Riparian buffers maintain aquatic trophic structure in agricultural landscapes.

Local and regional habitat conditions associated with agricultural activity can fundamentally alter aquatic ecosystems. Increased nutrient inputs, channelization and reduced riparian habitat both upstream and locally contribute to the degradation of stream ecosystems and their function. Here, we examine stream food webs in watersheds that feed into Lake Erie to determine the effects of agricultural land cover on major food web energy pathways and trophic structure. Given that higher agricultural intensity can increase nutrient runoff and reduce the riparian zone and litter in-fall into streams, we predicted that generalist fish would derive less energy from the terrestrial pathway and become more omnivorous. Consistent with these predictions, we show that both mean terrestrial energy use and trophic position of the resident top consumer, creek chub (Semotilus atromaculatus), decrease with local agricultural intensity but not with watershed-level agriculture intensity. These findings suggest that local riparian buffers can maintain trophic structure even in the face of high whole-watershed agricultural intensity.

Winter in water: differential responses and the maintenance of biodiversity.

The ecological consequences of winter in freshwater systems are an understudied but rapidly emerging research area. Here, we argue that winter periods of reduced temperature and light (and potentially oxygen and resources) could play an underappreciated role in mediating the coexistence of species. This may be especially true for temperate and subarctic lakes, where seasonal changes in the thermal environment might fundamentally structure species interactions. With climate change already shortening ice-covered periods on temperate and polar lakes, consideration of how winter conditions shape biotic interactions is urgently needed. Using freshwater fishes in northern temperate lakes as a case study, we demonstrate how physiological trait differences (e.g. thermal preference, light sensitivity) drive differential behavioural responses to winter among competing species. Specifically, some species have a higher capacity for winter activity than others. Existing and new theory is presented to argue that such differential responses to winter can promote species coexistence. Importantly, if winter is a driver of niche differences that weaken competition between, relative to within species, then shrinking winter periods could threaten coexistence by tipping the scales in favour of certain sets of species over others.

Behavioral responses to annual temperature variation alter the dominant energy pathway, growth, and condition of a cold-water predator.

There is a pressing need to understand how ecosystems will respond to climate change. To date, no long-term empirical studies have confirmed that fish populations exhibit adaptive foraging behavior in response to temperature variation and the potential implications this has on fitness. Here, we use an unparalleled 11-y acoustic telemetry, stable isotope, and mark-recapture dataset to test if a population of lake trout (Salvelinus namaycush), a cold-water stenotherm, adjusted its use of habitat and energy sources in response to annual variations in lake temperatures during the open-water season and how these changes translated to the growth and condition of individual fish. We found that climate influenced access to littoral regions in spring (data from telemetry), which in turn influenced energy acquisition (data from isotopes), and growth (mark-recapture data). In more stressful years, those with shorter springs and longer summers, lake trout had reduced access to littoral habitat and assimilated less littoral energy, resulting in reduced growth and condition. Annual variation in prey abundance influenced lake trout foraging tactics (i.e., the balance of the number and duration of forays) but not the overall time spent in littoral regions. Lake trout greatly reduced their use of littoral habitat and occupied deep pelagic waters during the summer. Together, our results provide clear evidence that climate-mediated behavior can influence the dominant energy pathways of top predators, with implications ranging from individual fitness to food web stability.

Homogenization of freshwater lakes: Recent compositional shifts in fish communities are explained by gamefish movement and not climate change.

Globally, lake fish communities are being subjected to a range of scale-dependent anthropogenic pressures, from climate change to eutrophication, and from overexploitation to species introductions. As a consequence, the composition of these communities is being reshuffled, in most cases leading to a surge in taxonomic similarity at the regional scale termed homogenization. The drivers of homogenization remain unclear, which may be a reflection of interactions between various environmental changes. In this study, we investigate two potential drivers of the recent changes in the composition of freshwater fish communities: recreational fishing and climate change. Our results, derived from 524 lakes of Ontario, Canada sampled in two periods (1965-1982 and 2008-2012), demonstrate that the main contributors to homogenization are the dispersal of gamefish species, most of which are large predators. Alternative explanations relating to lake habitat (e.g., area, phosphorus) or variations in climate have limited explanatory power. Our analysis suggests that human-assisted migration is the primary driver of the observed compositional shifts, homogenizing freshwater fish community among Ontario lakes and generating food webs dominated by gamefish species.

Effects of repeated daily acute heat challenge on the growth and metabolism of a cold water stenothermal fish.

Temperature is an important environmental factor influencing fish physiology that varies both spatially and temporally in ecosystems. In small north temperate zone lakes, cold water piscivores rely on nearshore prey; however, this region exceeds the optimal temperature of the foraging species during summer. To cope, piscivores make short excursions into the nearshore to feed and return to cold water to digest their meal, but the physiological impacts of these repeated acute exposures to warm water are not well understood. We exposed juvenile lake trout (Salvelinus namaycush) to treatments where they were held at ∼10°C and exposed to either 17 or 22°C for 5-10 min daily for 53 days mimicking warm-water forays. Control fish, held at an average temperature of ∼10°C but not exposed to thermal variation, consumed more food and grew slightly faster than heat challenged fish, with no clear differences in body condition, hepatosomatic index, ventricle mass, or muscle concentrations of lactate dehydrogenase and cytochrome c oxidase. Aerobic metabolic rates measured at 10°C indicated that standard metabolic rates (SMR) were similar among treatments; however, fish that were repeatedly exposed to 17°C had higher maximum metabolic rates (MMR) and aerobic scopes (AS) than control fish and those repeatedly exposed to 22°C. There were no differences in MMR or AS between fish exposed to 22°C and control fish. These results suggest that although SMR of fish are robust to repeated forays into warmer environments, MMR displays plasticity, allowing fish to be less constrained aerobically in cold water after briefly occupying warmer waters.

Seasonal variation in activity and nearshore habitat use of Lake Trout in a subarctic lake.

BACKGROUND: In lake ecosystems, predatory fish can move and forage across both nearshore and offshore habitats. This coupling of sub-habitats, which is important in stabilizing lake food webs, has largely been assessed from a dietary perspective and has not included movement data. As such, empirical estimates of the seasonal dynamics of these coupling movements by fish are rarely quantified, especially for northern lakes. Here we collect fine-scale fish movement data on Lake Trout (Salvelinus namaycush), a predatory cold-water fish known to link nearshore and offshore habitats, to test for seasonal drivers of activity, habitat use and diet in a subarctic lake. METHODS: We used an acoustic telemetry positioning array to track the depth and spatial movements of 43 Lake Trout in a subarctic lake over two years. From these data we estimated seasonal 50% home ranges, movements rates, tail beat activity, depth use, and nearshore habitat use. Additionally, we examined stomach contents to quantify seasonal diet. Data from water temperature and light loggers were used to monitor abiotic lake conditions and compare to telemetry data. RESULTS: Lake Trout showed repeatable seasonal patterns of nearshore habitat use that peaked each spring and fall, were lower throughout the long winter, and least in summer when this habitat was above preferred temperatures. Stomach content data showed that Lake Trout acquired the most nearshore prey during the brief spring season, followed by fall, and winter, supporting telemetry results. Activity rates were highest in spring when feeding on invertebrates and least in summer when foraging offshore, presumably on large-bodied prey fish. High rates of nearshore activity in fall were associated with spawning. Nearshore habitat use was widespread and not localized to specific regions of the lake, although there was high overlap of winter nearshore core areas between years. CONCLUSIONS: We provide empirical demonstrations of the seasonal extent to which a mobile top predator links nearshore and offshore habitats in a subarctic lake. Our findings suggest that the nearshore is an important foraging area for Lake Trout for much of the year, and the role of this zone for feeding should be considered in addition to its traditional importance as spawning habitat.

Species portfolio effects dominate seasonal zooplankton stabilization within a large temperate lake.

Portfolio effects (PEs) in ecology refer to the suite of phenomenon where the temporal variation of aggregate ecosystem properties (i.e., abundance) is lower than that of their ecosystem components. An example of this is where differential responses of species to environmental variation generate stability at higher levels of ecological organization (e.g., local community, metapopulation, metacommunity). Most of the research examining such PEs has focused on spatial or interannual variation of ecosystems; however, as global change continues to alter seasonality and ecosystem functioning, understanding the underlying food web structures that help maintain stability at multiple spatial and temporal scales is critical to managing ecological systems. Recent advances investigating diversity-stability relationships has led to the development of frameworks that incorporate a metacommunity perspective which allows for the partitioning of PEs across organizational scales (i.e., local community, metapopulation, cross-community, metacommunity) from local population dynamics (total). This partitioning yields insights into the mechanisms that generate observed PEs in nature. Here, we employed one of these recently developed frameworks on a temporally (1986-1999, 2008-2019) and spatially (five sampling stations, local communities) extensive data set of zooplankton abundance (e.g., density) within a large temperate lake to investigate how temporal (seasonal) and spatial (among site) PEs influence stability within the zooplankton metacommunity. We found that seasonal asynchrony of different zooplankton species within local communities and across communities generated the vast majority of stabilization, while spatial (i.e., metapopulation) dynamics were more synchronous and contributed little to overall system stability. Furthermore, significantly positive diversity-asynchrony relationships at the total, local- and cross-community scales were found as asynchrony was positively correlated with local Shannon diversity. Last, a comparison of PEs over the time periods, during which significant local and global changes (i.e., climate warming, invasive species) have occurred suggests that PEs may be eroding, as increasingly synchronous dynamics and declining diversity in recent years have led to a rise in metacommunity variability. We end by arguing for the critical importance of understanding seasonally driven stabilizing mechanisms as local and global changes threaten to fundamentally alter seasonal signals with potentially strong implications for the structures that lend stability to ecosystems.

Subsidy accessibility drives asymmetric food web responses.

Global change is fundamentally altering flows of natural and anthropogenic subsidies across space and time. After a pointed call for research on subsidies in the 1990s, an industry of empirical work has documented the ubiquitous role subsidies play in ecosystem structure, stability, and function. Here, we argue that physical constraints (e.g., water temperature) and species traits can govern a species' accessibility to resource subsidies, which has been largely overlooked in the subsidy literature. We examined the input of a high-quality, point-source anthropogenic subsidy (aquaculture feed) into a recipient freshwater lake food web. Using a combined bio-tracer approach, we detect a gradient in accessibility of the anthropogenic subsidy within the surrounding food web driven by the thermal preferences of three constituent species, effectively rewiring the recipient lake food web. Because aquaculture is predicted to increase significantly in coming decades to support growing human populations, and global change is altering temperature regimes, then this form of food web alteration may be expected to occur frequently. We argue that subsidy accessibility is a key characteristic of recipient food web interactions that must be considered when trying to understand the impacts of subsidies on ecosystem stability and function under continued global change.

Seasonal variation of behavior and brain size in a freshwater fish.

Teleost fishes occupy a range of ecosystem, and habitat types subject to large seasonal fluctuations. Temperate fishes, in particular, survive large seasonal shifts in temperature, light availability, and access to certain habitats. Mobile species such as lake trout (Salvelinus namaycush) can behaviorally respond to seasonal variation by shifting their habitat deeper and further offshore in response to warmer surface water temperatures during the summer. During cooler seasons, the use of more structurally complex nearshore zones by lake trout could increase cognitive demands and potentially result in a larger relative brain size during those periods. Yet, there is limited understanding of how such behavioral responses to a seasonally shifting environment might shape, or be shaped by, the nervous system.Here, we quantified variation in relative brain size and the size of five externally visible brain regions in lake trout, across six consecutive seasons in two different lakes. Acoustic telemetry data from one of our study lakes were collected during the study period from a different subset of individuals and used to infer relationships between brain size and seasonal behaviors (habitat use and movement rate).Our results indicated that lake trout relative brain size was larger in the fall and winter compared with the spring and summer in both lakes. Larger brains coincided with increased use of nearshore habitats and increased horizontal movement rates in the fall and winter based on acoustic telemetry. The telencephalon followed the same pattern as whole brain size, while the other brain regions (cerebellum, optic tectum, olfactory bulbs, and hypothalamus) were only smaller in the spring.These findings provide evidence that flexibility in brain size could underpin shifts in behavior, which could potentially subserve functions associated with differential habitat use during cold and warm seasons and allow fish to succeed in seasonally variable environments.

Food availability modulates temperature-dependent effects on growth, reproduction, and survival in Daphnia magna.

Reduced body size and accelerated life cycle due to warming are considered major ecological responses to climate change with fitness costs at the individual level. Surprisingly, we know little about how relevant ecological factors can alter these life history trade-offs and their consequences for individual fitness. Here, we show that food modulates temperature-dependent effects on body size in the water flea Daphnia magna and interacts with temperature to affect life history parameters. We exposed 412 individuals to a factorial manipulation of food abundance and temperature, tracked each reproductive event, and took daily measurements of body size from each individual. High temperature caused a reduction in maximum body size in both food treatments, but this effect was mediated by food abundance, such that low food conditions resulted in a reduction of 20% in maximum body size, compared with a reduction of 4% under high food conditions. High temperature resulted in an accelerated life cycle, with pronounced fitness cost at low levels of food where only a few individuals produced a clutch. These results suggest that the mechanisms affecting the trade-off between fast growth and final body size are food-dependent, and that the combination of low levels of food and high temperature could potentially threaten viability of ectotherms.

Food web rewiring in a changing world.

Climate change is asymmetrically altering environmental conditions in space, from local to global scales, creating novel heterogeneity. Here, we argue that this novel heterogeneity will drive mobile generalist consumer species to rapidly respond through their behaviour in ways that broadly and predictably reorganize - or rewire - food webs. We use existing theory and data from diverse ecosystems to show that the rapid behavioural responses of generalists to climate change rewire food webs in two distinct and critical ways. First, mobile generalist species are redistributing into systems where they were previously absent and foraging on new prey, resulting in topological rewiring - a change in the patterning of food webs due to the addition or loss of connections. Second, mobile generalist species, which navigate between habitats and ecosystems to forage, will shift their relative use of differentially altered habitats and ecosystems, causing interaction strength rewiring - changes that reroute energy and carbon flows through existing food web connections and alter the food web's interaction strengths. We then show that many species with shared traits can exhibit unified aggregate behavioural responses to climate change, which may allow us to understand the rewiring of whole food webs. We end by arguing that generalists' responses present a powerful and underutilized approach to understanding and predicting the consequences of climate change and may serve as much-needed early warning signals for monitoring the looming impacts of global climate change on entire ecosystems.

Field testing a novel high residence positioning system for monitoring the fine-scale movements of aquatic organisms.

Acoustic telemetry is an important tool for studying the behaviour of aquatic organisms in the wild.VEMCO high residence (HR) tags and receivers are a recent introduction in the field of acoustic telemetry and can be paired with existing algorithms (e.g. VEMCO positioning system [VPS]) to obtain high-resolution two-dimensional positioning data.Here, we present results of the first documented field test of a VPS composed of HR receivers (hereafter, HR-VPS). We performed a series of stationary and moving trials with HR tags (mean HR transmission period = 1.5 s) to evaluate the precision, accuracy and temporal capabilities of this positioning technology. In addition, we present a sample of data obtained for five European perch Perca fluviatilis implanted with HR tags (mean HR transmission period = 4 s) to illustrate how this technology can estimate the fine-scale behaviour of aquatic animals.Accuracy and precision estimates (median [5th-95th percentile]) of HR-VPS positions for all stationary trials were 5.6 m (4.2-10.8 m) and 0.1 m (0.02-0.07 m), respectively, and depended on the location of tags within the receiver array. In moving tests, tracks generated by HR-VPS closely mimicked those produced by a handheld GPS held over the tag, but these differed in location by an average of ≈9 m.We found that estimates of animal speed and distance travelled for perch declined when positional data for acoustically tagged perch were thinned to mimic longer transmission periods. These data also revealed a trade-off between capturing real nonlinear animal movements and the inclusion of positioning error.Our results suggested that HR-VPS can provide more representative estimates of movement metrics and offer an advancement for studying fine-scale movements of aquatic organisms, but high-precision survey techniques may be needed to test these systems.

Development and testing of a simple field-based intermittent-flow respirometry system for riverine fishes.

By understanding range-wide intraspecific variation in metabolic rate we can better understand how organisms have adapted to their environment. However, methods to quantify metabolic rate of fishes from remote areas or those that cannot be brought back to the laboratory because of imperilment status are lacking. Consequently, practical and reliable field-based methods are needed. To address this need, we developed a simple yet robust intermittent-flow respirometry system, adapted from a design commonly used in the laboratory that is readily suited for field use. Standard metabolic rate (SMR), maximum metabolic rate (MMR) and aerobic scope (AS) estimates were obtained from juvenile lake trout (Salvelinus namaycush) and brook trout (Salvelinus fontinalis) using both field- and laboratory-based systems. Whole-fish SMR, MMR and AS estimates from the field and laboratory methods did not differ from one another (ANCOVA and LMM: all P > 0.05) for either species and were comparable to estimates previously reported. Our field setup is a simpler system than the conventional laboratory-based system that requires less power and equipment to operate, yet still offers users the ability to: (1) acclimate fish to the respirometry chamber; (2) measure oxygen consumption during a shorter period (1 h), which yield metabolic rate estimates comparable to systems that take measurements over longer periods; and (3) take repeated oxygen consumption measurements with manual user-defined flush and measurement phase routines. Developing practical and reliable field respirometry methods, as demonstrated here, is important if we wish to improve our ability to predict how imperiled species will respond to changes in their environment. Such knowledge is critical for informing conservation strategies.

Persistent organic pollutant concentrations in fledglings of two arctic seabird species.

Persistent organic pollutants (POPs) and stable isotopes were measured in muscle from fledglings of two arctic seabird species, Northern fulmar (Fulmarus glacialis) and Black-legged kittiwakes (Rissa tridactyla). The purpose was to compare POP concentrations between species, in an age class that is highly vulnerable to POPs but little studied, relate to diet using stable isotopes, and quantify differences across life stages (egg to adult). Northern fulmar fledglings had significantly higher POP concentrations than kittiwake, consistent with results reported for adults of these species. Surprisingly, carbon and nitrogen stable isotopes did not differ between species, which does not match data for, or the known feeding ecology, of the adults. Fulmar/kittiwake POP concentration ratios varied across life stages indicating variable POP exposure and accumulation with age in seabirds, indicating that of the use of avian species-specific thresholds should only be done with caution in ecosystem-based POP risk management.