Species interactions, environmental gradients and body size shape population niche width.

Competition for shared resources is commonly assumed to restrict population-level niche width of coexisting species. However, the identity and abundance of coexisting species, the prevailing environmental conditions, and the individual body size may shape the effects of interspecific interactions on species' niche width. Here we study the effects of interspecific and intraspecific interactions, lake area and altitude, and fish body size on the trophic niche width and resource use of a generalist predator, the littoral-dwelling large, sparsely rakered morph of European whitefish (Coregonus lavaretus; hereafter LSR whitefish). We use stable isotope, diet and survey fishing data from 14 subarctic lakes along an environmental gradient in northern Norway. The isotopic niche width of LSR whitefish showed a humped-shaped relationship with increasing relative abundance of sympatric competitors, suggesting widest population niche at intermediate intensity of interspecific interactions. The isotopic niche width of LSR whitefish tended to decrease with increasing altitude, suggesting reduced niche in colder, less productive lakes. LSR whitefish typically shifted to a higher trophic position and increased reliance on littoral food resources with increasing body size, although between-lake differences in ontogenetic niche shifts were evident. In most lakes, LSR whitefish relied less on littoral food resources than coexisting fishes and the niche overlap between sympatric competitors was most evident among relatively large individuals (>250 mm). Individual niche variation was highest among >200 mm long LSR whitefish, which likely have escaped the predation window of sympatric predators. We demonstrate that intermediate intensity of interspecific interactions may broaden species' niche width, whereas strong competition for limited resources and high predation risk may suppress niche width in less productive environments. Acknowledging potential humped-shaped relationships between population niche width and interspecific interactions can help us understand species' responses to environmental disturbance (e.g. climate change and species invasions) as well as the driving forces of niche specialization.

Community structure influences species' abundance along environmental gradients.

Species' response to abiotic environmental variation can be influenced by local community structure and interspecific interactions, particularly in restricted habitats such as islands and lakes. In temperate lakes, future increase in water temperature and run-off of terrestrial (allochthonous) dissolved organic carbon (DOC) are predicted to alter community composition and the overall ecosystem productivity. However, little is known about how the present community structure and abiotic environmental variation interact to affect the abundance of native fish populations. We used a space-for-time approach to study how local community structure interact with lake morphometric and climatic characteristics (i.e. temperature and catchment productivity) to affect brown trout (Salmo trutta L.) yield in 283 Norwegian lakes located in different biogeographical regions. Brown trout yield (based on data from standardized survey gill net fishing; g 100 m(-2) gill net night(-1)) was generally lower in lakes where other fish species were present than in lakes with brown trout only. The yield showed an overall negative relationship with increasing temperature and a positive relationship with lake shoreline complexity. Brown trout yield was also negatively correlated with DOC load (measured using Normalized Difference Vegetation Index as a proxy) and lake size and depth (measured using terrain slope as a proxy), but only in lakes where other fish species were present. The observed negative response of brown trout yield to increasing DOC load and proportion of the pelagic open-water area is likely due to restricted (littoral) niche availability and competitive dominance of more pelagic fishes such as Arctic charr (Salvelinus alpinus (L.)). Our study highlights that, through competitive interactions, the local community structure can influence the response of a species' abundance to variation in abiotic conditions. Changes in biomass and niche use of top predators (such as the brown trout), associated with predicted changes in direct and indirect climatic factors, may have further influences on the structure and function of temperate lake ecosystems.

Consistent isotopic differences between Schistocephalus spp. parasites and their stickleback hosts.

Parasite-host systems show markedly variable patterns in isotopic fractionation: parasites can be either depleted or enriched in ¹5N and ¹³C as compared to their hosts. However, it remains unknown whether isotopic fractionation patterns are similar in comparable parasite-host systems from markedly different ecosystems. Results of this study show that large-sized Schistocephalus spp. endoparasites are consistently depleted in ¹5N (by on average -2.13 to -2.20 ‰) as compared to their nine-spined stickleback Pungitius pungitius and three-spined stickleback Gasterosteus aculeatus hosts. The differences between parasites and host for both δ¹5N and δ¹³C were consistent in both study systems despite marked biogeographical differences between the study localities. Although the stable isotope values in general were strongly correlated between the hosts and their parasites, Schistocephalus specimens occupying the same nine-spined stickleback host showed sometimes substantial individual variation in δ¹³C. This might be due to selective use of different carbon sources, or different metabolic or feeding rates. Further studies on selective feeding, physiology and metabolism of parasites are needed to better understand the role of parasites in the structure and functioning of aquatic food webs.

Effects of temporal abiotic drivers on the dynamics of an allometric trophic network model.

Current ecological research and ecosystem management call for improved understanding of the abiotic drivers of community dynamics, including temperature effects on species interactions and biomass accumulation. Allometric trophic network (ATN) models, which simulate material (carbon) transfer in trophic networks from producers to consumers based on mass-specific metabolic rates, provide an attractive framework to study consumer-resource interactions from organisms to ecosystems. However, the developed ATN models rarely consider temporal changes in some key abiotic drivers that affect, for example, consumer metabolism and producer growth. Here, we evaluate how temporal changes in carrying capacity and light-dependent growth rate of producers and in temperature-dependent mass-specific metabolic rate of consumers affect ATN model dynamics, namely seasonal biomass accumulation, productivity, and standing stock biomass of different trophic guilds, including age-structured fish communities. Our simulations of the pelagic Lake Constance food web indicated marked effects of temporally changing abiotic parameters on seasonal biomass accumulation of different guild groups, particularly among the lowest trophic levels (primary producers and invertebrates). While the adjustment of average irradiance had minor effect, increasing metabolic rate associated with 1-2°C temperature increase led to a marked decline of larval (0-year age) fish biomass, but to a substantial biomass increase of 2- and 3-year-old fish that were not predated by ≥4-year-old top predator fish, European perch (Perca fluviatilis). However, when averaged across the 100 simulation years, the inclusion of seasonality in abiotic drivers caused only minor changes in standing stock biomasses and productivity of different trophic guilds. Our results demonstrate the potential of introducing seasonality in and adjusting the average values of abiotic ATN model parameters to simulate temporal fluctuations in food-web dynamics, which is an important step in ATN model development aiming to, for example, assess potential future community-level responses to ongoing environmental changes.

Impacts of piscicide-induced fish removal on resource use and trophic diversity of lake invertebrates.

Chemical eradication of non-native species has become a widely used method to mitigate the potential negative impacts of altered competitive or predatory dynamics on biodiversity and natural ecosystem processes. However, the responses of non-target species can vary from rapid full recovery to delayed or absent recolonization, and little is known about the potential shifts in resource use and trophic diversity of native species following chemical treatments. We used a before-after-control-impact approach to study the effects of rotenone piscicide treatment on abundance and trophic niche of benthic invertebrates in three untreated and three treated lakes in central Norway, the latter group hosting non-native roach (Rutilus rutilus) and pike (Esox lucius) prior to rotenone treatment. Based on community composition data, the relative abundance of invertebrate grazers and collectors decreased while that of predators increased following fish removal in the treated lakes. The stable isotope data indicated minor shifts in resource use of, and trophic diversity among, benthic invertebrate communities. While the predatory dragonfly larvae (Odonata) and grazer snails (Lymnaeidae) showed increased δ13C values indicating increased reliance on littoral benthic algae, the collector mayfly larvae (Leptophlebia) showed decreased δ13C values following fish removal in treated lakes. Grazer snails also showed a shift to a lower trophic position, while the predatory dragonflies and collector mayflies showed no changes in δ15N values following fish removal. The community-level isotopic niches of benthic invertebrates showed no consistent changes, although the sample-size corrected and Bayesian estimates of standard ellipse areas (SEAC and SEAB) slightly increased in two of the three treated lakes due to an increased range in δ15N. In conclusion, our study findings indicate some changes in species assemblages but minor shifts in the resource use and trophic diversity of benthic invertebrate communities following fish removal in rotenone treated lakes.

Contrasting structural complexity differentiate hunting strategy in an ambush apex predator.

Structural complexity is known to influence prey behaviour, mortality and population structure, but the effects on predators have received less attention. We tested whether contrasting structural complexity in two newly colonised lakes (low structural complexity lake-LSC; high structural complexity-HSC) was associated with contrasting behaviour in an aquatic apex predator, Northern pike (Esox lucius; hereafter pike) present in the lakes. Behaviour of pike was studied with whole-lake acoustic telemetry tracking, supplemented by stable isotope analysis of pike prey utilization and survey fishing data on the prey fish community. Pike displayed increased activity, space use, individual growth as well as behavioural differentiation and spent more time in open waters in the LSC lake. Despite observed differences between lakes, stable isotopes analyses indicated a high dependency on littoral food sources in both lakes. We concluded that pike in the HSC lake displayed a behaviour consistent with a prevalent ambush predation behaviour, whereas the higher activity and larger space use in the LSC lake indicated a transition to more active search behaviour. It could lead to increased prey encounter and cause better growth in the LSC lake. Our study demonstrated how differences in structural complexity mediated prominent changes in the foraging behaviour of an apex predator, which in turn may have effects on the prey community.

Hydropower impacts on reservoir fish populations are modified by environmental variation.

Global transition towards renewable energy production has increased the demand for new and more flexible hydropower operations. Before management and stakeholders can make informed choices on potential mitigations, it is essential to understand how the hydropower reservoir ecosystems respond to water level regulation (WLR) impacts that are likely modified by the reservoirs' abiotic and biotic characteristics. Yet, most reservoir studies have been case-specific, which hampers large-scale planning, evaluation and mitigation actions across various reservoir ecosystems. Here, we investigated how the effect of the magnitude, frequency and duration of WLR on fish populations varies along environmental gradients. We used biomass, density, size, condition and maturation of brown trout (Salmo trutta L.) in Norwegian hydropower reservoirs as a measure of ecosystem response, and tested for interacting effects of WLR and lake morphometry, climatic conditions and fish community structure. Our results showed that environmental drivers modified the responses of brown trout populations to different WLR patterns. Specifically, brown trout biomass and density increased with WLR magnitude particularly in large and complex-shaped reservoirs, but the positive relationships were only evident in reservoirs with no other fish species. Moreover, increasing WLR frequency was associated with increased brown trout density but decreased condition of individuals within the populations. WLR duration had no significant impacts on brown trout, and the mean weight and maturation length of brown trout showed no significant response to any WLR metrics. Our study demonstrates that local environmental characteristics and the biotic community strongly modify the hydropower-induced WLR impacts on reservoir fishes and ecosystems, and that there are no one-size-fits-all solutions to mitigate environmental impacts. This knowledge is vital for sustainable planning, management and mitigation of hydropower operations that need to meet the increasing worldwide demand for both renewable energy and ecosystem services delivered by freshwaters.

Community structure affects trophic ontogeny in a predatory fish.

While most studies have focused on the timing and nature of ontogenetic niche shifts, information is scarce about the effects of community structure on trophic ontogeny of top predators. We investigated how community structure affects ontogenetic niche shifts (i.e., relationships between body length, trophic position, and individual dietary specialization) of a predatory fish, brown trout (Salmo trutta). We used stable isotope and stomach content analyses to test how functional characteristics of lake fish community compositions (competition and prey availability) modulate niche shifts in terms of (i) piscivorous behavior, (ii) trophic position, and (iii) individual dietary specialization. Northern Scandinavian freshwater fish communities were used as a study system, including nine subarctic lakes with contrasting fish community configurations: (i) trout-only systems, (ii) two-species systems (brown trout and Arctic charr [Salvelinus alpinus] coexisting), and (iii) three-species systems (brown trout, Arctic charr, and three-spined sticklebacks [Gasterosteus aculeatus] coexisting). We expected that the presence of profitable small prey (stickleback) and mixed competitor-prey fish species (charr) supports early piscivory and high individual dietary specialization among trout in multispecies communities, whereas minor ontogenetic shifts were expected in trout-only systems. From logistic regression models, the presence of a suitable prey fish species (stickleback) emerged as the principal variable determining the size at ontogenetic niche shifts. Generalized additive mixed models indicated that fish community structure shaped ontogenetic niche shifts in trout, with the strongest positive relationships between body length, trophic position, and individual dietary specialization being observed in three-species communities. Our findings revealed that the presence of a small-sized prey fish species (stickleback) rather than a mixed competitor-prey fish species (charr) was an important factor affecting the ontogenetic niche-shift processes of trout. The study demonstrates that community structure may modulate the ontogenetic diet trajectories of and individual niche specialization within a top predator.

European catfish (Silurus glanis) as a freshwater apex predator drives ecosystem via its diet adaptability.

Apex predators play a key role in ecosystem stability across environments but their numbers in general are decreasing. By contrast, European catfish (Silurus glanis), the European freshwater apex predator, is on the increase. However, studies concerning apex predators in freshwaters are scarce in comparison to those in terrestrial and marine ecosystems. The present study combines stomach content and stable isotope analyses with diet preferences of catfish to reveal its impact on the ecosystem since stocking. Catfish niche width is extremely wide in comparison to the typical model predator, Northern pike (Esox lucius). Catfish and pike have different individual dietary specialization that results in different functional roles in coupling or compartmentalizing distinct food webs. The role of both species in the ecosystem is irreplaceable due to multiple predator effects. The impact of catfish is apparent across the entire aquatic ecosystem, but herbivores are the most affected ecological group. The key feature of catfish, and probably a common feature of apex predators in general, is utilization of several dietary strategies by individuals within a population: long-term generalism or specialization and also short-term specialization. Catfish, similar to other large-bodied apex predators, have two typical features: enormous generalism and adaptability to new prey sources.

Macrophytes shape trophic niche variation among generalist fishes.

Generalist species commonly have a fundamental role in ecosystems as they can integrate spatially distinct habitats and food-web compartments, as well as control the composition, abundance and behavior of organisms at different trophic levels. Generalist populations typically consist of specialized individuals, but the potential for and hence degree of individual niche variation can be largely determined by habitat complexity. We compared individual niche variation within three generalist fishes between two comparable lakes in the Czech Republic differing in macrophyte cover, i.e. macrophyte-rich Milada and macrophyte-poor Most. We tested the hypothesis that large individual niche variation among generalist fishes is facilitated by the presence of macrophytes, which provides niches and predation shelter for fish and their prey items. Based on results from stable nitrogen (δ15N) and carbon (δ13C) isotopic mixing models, perch (Perca fluviatilis L.) and rudd (Scardinius erythrophthalmus (L.)) showed larger individual variation (i.e., variance) in trophic position in Milada as compared to Most, whereas no significant between-lake differences were observed for roach (Rutilus rutilus (L.)). Contrary to our hypothesis, all the three species showed significantly lower individual variation in the relative reliance on littoral food resources in Milada than in Most. Rudd relied significantly more whereas perch and roach relied less on littoral food resources in Milada than in Most, likely due to prevalent herbivory by rudd and prevalent zooplanktivory by perch and roach in the macrophyte-rich Milada as compared to macrophyte-poor Most. Our study demonstrates how the succession of macrophyte vegetation, via its effects on the physical and biological complexity of the littoral zone and on the availability of small prey fish and zooplankton, can strongly influence individual niche variation among generalist fishes with different ontogenetic trajectories, and hence the overall food-web structures in lake ecosystems.

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.