Introduced trout sever trophic connections in watersheds: consequences for a declining amphibian

Trophic linkages between terrestrial and aquatic ecosystems are increasingly recognized as important yet poorly known features of food webs. Here we describe research to understand the dynamics of lake food webs in relation to a native riparian amphibian and its interaction with introduced trout. The mountain yellow-legged frog Rana muscosa is endemic to alpine watersheds of the Sierra Nevada Mountains and the Transverse Ranges of California, but it has declined to a small fraction of its historical distribution and abundance. Although remaining frogs and introduced trout feed in different habitats of alpine lakes, our stable-isotope analyses clearly show that the same resource base of benthic invertebrates sustains their growth. During one period, insect emergence from naturally fishless lakes was nearly 20-fold higher compared to adjacent lakes with trout, showing that fish reduce availability of aquatic prey to amphibious and terrestrial consumers. Although trout cannot prey on adult frogs due to gape limitation, foraging post-metamorphic frogs are 10 times more abundant in the absence of trout, suggesting an important role for competition for prey by trout in highly unproductive alpine watersheds. Most Sierran lakes contain fish, and those that do not are usually small isolated ponds; in our study, these two lake types supported the lowest densities of post-metamorphic frogs, and these frogs were less reliant on local, benthic sources of productivity. Since Rana muscosa was formerly the most abundant vertebrate in the Sierra Nevada, the reduction in energy flow from lake benthos to this consumer due to fish introductions may have had negative consequences for its numerous terrestrial predators, many of which have also declined. We suggest that disruptions of trophic connections between aquatic and terrestrial food webs are an important but poorly understood consequence of fish introduction to many thousands of montane lakes and streams worldwide and may contribute to declines of native consumers in riparian habitats.     

Lake of flies, or lake of fish? A trophic model of Lake Malawi

Ecosystem-focused models have, for the first time, become available for the combined demersal and pelagic components of a large tropical lake ecosystem, Lake Malawi. These provide the opportunity to explore continuing controversies over the production efficiencies and ecological functioning of large tropical lakes. In Lake Malawi these models can provide important insight to the effect of fishing on fish composition, and the potential competition that the lakefly Chaoborus edulis may have with fisheries production. A mass-balanced trophic model developed for the demersal fish community of the southern and western areas of Lake Malawi was integrated with an existing trophic model developed for the open-water pelagic. Input parameters for the demersal model were obtained from a survey of fish distributions, fish food consumption studies, and from additional published quantitative and qualitative information on the various biotic components of the community. The model was constructed using the Ecopath approach and software. The graphically presented demersal food web spanned four trophic levels and was based primarily on consumption of detritus, zooplankton and sedimented diatoms. Zooplankton was imported into the system at trophic levels three and four through fish predation on carnivorous and herbivorous copepods and Chaoborus larvae. It is proposed that the primary consumption of copepods was by fish migrating into the pelagic zone. Chaoborus larvae in the demersal were probably consumed near the lakebed as they conducted a daily migration from the pelagic to seek refuge in the sediments. This evidence for strong benthic-pelagic coupling provided the opportunity for linking the demersal model to the existing model for the pelagic community so producing the first model for the complete ecosystem. Energy fluxes through the resulting combined model demonstrated that the primary import of biomass to the demersal system was detritus of pelagic origin (72.1%) and pelagic zooplankton (10.6%). Only 15.8% of the biomass consumed within the demersal system was of demersal origin. Lakefly production is efficiently utilised by the lake fish community, and any attempt to improve fishery production through introduction of a non-native plantivorous fish species would have a negative impact on the stability and productivity of the lake ecosystem.     

Role of the fish Astyanax aeneus (Characidae) as a keystone nutrient recycler in low-nutrient neotropical streams

Nutrient recycling by animals is a potentially important biogeochemical process in both terrestrial and aquatic ecosystems. Stoichiometric traits of individual species may result in some taxa playing disproportionately important roles in the recycling of nutrients relative to their biomass, acting as keystone nutrient recyclers. We examined factors controlling the relative contribution of 12 Neotropical fish species to nutrient recycling in four streams spanning a range of phosphorus (P) levels. In high-P conditions (135 microg/L soluble reactive phosphorus, SRP), most species fed on P-enriched diets and P excretion rates were high across species. In low-P conditions (3 microg/L SRP), aquatic food resources were depleted in P, and species with higher body P content showed low rates of P recycling. However, fishes that were subsidized by terrestrial inputs were decoupled from aquatic P availability and therefore excreted P at disproportionately high rates. One of these species, Astyanax aeneus (Characidae), represented 12% of the total population and 18% of the total biomass of the fish assemblage in our focal low-P study stream but had P excretion rates > 10-fold higher than other abundant fishes. As a result, we estimated that P excretion by A. aeneus accounted for 90% of the P recycled by this fish assemblage and also supplied approximately 90% of the stream P demand in this P-limited ecosystem. Nitrogen excretion rates showed little variation among species, and the contribution of a given species to ecosystem N recycling was largely dependent upon the total biomass of that species. Because of the high variability in P excretion rates among fish species, ecosystem-level P recycling could be particularly sensitive to changes in fish community structure in P-limited systems.     

Quantity and quality: unifying food web and ecosystem perspectives on the role of resource subsidies in freshwaters

Although the study of resource subsidies has emerged as a key topic in both ecosystem and food web ecology, the dialogue over their role has been limited by separate approaches that emphasize either subsidy quantity or quality. Considering quantity and quality together may provide a simple, but previously unexplored, framework for identifying the mechanisms that govern the importance of subsidies for recipient food webs and ecosystems. Using a literature review of > 90 studies of open-water metabolism in lakes and streams, we show that high-flux, low-quality subsidies can drive freshwater ecosystem dynamics. Because most of these ecosystems are net heterotrophic, allochthonous inputs must subsidize respiration. Second, using a literature review of subsidy quality and use, we demonstrate that animals select for high-quality food resources in proportions greater than would be predicted based on food quantity, and regardless of allochthonous or autochthonous origin. This finding suggests that low-flux, high-quality subsidies may be selected for by animals, and in turn may disproportionately affect food web and ecosystem processes (e.g., animal production, trophic energy or organic matter flow, trophic cascades). We then synthesize and review approaches that evaluate the role of subsidies and explicitly merge ecosystem and food web perspectives by placing food web measurements in the context of ecosystem budgets, by comparing trophic and ecosystem production and fluxes, and by constructing flow food webs. These tools can and should be used to address future questions about subsidies, such as the relative importance of subsidies to different trophic levels and how subsidies may maintain or disrupt ecosystem stability and food web interactions.     

Trophic relationships in the recently impounded Bagré reservoir in Burkina Faso

The trophic dynamics of Bagré reservoir which has been recently impounded in Burkina Faso was based on the data collected during 1997–1998 period using the Ecopath model and software. Total fish biomass is 22.63 t km⁻² and mainly represents trophic levels (TLs) 2 and 3. The trophic food chain is relatively long and the overall transfer efficiency is quite low. Grazing foodweb based on primary producers is prominent in the reservoir ecosystem and detritus plays a less significant role. Seasonal and long-term variations in water quality have significant influences on the lower TLs clearly showing a bottom-up functioning of the ecosystem. Environmental degradations, such as siltation occurring in the lake, suggest possible risks in limiting ecosystem productivity.     

Harvesting in an eight-species ecosystem

The theory for a general equilibrium ecosystem model that can include large number of interacting species is presented. Features include: (1) individual plants and animals are assumed to behave as if they are maximizing their net energy intake, (2) short- and long-run equilibriums are obtained, (3) species’ population adjustments depend on individual net energies. The theory is applied using simulations of an eight-species Alaskan marine ecosystem for which a “natural” equilibrium is calculated. Humans are introduced by adding a regulated open access fishery that harvests one of the species. Fishing impacts the fish population as well as the populations of other species, including Stellar sea lions, an endangered species. The sensitivity of fish and nonfish species populations to harvesting are calculated.     

Interacting trophic forcing and the population dynamics of herring

Small pelagic fish occupy a central position in marine ecosystems worldwide, largely by determining the energy transfer from lower trophic levels to predators at the top of the food web, including humans. Population dynamics of small pelagic fish may therefore be regulated neither strictly bottom-up nor top-down, but rather through multiple external and internal drivers. While in many studies single drivers have been identified, potential synergies of multiple factors, as well as their relative importance in regulating population dynamics of small pelagic fish, is a largely unresolved issue. Using a statistical, age-structured modeling approach, we demonstrate the relative importance and influence of bottom-up (e.g., climate, zooplankton availability) and top-down (i.e., fishing and predation) factors on the population dynamics of Bothnian Sea herring (Clupea harengus) throughout its life cycle. Our results indicate significant bottom-up effects of zooplankton and interspecific competition from sprat (Sprattus sprattus), particularly on younger age classes of herring. Although top-down forcing through fishing and predation by grey seals (Halichoerus grypus) and Atlantic cod (Gadus morhua) also was evident, these factors were less important than resource availability and interspecific competition. Understanding key ecological processes and interactions is fundamental to ecosystem-based management practices necessary to promote sustainable exploitation of small pelagic fish.     

How might we model an ecosystem?

Predicting ecosystem effects is of crucial importance in a world at threat from natural and human-mediated change. Here we propose an ecologically defensible representation of an ecosystem that facilitates predictive modelling. The representation has its roots in the early trophic and energetic theory of ecosystem dynamics and more recent functional ecology and network theory. Using the arable ecosystem of the UK as an example, we show that the representation allows simplification from the many interacting plant and invertebrate species, typically present in arable fields, to a more tractable number of trophic-functional types. Our compound hypothesis is that “trophic-functional types of plants and invertebrates can be used to explain the structure, diversity and dynamics of arable ecosystems”. The trophic-functional types act as containers for individuals, within an individual-based model, sharing similar trophic behaviour and traits of biomass transformation. Biomass, or energy, flows between the types and this allows the key ecological properties of individual abundance and body mass, at each trophic height, to be followed through simulations. Our preliminary simulation results suggest that the model shows great promise. The simulation output for simple ecosystems, populated with realistic parameter values, is consistent with current laboratory observations and provides exciting indications that it could reproduce field scale phenomena. The model also produces output that links the individual, population and community scales, and may be analysed and tested using community, network (food web) and population dynamic theory. We show that we can include management effects, as perturbations to parameter values, for modelling the effects of change and indicating management responses to change. This model will require robust analysis, testing and validation, and we discuss how we will achieve this in the future.     

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

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

Biotic control of stream fluxes: spawning salmon drive nutrient and matter export

Organisms can control movements of nutrients and matter by physically modifying habitat. We examined how an ecosystem engineer, sockeye salmon (Oncorhynchus nerka), influences seasonal fluxes of sediments, nitrogen (N), and phosphorus (P) in streams of southwestern Alaska. The purpose of this study was to investigate whether salmon act as net importers or net exporters of matter and nutrients from streams and how these roles change as a function of salmon population density. We measured discharge and concentrations of suspended sediments and total N and P every 7-14 days for up to four summers in 10 streams spanning a gradient in salmon densities. We statistically allocated whole-season fluxes to salmon activities, such as excretion and bioturbation, and to export by hydrologic discharge. In addition, we used counts of spawning salmon to estimate nutrient and matter imports by salmon to streams. Large seasonal pulses of suspended sediments, P, and N were associated with salmon spawning activities, often increasing export an order of magnitude higher than during pre-salmon levels. Years and streams with more salmon had significantly higher levels of export of sediments and nutrients. In addition, years with higher precipitation had higher background export of P and N. Salmon exported an average of the equivalent of 189%, 60%, and 55% of total matter, P, and N that salmon imported in their bodies. The relative magnitude of export varied; salmon exported more than their bodies imported in 80%, 20%, and 16% across all streams and years for sediments, P, and N, respectively. A bioassay experiment indicated that the P exported by salmon is directly available for use by primary producers in the downstream lake. These results demonstrate that salmon not only move nutrients upstream on large spatial scales via their migration from the ocean and subsequent death, but also redistribute matter and nutrients on finer spatial scales through their spawning activities.     

From single fine roots to a black alder forest ecosystem: How system behaviour emerges from single component activities

Based on empirical findings in a natural black alder ecosystem in Northern Germany we developed an individual based model that integrates components of a black alder ecosystem interacting on different levels of organisation. The factors determining seasonal fine root biomass development of forest ecosystems are not yet fully understood.We used an object oriented model approach to investigate this complex matter for black alder trees. Processes like growth, storage, respiration, transport, nutrient mineralisation and uptake as well as interactions among these factors are described on the level of functionally differentiated plant organs (fine roots, coarse roots, stem, branches, leaves) and soil units. The object structure of the model is determined by spatial relations between plant modules as well as between plant modules and their local environment modules.As results of model application we found that (i) on the organ level, spatio-temporal plasticity of (root) growth allocation is related to spatio-temporal variation of resource availability, (ii) on the plant level, balanced root:shoot growth appears in response to variation of available resources light and nutrients, (iii) on the population level, tree stand development (population structure, self-thinning) resulted from coexistence and competition between plant individuals.For the understanding of the root compartment it seems relevant that the model implementation of local scale fine root dynamics is consistent with a self-organised large scale spatial heterogeneity of fine root activity pattern. On the other hand, fine-root dynamics cannot be explained as a result of autonomous dynamics. A reference to above-ground processes is a necessary condition and the overall plant seems to act as an integrator providing boundary conditions for local activity pattern. At the same time fine-root characteristics are of some importance for properties on hierarchically higher levels, e.g. co-existence in a tree population or element cycling in the ecosystem.As a conclusion, modelling of the spatio-temporal dynamics of tree root systems appears as a paradigmatic example of scale and organisation level integrating processes.     

Underappreciated species in ecology: "ugly fish" in the northwest Atlantic Ocean.

Species shifts and replacements are common in ecological studies. Observations thereof serve as the impetus for many ecological endeavors. Many of the species now known to dominate ecosystem functioning were largely ignored until studies of those underappreciated species elucidated their critical roles. Recognizing the potential importance of underappreciated species has implications for functional redundancies in ecosystems and should alter our approach to long-term monitoring. One example of an applied ecological system containing species shifts, underappreciated species, and potential changes in functional redundancies is the topic of fisheries. The demersal component of many fish communities usually consists of high-profile and commercially valuable species that are targets of fisheries, plus a diverse group of lesser known species that have minimal commercial value and focus. Yet ecologically these traditionally nontargeted species are often a major biomass sink in marine ecosystems and can also be critical in the functioning of bentho-demersal food webs. I examined the biomass trajectories of several species of skates, cottids, lophiids, anarhichadids, zooarcids, and similar species in the northeast U.S. Atlantic ecosystem to determine whether their relative abundance has changed across the past four decades. Distribution and stomach contents of these species were also evaluated over time to further elucidate the relative importance of these species. Landings of these underappreciated bentho-demersal fish were also examined in comparison to those species that historically have been commercially targeted. Of particular emphasis was the evaluation of evidence for sequential stock depletion and the ramifications for functional redundancy for this ecosystem. Results indicate that some of these fish species are now the dominant piscivores, benthivores, and scavengers in this ecosystem. These formerly under-studied species generally have either maintained a consistent population size or have increased in abundance (and expanded in distribution) over the past several decades. Nontraditionally targeted fish species are an often overlooked but important component of bentho-demersal fish communities. Implications for the energy flow and resilience specifically for future fisheries and generally for harvesting biological resources are significant, remaining critical issues for the world's ecosystems.     

Elucidation of ecosystem attributes of an oligotrophic lake in Hokkaido,Japan, using Ecopath with Ecosim (EwE)

The fishing practices in the oligotrophic Lake Toya, Hokkaido, Japan, have profound implications in the ecosystem sustainability. The status of the sockeye salmon (Oncorhynchus nerka) population has become a serious concern among the lake managers and policy makers during the last decades. While the decline of the sockeye salmon population has been well documented in Lake Toya, there is considerable uncertainty with regards to the impact on the broader system dynamics. In this study, our objective is to address this knowledge gap by undertaking a synthesis of the Lake Toya food web using the mass-balance modeling software Ecopath with Ecosim (EwE). Our primary research question is to examine the repercussions of the declining sockeye salmon population on the trophic dynamics of the lake. Namely, we assess if there are any competing species that might have benefited from the decrease of sockeye salmon standing biomass and to what extent do these changes propagate through the Lake Toya food web? Our analysis pinpoints the critical role of the Japanese smelt (Hypomesus transpacificus nipponensis) in the system, which demonstrates a wide range of effects on several functional groups at both higher and lower trophic levels in Lake Toya. In particular, being a substantial portion of the masu salmon (Oncorhynchus masou) and adult sockeye salmon diets, the Japanese smelt has a positive impact on the top predators of the system. Amphipods, insects, and shrimp strongly benefit from the autochthonous and allochthonous organic matter in the system, while the tight coupling between phytoplankton and zooplankton seems to be particularly critical for the integrity of the Lake Toya food web. Whereas the values of the different ecosystem attributes (e.g., primary production/biomass, biomass/total throughput, system omnivory index, amount of recycled throughput, Finn's cycling index) provide evidence that Lake Toya is an immature system, we note that the internal redundancy and the system overhead estimates suggest that the lake possesses substantial reserves to overcome external perturbations. We also examined the effects of a variety of fishing policies on the biomass of masu salmon and adult sockeye salmon, which verify the belief that the adult sockeye population is quite fragile with high likelihood to collapse. Our analysis also predicts that sockeye will not rebound unless the fishing pressure exerted is substantially reduced (>50% of the reference levels used). Masu salmon seems to benefit under all the scenarios examined indicating that the intensity of the current fishing activities is significantly lower than its biomass accumulation rate in the system.     

Modelling coral reef ecosystems with limited observational data

Ecosystem models represent potentially powerful tools for coral reef ecosystem managers. They can provide insight into ecosystem dynamics not achievable through alternative means allowing coral reef managers to assess the potential outcome of any given management decision. One of the main limitations in the applicability of ecosystem models is that they often require detailed empirical data and this can restrict their applicability to ecosystems that are either currently well studied or have the resources available to collect the required data. This study describes the development of a coral reef ecosystem model that can be calibrated to an ecosystem with limited empirical data. Based on the assumption that coral reef ecological structure is generic across all tropical coral reefs and that the magnitude of the interactions between ecological components is reef specific, the dynamics of the ecosystem can be replicated based on limited empirical data. The model successfully replicated the dynamics of three individual reef systems including an inshore and oceanic reef within the Great Barrier Reef and a Caribbean reef system. It highlighted the importance of understanding the specific dynamics of a given reef and that a positive management intervention in one system may result in a negative outcome for another. The model was also used to assess the importance of various interactions within coral reef ecosystems. It identified the interactions between hard corals and other non-algal benthic components as being an important (but currently understudied) facet of coral reef ecology. The development of this modelling approach provides access to ecosystem modelling tools for coral reef managers previously excluded due to a lack of resources or technical expertise.     

Marine-derived nutrients, bioturbation, and ecosystem metabolism: reconsidering the role of salmon in streams

In coastal areas of the North Pacific Ocean, annual returns of spawning salmon provide a substantial influx of nutrients and organic matter to streams and are generally believed to enhance the productivity of recipient ecosystems. Loss of this subsidy from areas with diminished salmon runs has been hypothesized to limit ecosystem productivity in juvenile salmon rearing habitats (lakes and streams), thereby reinforcing population declines. Using five to seven years of data from an Alaskan stream supporting moderate salmon densities, we show that salmon predictably increased stream water nutrient concentrations, which were on average 190% (nitrogen) and 390% (phosphorus) pre-salmon values, and that primary producers incorporated some of these nutrients into tissues. However, benthic algal biomass declined by an order of magnitude despite increased nutrients. We also measured changes in stream ecosystem metabolic properties, including gross primary productivity (GPP) and ecosystem respiration (ER), from three salmon streams by analyzing diel measurements of oxygen concentrations and stable isotopic ratios (delta O-O2) within a Bayesian statistical model of oxygen dynamics. Our results do not support a shift toward higher primary productivity with the return of salmon, as is expected from a nutrient fertilization mechanism. Rather, net ecosystem metabolism switched from approximately net autotrophic (GPP > or = ER) to a strongly net heterotrophic state (GPP < ER) in response to bioturbation of benthic habitats by salmon. Following the seasonal arrival of salmon, GPP declined to <12% of pre-salmon rates, while ER increased by over threefold. Metabolism by live salmon could not account for the observed increase in ER early in the salmon run, suggesting salmon nutrients and disturbance enhanced in situ heterotrophic respiration. Salmon also changed the physical properties of the stream, increasing air-water gas exchange by nearly 10-fold during peak spawning. We suggest that management efforts to restore salmon ecosystems should consider effects on ecosystem metabolic properties and how salmon disturbance affects the incorporation of marine-derived nutrients into food webs.     

An integrated modeling framework for performing environmental assessments: Application to ecosystem services in the Albemarle-Pamlico basins (NC and VA, USA)

The U.S. Environmental Protection Agency uses environmental models to inform rulemaking and policy decisions at multiple spatial and temporal scales. As decision-making has moved towards integrated thinking and assessment (e.g. media, site, region, services), the increasing complexity and interdisciplinary nature of modern environmental problems has necessitated a new generation of integrated modeling technologies. Environmental modelers are now faced with the challenge of determining how data from manifold sources, types of process-based and empirical models, and hardware/software computing infrastructure can be reliably integrated and applied to protect human health and the environment.In this study, we demonstrate an Integrated Modeling Framework that allows us to predict the state of freshwater ecosystem services within and across the Albemarle-Pamlico Watershed, North Carolina and Virginia (USA). The Framework consists of three facilitating technologies: Data for Environmental Modeling automates the collection and standardization of input data; the Framework for Risk Assessment of Multimedia Environmental Systems manages the flow of information between linked models; and the Supercomputer for Model Uncertainty and Sensitivity Evaluation is a hardware and software parallel-computing interface with pre/post-processing analysis tools, including parameter estimation, uncertainty and sensitivity analysis. In this application, five environmental models are linked within the Framework to provide multimedia simulation capabilities: the Soil Water Assessment Tool predicts watershed runoff; the Watershed Mercury Model simulates mercury runoff and loading to streams; the Water quality Analysis and Simulation Program predicts water quality within the stream channel; the Habitat Suitability Index model predicts physicochemical habitat quality for individual fish species; and the Bioaccumulation and Aquatic System Simulator predicts fish growth and production, as well as exposure and bioaccumulation of toxic substances (e.g., mercury).Using this Framework, we present a baseline assessment of two freshwater ecosystem services-water quality and fisheries resources-in headwater streams throughout the Albemarle-Pamlico. A stratified random sample of 50 headwater streams is used to draw inferences about the target population of headwater streams across the region. Input data is developed for a twenty-year baseline simulation in each sampled stream using current land use and climate conditions. Monte Carlo sampling (n = 100 iterations per stream) is also used to demonstrate some of the Framework's experimental design and data analysis features. To evaluate model performance and accuracy, we compare initial (i.e., uncalibrated) model predictions (water temperature, dissolved oxygen, fish density, and methylmercury concentration within fish tissue) against empirical field data. Finally, we ‘roll-up’ the results from individual streams, to assess freshwater ecosystem services at the regional scale.     

Structural changes in lake functioning induced from nutrient loading and climate variability

Climate variability is increasingly recognized as an important regulatory factor, capable of influencing the structural properties of aquatic ecosystems. Lakes appear to be particularly sensitive to the ecological impacts of climate variability, and several long time series have shown a close coupling between climate, lake thermal properties and individual organism physiology, population abundance, community structure, and food web dynamics. Thus, understanding the complex interplay among meteorological forcing, hydrological variability, and ecosystem functioning is essential for improving the credibility of model-based water resources/fisheries management. Our objective herein is to examine the relative importance of the ecological mechanisms underlying plankton seasonal variability in Lake Washington, Washington State (USA), over a 35-year period (1964–1998). Our analysis is founded upon an intermediate complexity plankton model that is used to reproduce the limiting nutrient (phosphate)–phytoplankton–zooplankton–detritus (particulate phosphorus) dynamics in the lake. Model parameterization is based on a Bayesian calibration scheme that offers insights into the degree of information the data contain about model inputs and allows obtaining predictions along with uncertainty bounds for modeled output variables. The model accurately reproduces the key seasonal planktonic patterns in Lake Washington and provides realistic estimates of predictive uncertainty for water quality variables of environmental management interest. A principal component analysis of the annual estimates of the underlying ecological processes highlighted the significant role of the phosphorus recycling stemming from the zooplankton excretion on the planktonic food web variability. We also identified a moderately significant signature of the local climatic conditions (air temperature) on phytoplankton growth (r = 0.41), herbivorous grazing (r = 0.38), and detritus mineralization (r = 0.39). Our study seeks linkages with the conceptual food web model proposed by Hampton et al. [Hampton, S.E., Scheuerell, M.D., Schindler, D.E., 2006b. Coalescence in the Lake Washington story: interaction strengths in a planktonic food web. Limnol. Oceanogr. 51, 2042–2051.] to emphasize the “bottom-up” control of the Lake Washington plankton phenology. The posterior predictive distributions of the plankton model are also used to assess the exceedance frequency and confidence of compliance with total phosphorus (15 μg L⁻¹) and chlorophyll a (4 μg L⁻¹) threshold levels during the summer-stratified period in Lake Washington. Finally, we conclude by underscoring the importance of explicitly acknowledging the uncertainty in ecological forecasts to the management of freshwater ecosystems under a changing global environment.     

Mating system and individual reproductive success of sympatric anadromous and resident brook charr, <Emphasis Type="Italic">Salvelinus fontinalis</Emphasis>, under natural conditions

Salmonids are known for the occurrence in sympatry of two life-history forms, one that undergoes migration to sea before returning to freshwater to reproduce (anadromous) and one that inhabits freshwater without a migration phase (resident). Whereas one breeding population is often suggested by population genetic studies, mating patterns have rarely been directly assessed, especially when both sexes are found within each life-history form. By using highly polymorphic microsatellite loci and parentage analysis in a natural population of sympatric anadromous and resident brook charr (Salvelinus fontinalis), we found that gene flow occurred between the two forms and was mediated by resident males mating with both resident and anadromous females. Determinants of reproductive success, estimated by the number of surviving juveniles (ages 1 and 2 years), differed between the sexes. No strong evidence of the influence of size on individual reproductive success was found for males, whereas larger females (and hence most likely to be anadromous) were more successful. The higher individual reproductive success of anadromous fish compared to residents was mainly explained by this higher reproductive success of anadromous females. We suggest that resident males adopt a “sneaking” reproductive tactic as a way of increasing their reproductive success by mating with females of all sizes in all habitats. The persistence of the resident tactic among females may be linked to their advantage in accessing spatially constrained spawning areas in small tributary streams unavailable to larger females.     

The importance of an infrequently flooded intertidal marsh surface as an energy source for the mummichog Fundulus heteroclitus: An experimental approach

Fundulus heteroclitus is known to ascend onto the marsh surface to feed. Our study investigated whether the marsh surface food items are a necessary source of caloric intake for the Canary Creek, Delaware, USA population of this species. Enclosure techniques were used to restrict mummichogs from the marsh surface and the growth rates of these fish were compared to those having access to the marsh surface. Growth rates were significantly higher for mummichogs allowed access to the marsh surface. Food addition and density reduction experiments showed that food availability per fish, rather than behavioral responses due to fish crowding, was responsible for the increased growth. Although food was available in the subtidal portion of the habitat, it was of insufficient quantity for fish at natural density to grow at a normal rate, and mummichogs must utilize the marsh surface for at least a portion of their energy intake.     

Effect of gyrodactylid ectoparasites on host behaviour and social network structure in guppies <Emphasis Type="Italic">Poecilia reticulata</Emphasis>

Understanding how individuals modify their social interactions in response to infectious disease is of central importance for our comprehension of how disease dynamics operate in real-world populations. Whilst a significant amount of theoretical work has modelled disease transmission using network models, we have comparatively little understanding of how infectious disease impacts on the social behaviour of individuals and how these effects scale up to the level of the population. We experimentally manipulated the parasite load of female guppies (Poecilia reticulata) and introduced fish either infected with the ectoparasites Gyrodactylus spp. (experimental) or uninfected (control) into replicated semi-natural populations of eight size-matched female guppies. We quantified the behaviour and social associations of both the introduced fish and the population fish. We found that infected experimental fish spent less time associating with the population fish than the uninfected control fish. Using information on which fish initiated shoal fission (splitting) events, our results demonstrate that the population fish actively avoided infected experimental fish. We also found that the presence of an infected individual resulted in a continued decline in social network clustering up to at least 24 h after the introduction of the infected fish, whereas in the control treatment, the clustering coefficient showed an increase at this time point. These results demonstrate that the presence of a disease has implications for both the social associations of infected individuals and for the social network structure of the population, which we predict will have consequences for infectious disease transmission.