photographs and expert review, and a binary consumption matrix was used to determine the characteristics of the trophic network. We characterized the diet composition at each site for each season using discriminant analysis. Trophic networks during the dry seasons showed higher trophic species richness and linkage density, and the predominance of coarse particulate organic matter (CPOM) and fine particulate organic matter (FPOM) in the high and medium sections of the river. During the dry seasons the diets had a lower number of basal species, but in the low river section there was a high percentage of fungi and microalgae. During the rainy seasons, no patterns were observed for the percentage of resources. Results indicated a direct relation between periods of hydrologic stability and an increase of CPOM during dry seasons and an increase of resource diversity consumed by macroinvertebrates at all sites during the rainy season, showing that changes in trophic networks of the Gaira River were more important during seasonal periods than along the altitudinal gradient.

ECOSYSTEM SIZE, BUT NOT DISTURBANCE, DETERMINES FOOD-CHAIN LENGTH ON ISLANDS OF THE BAHAMAS.

Ecologists have long struggled to explain variation in food-chain length among natural ecosystems. Food-chain length is predicted to be shorter in ecosystems subjected to greater disturbance because longer chains are theoretically less resilient to perturbation. Moreover, food-chain length is expected to be longer in larger ecosystems because increasing ecosystem size increases species richness and stabilizes predator-prey interactions, or increases total resource availability. Here we test the roles of disturbance and ecosystem size in determining the food-chain length of terrestrial food webs on Bahamian islands. We found that disturbance affected the identity of top predators, but did not change food-chain length because alternative top predators occupied similar trophic positions. On the other hand, a 106 -fold increase in ecosystem size elevated food-chain length by one trophic level. We suggest that the effect of disturbance on food-chain length is weak when alternate top predators are trophic omnivores and have similar trophic positions. This and previous work in lakes suggest that ecosystem size may be a strong determinant of food-chain length in both aquatic and terrestrial ecosystems.