Species invasions shift microbial phenology in a two-decade freshwater time series.

Invasive species impart abrupt changes on ecosystems, but their impacts on microbial communities are often overlooked. We paired a 20 y freshwater microbial community time series with zooplankton and phytoplankton counts, rich environmental data, and a 6 y cyanotoxin time series. We observed strong microbial phenological patterns that were disrupted by the invasions of spiny water flea (Bythotrephes cederströmii) and zebra mussels (Dreissena polymorpha). First, we detected shifts in Cyanobacteria phenology. After the spiny water flea invasion, Cyanobacteria dominance crept earlier into clearwater; and after the zebra mussel invasion, Cyanobacteria abundance crept even earlier into the diatom-dominated spring. During summer, the spiny water flea invasion sparked a cascade of shifting diversity where zooplankton diversity decreased and Cyanobacteria diversity increased. Second, we detected shifts in cyanotoxin phenology. After the zebra mussel invasion, microcystin increased in early summer and the duration of toxin production increased by over a month. Third, we observed shifts in heterotrophic bacteria phenology. The Bacteroidota phylum and members of the acI Nanopelagicales lineage were differentially more abundant. The proportion of the bacterial community that changed differed by season; spring and clearwater communities changed most following the spiny water flea invasion that lessened clearwater intensity, while summer communities changed least following the zebra mussel invasion despite the shifts in Cyanobacteria diversity and toxicity. A modeling framework identified the invasions as primary drivers of the observed phenological changes. These long-term invasion-mediated shifts in microbial phenology demonstrate the interconnectedness of microbes with the broader food web and their susceptibility to long-term environmental change.

Nine Lessons about Aquatic Invasive Species from the North Temperate Lakes Long-Term Ecological Research (NTL-LTER) Program.

Freshwater ecosystems can serve as model systems that reveal insights into biological invasions. In this article, we summarize nine lessons about aquatic invasive species from the North Temperate Lakes Long-Term Ecological Research program and affiliated projects. The lessons about aquatic invasive species are as follows: Invasive species are more widespread than has been documented; they are usually at low abundance; they can irrupt from low-density populations in response to environmental triggers; they can occasionally have enormous and far-reaching impacts; they can affect microbial communities; reservoirs act as invasive species hotspots; ecosystem vulnerability to invasion can be estimated; invasive species removal can produce long-term benefits; and the impacts of invasive species control may be greater than the impacts of the invasive species. This synthesis highlights how long-term research on a freshwater landscape can advance our understanding of invasions.

Lakeshore residential development as a driver of aquatic habitat and littoral fish communities: A cross-system study.

Lakeshore riparian habitats have undergone intensive residential development in many parts of the world. Lakeshore residential development (LRD) is associated with aquatic habitat loss/alteration, including altered macrophyte communities and reduced coarse woody habitat. Yet habitat-mediated and other generalized effects of LRD on lake biotic communities are not well understood. We used two approaches to examine the relationships among LRD, habitat, and fish community in a set of 57 northern Wisconsin lakes. First, we examined how LRD affected aquatic habitat using mixed linear effects models. Second, we evaluated how LRD affected fish abundance and community structure at both whole-lake and site-level spatial scales using generalized linear mixed-effects models. We found that LRD did not have a significant relationship with the total abundance (all species combined) of fish at either scale. However, there were significant species-specific responses to LRD at the whole-lake scale. Species abundances varied across the LRD gradient, with bluegill (Lepomis macrochirus) and mimic shiners (Notropis volucellus) responding positively along the gradient and walleye (Sander vitreus) having the most negative response. We also quantified site-level habitat associations for each fish species. We found that habitat associations did not inform a species' overall response to LRD, as illustrated by species with similar responses to LRD having vastly different habitat associations. Finally, even with the inclusion of littoral habitat information in models, LRD still had significant effects on species abundances, reflecting a role of LRD in shaping littoral fish communities independent of our measure of littoral habitat alteration. Our results indicated that LRD altered littoral fish communities at the whole-lake scale through both habitat and non-habitat-mediated drivers.

Production dynamics reveal hidden overharvest of inland recreational fisheries.

Recreational fisheries are valued at $190B globally and constitute the predominant way in which people use wild fish stocks in developed countries, with inland systems contributing the main fraction of recreational fisheries. Although inland recreational fisheries are thought to be highly resilient and self-regulating, the rapid pace of environmental change is increasing the vulnerability of these fisheries to overharvest and collapse. Here we directly evaluate angler harvest relative to the biomass production of individual stocks for a major inland recreational fishery. Using an extensive 28-y dataset of the walleye (Sander vitreus) fisheries in northern Wisconsin, United States, we compare empirical biomass harvest (Y) and calculated production (P) and biomass (B) for 390 lake year combinations. Production overharvest occurs when harvest exceeds production in that year. Biomass and biomass turnover (P/B) declined by ∼30 and ∼20%, respectively, over time, while biomass harvest did not change, causing overharvest to increase. Our analysis revealed that ∼40% of populations were production-overharvested, a rate >10× higher than estimates based on population thresholds often used by fisheries managers. Our study highlights the need to adapt harvest to changes in production due to environmental change.

Invasive species triggers a massive loss of ecosystem services through a trophic cascade.

Despite growing recognition of the importance of ecosystem services and the economic and ecological harm caused by invasive species, linkages between invasions, changes in ecosystem functioning, and in turn, provisioning of ecosystem services remain poorly documented and poorly understood. We evaluate the economic impacts of an invasion that cascaded through a food web to cause substantial declines in water clarity, a valued ecosystem service. The predatory zooplankton, the spiny water flea (Bythotrephes longimanus), invaded the Laurentian Great Lakes in the 1980s and has subsequently undergone secondary spread to inland lakes, including Lake Mendota (Wisconsin), in 2009. In Lake Mendota, Bythotrephes has reached unparalleled densities compared with in other lakes, decreasing biomass of the grazer Daphnia pulicaria and causing a decline in water clarity of nearly 1 m. Time series modeling revealed that the loss in water clarity, valued at US$140 million (US$640 per household), could be reversed by a 71% reduction in phosphorus loading. A phosphorus reduction of this magnitude is estimated to cost between US$86.5 million and US$163 million (US$430-US$810 per household). Estimates of the economic effects of Great Lakes invasive species may increase considerably if cases of secondary invasions into inland lakes, such as Lake Mendota, are included. Furthermore, such extreme cases of economic damages call for increased investment in the prevention and control of invasive species to better maximize the economic benefits of such programs. Our results highlight the need to more fully incorporate ecosystem services into our analysis of invasive species impacts, management, and public policy.

The consistency of a species' response to press perturbations with high food web uncertainty.

Predicting species responses to perturbations is a fundamental challenge in ecology. Decision makers must often identify management perturbations that are the most likely to deliver a desirable management outcome despite incomplete information on the pattern and strength of food web links. Motivated by a current fishery decline in inland lakes of the Midwestern United States, we evaluate consistency of the responses of a target species (walleye [Sander vitreus]) to press perturbations. We represented food web uncertainty with 193 plausible topological models and applied four perturbations to each one. Frequently the direction of the focal predator response to the same perturbation is not consistent across food web topologies. Simultaneous application of management perturbations led to less consistent outcomes compared to the best single perturbation. However, direct manipulation of the adult focal predator produced a desirable outcome in 77% of 193 plausible topologies. Identifying perturbations that produce consistent outcomes in the face of food web uncertainty can have important implications for natural resource conservation and management efforts.

Quantifying aquatic insect deposition from lake to land.

Adjacent ecosystems are influenced by organisms that move across boundaries, such as insects with aquatic larval stages and terrestrial adult stages, which transport energy and nutrients from water to land. However, the ecosystem-level effect of aquatic insects on land has generally been ignored, perhaps because the organisms themselves are individually small. At the naturally productive Lake Mývatn, Iceland, we used two readily measured quantities: total insect emergence from water and relative insect density on land, to demonstrate an approach for estimating aquatic insect deposition (e.g., kg N x m(-2) x yr(-1)) to shore. Estimates from emergence traps between 2008 and 20.11 indicated a range of 0.15-3.7 g x m(-2) x yr(-1), or a whole-lake emergence of 3.1-76 Mg/yr; all masses are given as dry mass. Using aerial infall trap measurements of midge relative abundance over land, we developed a local-maximum decay function model to predict proportional midge deposition with distance from the lake. The dispersal model predicted midge abundance with R2 = 0.89, a pattern consistent among years, with peak midge deposition occurring 20-25 m inland and 70% of midges deposited within 100 m of shore. During a high-midge year (2008), we estimate midge deposition within the first 50 m of shoreline to be 100 kg xha(-1) x yr(-1), corresponding to inputs of 10 kg N x ha(-1) x yr(-1) and 1 kg P x ha(-1) x yr(-1), or about three to five times above background terrestrial N deposition rates. Consistent with elevated N input where midges are most dense, we observed that soil available nitrate in resin bags decreases with increasing distance from the lake. Our approach, generalizable to other systems, shows that aquatic insects can be a major source of nutrients to terrestrial ecosystems and have the capacity to significantly affect ecosystem processes.

Taking the trophic bypass: aquatic-terrestrial linkage reduces methylmercury in a terrestrial food web.

Ecosystems can be linked by the movement of matter and nutrients across habitat boundaries via aquatic insect emergence. Aquatic organisms tend to have higher concentrations of certain toxic contaminants such as methylmercury (MeHg) compared to their terrestrial counterparts. If aquatic organisms come to land, terrestrial organisms that consume them are expected to have elevated MeHg concentrations. But emergent aquatic insects could have other impacts as well, such as altering consumer trophic position or increasing ecosystem productivity as a result of nutrient inputs from insect carcasses. We measure MeHg in terrestrial arthropods at two lakes in northeastern Iceland and use carbon and nitrogen stable isotopes to quantify aquatic reliance and trophic position. Across all terrestrial focal arthropod taxa (Lycosidae, Linyphiidae, Acari, Opiliones), aquatic reliance had significant direct and indirect (via changes in trophic position) effects on terrestrial consumer MeHg. However, contrary to our expectations, terrestrial consumers that consumed aquatic prey had lower MeHg concentrations than consumers that ate mostly terrestrial prey. We hypothesize that this is due to the lower trophic position of consumers feeding directly on midges relative to those that fed mostly on terrestrial prey and that had, on average, higher trophic positions. Thus, direct consumption of aquatic inputs results in a trophic bypass that creates a shorter terrestrial food web and reduced biomagnification of MeHg across the food web. Our finding that MeHg was lower at terrestrial sites with aquatic inputs runs counter to the conventional wisdom that aquatic systems are a source of MeHg contamination to surrounding terrestrial ecosystems.

Experimental evidence that ecological effects of an invasive fish are reduced at high densities.

Understanding the relationship between invasive species density and ecological impact is a pressing topic in ecology, with implications for environmental management and policy. Although it is widely assumed that invasive species impact will increase with density, theory suggests interspecific competition may diminish at high densities due to increased intraspecific interactions. To test this theory, we experimentally examined intra- and interspecific interactions between a globally invasive fish, round goby (Neogobius melanostomus), and three native species at different round goby densities in a tributary of the Laurentian Great Lakes. Eighteen 2.25 m(2) enclosures were stocked with native fish species at natural abundances, while round gobies were stocked at three different densities: 0 m(-2), 2.7 m(-2), and 10.7 m(-2). After 52 days, native fish growth rate was significantly reduced in the low density goby treatment, while growth in the high density goby treatment mirrored the goby-free treatment for two of three native species. Invertebrate density and gut content weight of native fishes did not differ among treatments. Conversely, gut content weight and growth of round gobies were lower in the high goby density treatment, suggesting interactions between round gobies and native fishes are mediated by interference competition amongst gobies. Our experiment provides evidence that invasive species effects may diminish at high densities, possibly due to increased intraspecific interactions. This is consistent with some ecological theory, and cautions against the assumption that invasive species at moderate densities have low impact.

Are rapid transitions between invasive and native species caused by alternative stable states, and does it matter?

Rapid transitions in ecosystem structure, or regime shifts, are a hallmark of alternative stable states (ASS). However, regime shifts can occur even when feedbacks are not strong enough to cause ASS. We investigated the potential for ASS to explain transitions between dominance of an invasive species, rusty crayfish (Orconectes rusticus), and native sunfishes (Lepomis spp.) in northern Wisconsin (USA) lakes. A rapid transition from Lepomis to rusty crayfish dominance occurred as rusty crayfish invaded Trout Lake, and the reverse transition resulted from an eight-year experimental removal of rusty crayfish from Sparkling Lake. We fit a stage-structured population model of species interactions to 31 years of time-series data from each lake. The model identified water level as an important driver, with drought conditions reducing rusty crayfish recruitment and allowing Lepomis dominance. The maximum-likelihood parameter estimates of the negative interaction between rusty crayfish and Lepomis led to ASS in the model, where each species was capable of excluding the other within a narrow range of environmental conditions. However, uncertainty in parameter estimates made it impossible to exclude the potential that rapid transitions were caused by a simpler threshold response lacking alternative equilibria. Simulated forward and backward transitions between species dominance occurred at different environmental conditions (i.e., hysteresis), even when the parameters used for simulation did not predict ASS as a result of slow species responses to environmental drivers. Thus, ASS are possible, but by no means certain, explanations for rapid transitions in this system, and our results highlight the difficulties associated with distinguishing ASS from other types of threshold responses. However, whether regime shifts are caused by ASS may be relatively unimportant in this system, as the range of conditions over which transitions occur is narrow, and under most conditions, the system is predicted to exist in only a single state.

Subsidies to predators, apparent competition and the phylogenetic structure of prey communities.

Ecosystems are fragmented by natural and anthropogenic processes that affect organism movement and ecosystem dynamics. When a fragmentation restricts predator but not prey movement, then the prey produced on one side of an ecosystem edge can subsidize predators on the other side. When prey flux is high, predator density on the receiving side increases above that possible by in situ prey productivity, and when low, the formerly subsidized predators can impose strong top-down control of in situ prey--in situ prey experience apparent competition from the subsidy. If predators feed on some evolutionary clades of in situ prey over others, then subsidy-derived apparent competition will induce phylogenetic structure in prey composition. Dams fragment the serial nature of river ecosystems by prohibiting movement of organisms and restricting flowing water. In the river tailwater just below a large central Mexican dam, fish density was high and fish gorged on reservoir-derived zooplankton. When the dam was closed, water flow and the zooplankton subsidy ceased, densely packed pools of fish formed, fish switched to feed on in situ prey, and the tailwater macroinvertebrate community was phylogenetic structured. We derived expectations of structure from trait-based community assembly models based on macroinvertebrate body size, tolerance to anthropogenic disturbance, and fish-diet selectivity. The diet-selectivity model best fit the observed tailwater phylogenetic structure. Thus, apparent competition from subsidies phylogenetically structures prey communities, and serial variation in phylogenetic community structure can be indicative of fragmentation in formerly continuous ecosystems.

Assessing ecosystem vulnerability to invasive rusty crayfish (Orconectes rusticus).

Despite the widespread introduction of nonnative species and the heterogeneity of ecosystems in their sensitivity to ecological impacts, few studies have assessed ecosystem vulnerability to the entire invasion process, from arrival to establishment and impacts. Our study addresses this challenge by presenting a probabilistic, spatially explicit approach to predicting ecosystem vulnerability to species invasions. Using the freshwater-rich landscapes of Wisconsin, USA, we model invasive rusty crayfish (Orconectes rusticus) as a function of exposure risk (i.e., likelihood of introduction and establishment of O. rusticus based on a species distribution model) and the sensitivity of the recipient community (i.e., likelihood of impacts on native O. virilis and O. propinquus based on a retrospective analysis of population changes). Artificial neural networks predicted that approximately 10% of 4200 surveyed lakes (n = 388) and approximately 25% of mapped streams (23 523 km total length) are suitable for O. rusticus introduction and establishment. A comparison of repeated surveys before vs. post-1985 revealed that O. virilis was six times as likely and O. propinquus was twice as likely to be extirpated in streams invaded by O. rusticus, compared to streams that were not invaded. Similarly, O. virilis was extirpated in over three-quarters of lakes invaded by O. rusticus compared to half of the uninvaded lakes, whereas no difference was observed for O. propinquus. We identified 115 lakes (approximately 3% of lakes) and approximately 5000 km of streams (approximately 6% of streams) with a 25% chance of introduction, establishment, and extirpation by O. rusticus of either native congener. By identifying highly vulnerable ecosystems, our study offers an effective strategy for prioritizing on-the-ground management action and informing decisions about the most efficient allocation of resources. Moreover, our results provide the flexibility for stakeholders to identify priority sites for prevention efforts given a maximum level of acceptable risk or based on budgetary or time restrictions. To this end, we incorporate the model predictions into a new online mapping tool with the intention of closing the communication gap between academic research and stakeholders that requires information on the prospects of future invasions.

Historical and contemporary trophic niche partitioning among Laurentian Great Lakes coregonines.

Anthropogenic activities have significantly altered freshwater fish communities. Extirpations of deepwater coregonines (Coregonus spp.), a diverse group of fish species, have left vast areas of the Laurentian Great Lakes devoid of a deepwater fish community. Currently, fisheries managers are considering restoring populations by reintroducing deepwater coregonines from Lake Superior and Lake Nipigon. However, little is known about the historical ecology of deepwater coregonines, and species characterization has proved difficult. We used stable isotope analysis of museum-preserved and contemporary specimens to investigate if (1) coregonine species historically occupied distinct niches and (2) the pattern of trophic niche partitioning has changed over the last century. Across all lakes, individual species occupied distinct trophic niches, confirming that these species were ecologically distinct. Understanding trophic niche partitioning helps resolve uncertainty about distinctness of species within and across lakes and may provide a better ecological basis for rehabilitation of Great Lakes food webs and ecosystems.

Flux of aquatic insect productivity to land: comparison of lentic and lotic ecosystems.

Recently, food web studies have started exploring how resources from one habitat or ecosystem influence trophic interactions in a recipient ecosystem. Benthic production in lakes and streams can be exported to terrestrial habitats via emerging aquatic insects and can therefore link aquatic and terrestrial ecosystems. In this study, we develop a general conceptual model that highlights zoobenthic production, insect emergence, and ecosystem geometry (driven principally by area-to-edge ratio) as important factors modulating the flux of aquatic production across the ecosystem boundary. Emerging insect flux, defined as total insect production emerging per meter of shoreline (g C x m(-1) x yr(-1)) is then distributed inland using decay functions and is used to estimate insect deposition rate to terrestrial habitats (g C x m(-2) x yr(-1)). Using empirical data from the literature, we simulate insect fluxes across the water-land ecosystem boundary to estimate the distribution of fluxes and insect deposition inland for lakes and streams. In general, zoobenthos in streams are more productive than in lakes (6.67 vs. 1.46 g C x m(-2) x yr(-1)) but have lower insect emergence to aquatic production ratios (0.19 vs. 0.30). However, as stream width is on average smaller than lake radius, this results in flux (F) estimates 2 1/2 times greater for lakes than for streams. Ultimately, insect deposition onto land (within 100 m of shore) adjacent to average-sized lakes (10-ha lakes, 0.021 g C x m(-2) x yr(-1)) is greater than for average-sized streams (4 m width, 0.002 g C x m(-2) x yr(-1)) used in our comparisons. For the average lake (both in size and productivity), insect deposition rate approaches estimates of terrestrial secondary production in low-productivity ecosystems (e.g., deserts and tundra, approximately 0.07 g C x m(-2) x yr(-1)). However, larger lakes (1300 ha) and streams (16 m) can have average insect deposition rates (approximately 0.01-2.4 g C x m(-2) x yr(-1)) comparable to estimates of secondary production of more productive ecosystems such as grasslands. Because of the potentially large inputs of emerging aquatic insects into terrestrial habitats, ecosystem processes and terrestrial consumers can be influenced by insect inputs. The relative contribution of lakes and streams to this flux will vary among landscapes depending on the number and size of these ecosystems types on the landscape.

Nitrogen stable isotopes in streams: effects of agricultural sources and transformations.

The nitrogen stable isotope ratio of biological tissue has been proposed as an indicator of anthropogenic N inputs to aquatic ecosystems, but overlap in the isotopic signatures of various N sources and transformations make definitive attribution of processes difficult. We collected primary consumer invertebrates from streams in agricultural settings in Wisconsin, U.S.A., to evaluate the relative influence of animal manure, inorganic fertilizer, and denitrification on biotic delta15N. Variance in biotic delta15N was explained by inorganic fertilizer inputs and the percentage of wetland land cover in the watershed, but not by animal manure inputs. These results suggest that denitrification of inorganic fertilizer is the primary driver of delta15N variability among the study sites. Comparison with previously collected stream water NO3-N concentrations at the same sites supports the role of denitrification; for a given N application rate, streams with high biotic delta15N had low NO3-N concentrations. The lack of a manure signal in biotic delta15N may be due its high ammonia content, which can be dispersed outside the range of its application by volatilization. Based on our findings and on agricultural census data for the entire United States, inorganic fertilizer is more likely than manure to drive variability in biotic delta15N and to cause excessive nitrogen concentrations in streams.

Landscape planning for agricultural non-point source pollution reduction. II. Balancing watershed size, number of watersheds, and implementation effort.

Agricultural non-point source (NPS) pollution poses a severe threat to water quality and aquatic ecosystems. In response, tremendous efforts have been directed toward reducing these pollution inputs by implementing agricultural conservation practices. Although conservation practices reduce pollution inputs from individual fields, scaling pollution control benefits up to the watershed level (i.e., improvements in stream water quality) has been a difficult challenge. This difficulty highlights the need for NPS reduction programs that focus efforts within target watersheds and at specific locations within target watersheds, with the ultimate goal of improving stream water quality. Fundamental program design features for NPS control programs--i.e., number of watersheds in the program, total watershed area, and level of effort expended within watersheds--have not been considered in any sort of formal analysis. Here, we present an optimization model that explores the programmatic and environmental trade-offs between these design choices. Across a series of annual program budgets ranging from $2 to $200 million, the optimal number of watersheds ranged from 3 to 27; optimal watershed area ranged from 29 to 214 km(2); and optimal expenditure ranged from $21,000 to $35,000/km(2). The optimal program configuration was highly dependent on total program budget. Based on our general findings, we delineated hydrologically complete and spatially independent watersheds ranging in area from 20 to 100 km(2). These watersheds are designed to serve as implementation units for a targeted NPS pollution control program currently being developed in Wisconsin.

Landscape planning for agricultural nonpoint source pollution reduction III: assessing phosphorus and sediment reduction potential.

Riparian buffers have the potential to improve stream water quality in agricultural landscapes. This potential may vary in response to landscape characteristics such as soils, topography, land use, and human activities, including legacies of historical land management. We built a predictive model to estimate the sediment and phosphorus load reduction that should be achievable following the implementation of riparian buffers; then we estimated load reduction potential for a set of 1598 watersheds (average 54 km(2)) in Wisconsin. Our results indicate that land cover is generally the most important driver of constituent loads in Wisconsin streams, but its influence varies among pollutants and according to the scale at which it is measured. Physiographic (drainage density) variation also influenced sediment and phosphorus loads. The effect of historical land use on present-day channel erosion and variation in soil texture are the most important sources of phosphorus and sediment that riparian buffers cannot attenuate. However, in most watersheds, a large proportion (approximately 70%) of these pollutants can be eliminated from streams with buffers. Cumulative frequency distributions of load reduction potential indicate that targeting pollution reduction in the highest 10% of Wisconsin watersheds would reduce total phosphorus and sediment loads in the entire state by approximately 20%. These results support our approach of geographically targeting nonpoint source pollution reduction at multiple scales, including the watershed scale.

Benthic algal production across lake size gradients: interactions among morphometry, nutrients, and light.

Attached algae play a minor role in conceptual and empirical models of lake ecosystem function but paradoxically form the energetic base of food webs that support a wide variety of fishes. To explore the apparent mismatch between perceived limits on contributions of periphyton to whole-lake primary production and its importance to consumers, we modeled the contribution of periphyton to whole-ecosystem primary production across lake size, shape, and nutrient gradients. The distribution of available benthic habitat for periphyton is influenced by the ratio of mean depth to maximum depth (DR = z/ z(max)). We modeled total phytoplankton production from water-column nutrient availability, z, and light. Periphyton production was a function of light-saturated photosynthesis (BPmax) and light availability at depth. The model demonstrated that depth ratio (DR) and light attenuation strongly determined the maximum possible contribution of benthic algae to lake production, and the benthic proportion of whole-lake primary production (BPf) declined with increasing nutrients. Shallow lakes (z < or =5 m) were insensitive to DR and were dominated by either benthic or pelagic primary productivity depending on trophic status. Moderately deep oligotrophic lakes had substantial contributions by benthic primary productivity at low depth ratios and when maximum benthic photosynthesis was moderate or high. Extremely large, deep lakes always had low fractional contributions of benthic primary production. An analysis of the world's largest lakes showed that the shapes of natural lakes shift increasingly toward lower depth ratios with increasing depth, maximizing the potential importance of littoral primary production in large-lake food webs. The repeatedly demonstrated importance of periphyton to lake food webs may reflect the combination of low depth ratios and high light penetration characteristic of large, oligotrophic lakes that in turn lead to substantial contributions of periphyton to autochthonous production.

Go big or don't? A field-based diet evaluation of freshwater piscivore and prey fish size relationships.

Body size governs predator-prey interactions, which in turn structure populations, communities, and food webs. Understanding predator-prey size relationships is valuable from a theoretical perspective, in basic research, and for management applications. However, predator-prey size data are limited and costly to acquire. We quantified predator-prey total length and mass relationships for several freshwater piscivorous taxa: crappie (Pomoxis spp.), largemouth bass (Micropterus salmoides), muskellunge (Esox masquinongy), northern pike (Esox lucius), rock bass (Ambloplites rupestris), smallmouth bass (Micropterus dolomieu), and walleye (Sander vitreus). The range of prey total lengths increased with predator total length. The median and maximum ingested prey total length varied with predator taxon and length, but generally ranged from 10-20% and 32-46% of predator total length, respectively. Predators tended to consume larger fusiform prey than laterally compressed prey. With the exception of large muskellunge, predators most commonly consumed prey between 16 and 73 mm. A sensitivity analysis indicated estimates can be very accurate at sample sizes greater than 1,000 diet items and fairly accurate at sample sizes greater than 100. However, sample sizes less than 50 should be evaluated with caution. Furthermore, median log10 predator-prey body mass ratios ranged from 1.9-2.5, nearly 50% lower than values previously reported for freshwater fishes. Managers, researchers, and modelers could use our findings as a tool for numerous predator-prey evaluations from stocking size optimization to individual-based bioenergetics analyses identifying prey size structure. To this end, we have developed a web-based user interface to maximize the utility of our models that can be found at www.LakeEcologyLab.org/pred_prey.

Quantitative approaches to the analysis of stable isotope food web data.

Ecologists use stable isotopes (delta13C, delta15N) to better understand food webs and explore trophic interactions in ecosystems. Traditionally, delta13C vs. delta15N bi-plots have been used to describe food web structure for a single time period or ecosystem. Comparisons of food webs across time and space are increasing, but development of statistical approaches for testing hypotheses regarding food web change has lagged behind. Here we present statistical methodologies for quantitatively comparing stable isotope food web data. We demonstrate the utility of circular statistics and hypothesis tests for quantifying directional food web differences using two case studies: an arthropod salt marsh community across a habitat gradient and a freshwater fish community from Lake Tahoe, USA, over a 120-year time period. We calculated magnitude and mean angle of change (theta) for each species in food web space using mean delta13C and delta15N of each species as the x, y coordinates. In the coastal salt marsh, arthropod consumers exhibited a significant shift toward dependence on Spartina, progressing from a habitat invaded by Phragmites to a restored Spartina habitat. In Lake Tahoe, we found that all species from the freshwater fish community shifted in the same direction in food web space toward more pelagic-based production with the introduction of nonnative Mysis relicta and onset of cultural eutrophication. Using circular statistics to quantitatively analyze stable isotope food web data, we were able to gain significant insight into patterns and changes in food web structure that were not evident from qualitative comparisons. As more ecologists incorporate a food web perspective into ecosystem analysis, these statistical tools can provide a basis for quantifying directional food web differences from standard isotope data.