Environmental warming increases the importance of high-turnover energy channels in stream food webs.

Warming temperatures are altering communities and trophic networks across Earth's ecosystems. While the overall influence of warming on food webs is often context-dependent, increasing temperatures are predicted to change communities in two fundamental ways: (1) by reducing average body size and (2) by increasing individual metabolic rates. These warming-induced changes have the potential to influence the distribution of food web fluxes, food web stability, and the relative importance of deterministic and stochastic ecological processes shaping community assembly. Here, we quantified patterns and the relative distribution of organic matter fluxes through stream food webs spanning a broad natural temperature gradient (5-27°C). We then related these patterns to species and community trait distributions of mean body size and population biomass turnover (P:B) within and across streams. We predicted that (1) communities in warmer streams would exhibit smaller body size and higher P:B and (2) organic matter fluxes within warmer communities would increasingly skew toward smaller, higher P:B populations. Across the temperature gradient, warmer communities were characterized by smaller body size (~9% per °C) and higher P:B (~7% faster turnover per °C) populations on average. Additionally, organic matter fluxes within warmer streams were increasingly skewed toward higher P:B populations, demonstrating that warming can restructure organic matter fluxes in both an absolute and relative sense. With warming, the relative distribution of organic matter fluxes was decreasingly likely to arise through the random sorting of species, suggesting stronger selection for traits driving high turnover with increasing temperature. Our study suggests that a warming world will favor energy fluxes through "smaller and faster" populations, and that these changes may be more predictable than previously thought.

Human activities shape global patterns of decomposition rates in rivers.

Rivers and streams contribute to global carbon cycling by decomposing immense quantities of terrestrial plant matter. However, decomposition rates are highly variable and large-scale patterns and drivers of this process remain poorly understood. Using a cellulose-based assay to reflect the primary constituent of plant detritus, we generated a predictive model (81% variance explained) for cellulose decomposition rates across 514 globally distributed streams. A large number of variables were important for predicting decomposition, highlighting the complexity of this process at the global scale. Predicted cellulose decomposition rates, when combined with genus-level litter quality attributes, explain published leaf-litter-decomposition rates with impressive accuracy (70% variance explained). Our global map provides estimates of rates across vast understudied areas of Earth, and reveals rapid decomposition across continental-scale areas dominated by human activities.

Facilitation strength across environmental and beneficiary trait gradients in stream communities.

Ecosystem engineers modify habitats in ways that facilitate other community members by ameliorating harsh conditions. The strength of such facilitation is predicted to be influenced by both beneficiary traits and abiotic context. One key trait of animals that could control the strength of facilitation is beneficiary body size because it should determine how beneficiaries fit within and exploit stress ameliorating habitat modifications. However, few studies have measured how beneficiary body size relates to facilitation strength along environmental gradients. We examined how the strength of facilitation by net-spinning caddisflies on invertebrate communities in streams varied along an elevation gradient and based on traits of the invertebrate beneficiaries. We measured whether use of silk retreats as habitat concentrated invertebrate density and biomass compared to surrounding rock surface habitat and whether the use of retreat habitat varied across body sizes of community members along the gradient. We found that retreats substantially concentrated the densities of a diversity of taxa including eight different Orders, and this effect was greatest at high elevations. Caddisfly retreats also concentrated invertebrate biomass more as elevation increased. Body size of invertebrates inhabiting retreats was lower than that of surrounding rock habitats at low elevation sites, however, body size between retreats and rocks converged at higher elevation sites. Additionally, the body size of invertebrates found in retreats varied within and across taxa. Specifically, caddisfly retreats functioned as a potential nursery for taxa with large maximal body sizes. However, the patterns of this taxon-specific nursery effect were not influenced by elevation unlike the patterns observed based on community-level body size. Collectively, our results indicate that invertebrates use retreats in earlier life stages or when they are smaller in body size independent of life stage. Furthermore, our analysis suggests that facilitation strength intensifies as elevation increases within stream invertebrate communities. Further consideration of how trait variation and environmental gradients interact to determine the strength and direction of biotic interactions will be important as species ranges and environmental conditions continue to shift.

Landscape diversity promotes stable food-web architectures in large rivers.

Uncovering relationships between landscape diversity and species interactions is crucial for predicting how ongoing land-use change and homogenization will impact the stability and persistence of communities. However, such connections have rarely been quantified in nature. We coupled high-resolution river sonar imaging with annualized energetic food webs to quantify relationships among habitat diversity, energy flux, and trophic interaction strengths in large-river food-web modules that support the endangered Pallid Sturgeon. Our results demonstrate a clear relationship between habitat diversity and species interaction strengths, with more diverse foraging landscapes containing higher production of prey and a greater proportion of weak and potentially stabilizing interactions. Additionally, rare patches of large and relatively stable river sediments intensified these effects and further reduced interaction strengths by increasing prey diversity. Our findings highlight the importance of landscape characteristics in promoting stabilizing food-web architectures and provide direct relevance for future management of imperilled species in a simplified and rapidly changing world.

Resource modification by ecosystem engineers generates hotspots of stream community assembly and ecosystem function.

Ecosystem engineers can generate hotspots of ecological structure and function by facilitating the aggregation of both resources and consumers. However, nearly all examples of such engineered hotspots come from long-lived foundation species, such as marine and freshwater mussels, intertidal cordgrasses, and alpine cushion plants, with less attention given to small-bodied and short-lived animals. Insects often have rapid life cycles and high population densities and are among the most diverse and ubiquitous animals on earth. Although these taxa have the potential to generate hotspots and heterogeneity comparable to that of foundation species, few studies have examined this possibility. We conducted a mesocosm experiment to examine the degree to which a stream insect ecosystem engineer, the net-spinning caddisfly (Tricoptera:Hydropsychidae), creates hotspots by facilitating invertebrate community assembly. Our experiment used two treatments: (1) stream benthic habitat with patches of caddisfly engineers present and (2) a control treatment with no caddisflies present. We show that compared to controls, caddisflies increased local resource availability measured as particulate organic matter (POM) by 43%, ecosystem respiration (ER) by 70%, and invertebrate density, biomass, and richness by 96%, 244%, and 72%, respectively. These changes resulted in increased spatial variation of POM by 25%, invertebrate density by 76%, and ER by 29% compared to controls, indicating a strong effect of caddisflies on ecological heterogeneity. We found a positive relationship between invertebrate density and ammonium concentration in the caddisfly treatment, but no such relationship in the control, indicating that either caddisflies themselves or the invertebrate aggregations they create increased nutrient availability. When accounting for the amount of POM, caddisfly treatments increased invertebrate density by 48% and richness by 40% compared to controls, suggesting that caddisflies may also enhance the nutritional quality of resources for the invertebrate assemblage. The caddisfly treatment also increased the rate of ecosystem respiration as a function of increasing POM compared to the control. Our study demonstrates that insect ecosystem engineers can generate heterogeneity by concentrating local resources and consumers, with consequences for carbon and nutrient cycling.

A global synthesis of human impacts on the multifunctionality of streams and rivers.

Human impacts, particularly nutrient pollution and land-use change, have caused significant declines in the quality and quantity of freshwater resources. Most global assessments have concentrated on species diversity and composition, but effects on the multifunctionality of streams and rivers remain unclear. Here, we analyse the most comprehensive compilation of stream ecosystem functions to date to provide an overview of the responses of nutrient uptake, leaf litter decomposition, ecosystem productivity, and food web complexity to six globally pervasive human stressors. We show that human stressors inhibited ecosystem functioning for most stressor-function pairs. Nitrate uptake efficiency was most affected and was inhibited by 347% due to agriculture. However, concomitant negative and positive effects were common even within a given stressor-function pair. Some part of this variability in effect direction could be explained by the structural heterogeneity of the landscape and latitudinal position of the streams. Ranking human stressors by their absolute effects on ecosystem multifunctionality revealed significant effects for all studied stressors, with wastewater effluents (194%), agriculture (148%), and urban land use (137%) having the strongest effects. Our results demonstrate that we are at risk of losing the functional backbone of streams and rivers if human stressors persist in contemporary intensity, and that freshwaters are losing critical ecosystem services that humans rely on. We advocate for more studies on the effects of multiple stressors on ecosystem multifunctionality to improve the functional understanding of human impacts. Finally, freshwater management must shift its focus toward an ecological function-based approach and needs to develop strategies for maintaining or restoring ecosystem functioning of streams and rivers.

Combined carbon flows through detritus, microbes, and animals in reference and experimentally enriched stream ecosystems.

Tracking carbon (C) flow through ecosystems requires quantification of myriad biophysical processes, including C routing through microbial and metazoan food webs. Yet detailed organic matter budgets are rarely combined with simultaneous measurement of C flows supporting microbial and animal production. Here, we synthesize concurrent data sets on organic matter, microbes, and macroinvertebrates from two detritus-based stream ecosystems, one of which was subject to experimental nitrogen (N) and phosphorus (P) enrichment. Our synthesis provides new insights into C flow through forest stream ecosystems. Over 3 yr, the reference stream showed a striking balance of inputs and outputs, with a mean surplus of only 7 g C·m-2 ·yr-1 (~1% of annual inputs), presumably stored in sediments as fine particulate organic matter (FPOM). In contrast, N and P enrichment over 2 yr resulted in severe deficits of C (-576 g C·m-2 ·yr-1 or ~170% of annual inputs), a shortfall presumably met by stored C. Our data set provides an ecosystem-based estimate of the fate of forest litter C at ambient nutrient concentrations: 6.2% was leached as dissolved organic C, 40.6% and 8.5% flowed to litter-associated fungi and bacteria, respectively, 7.5% was consumed by macroinvertebrates, 1.8% was exported as coarse particles, and the remainder (35.4%) was presumably fragmented by biophysical processes. Our calculations also allowed an estimate of inputs into the heterogeneous FPOM pool, which is otherwise difficult to obtain. At naturally low nutrient concentrations, 50.7% was derived from fragmented litter, 39.1% from microbial biomass (mostly fungal), and 10.2% from macroinvertebrate egesta. Nutrient addition drove large changes in C fluxes in the experimental stream, especially in flows of leaf litter to fungi (×1.7 pretreatment) and macroinvertebrates (×2.7), and of FPOM to hydrologic export (×2.6). Our results underscore the key roles of both microbes and metazoans in controlling C flow through detritus-based ecosystems, as well as how release from persistent nutrient limitation may perturb steady-state conditions of C inputs vs. outputs. Our analysis also suggests areas for future research, including assessing the relative importance of stored vs. recycled C in fueling detrital food webs subject to altered nutrient regimes and other global-change drivers.

Resource supply governs the apparent temperature dependence of animal production in stream ecosystems.

Rising global temperatures are changing how energy and materials move through ecosystems, with potential consequences for the role of animals in these processes. We tested a central prediction of the metabolic scaling framework-the temperature independence of animal community production-using a series of geothermally heated streams and a comprehensive empirical analysis. We show that the apparent temperature sensitivity of animal production was consistent with theory for individuals (Epind  = 0.64 vs. 0.65 eV), but strongly amplified relative to theoretical expectations for communities, both among (Epamong  = 0.67 vs. 0 eV) and within (Epwithin  = 1.52 vs. 0 eV) streams. After accounting for spatial and temporal variation in resources, we show that the apparent positive effect of temperature was driven by resource supply, providing strong empirical support for the temperature independence of invertebrate production and the necessary inclusion of resources in metabolic scaling efforts.

Thermal niche diversity and trophic redundancy drive neutral effects of warming on energy flux through a stream food web.

Climate warming is predicted to alter routing and flows of energy through food webs because of the critical and varied effects of temperature on physiological rates, community structure, and trophic dynamics. Few studies, however, have experimentally assessed the net effect of warming on energy flux and food web dynamics in natural intact communities. Here, we test how warming affects energy flux and the trophic basis of production in a natural invertebrate food web by experimentally heating a stream reach in southwest Iceland by ~4°C for 2 yr and comparing its response to an unheated reference stream. Previous results from this experiment showed that warming led to shifts in the structure of the invertebrate assemblage, with estimated increases in total metabolic demand but no change in annual secondary production. We hypothesized that elevated metabolic demand and invariant secondary production would combine to increase total consumption of organic matter in the food web, if diet composition did not change appreciably with warming. Dietary composition of primary consumers indeed varied little between streams and among years, with gut contents primarily consisting of diatoms (72.9%) and amorphous detritus (19.5%). Diatoms dominated the trophic basis of production of primary consumers in both study streams, contributing 79-86% to secondary production. Although warming increased the flux of filamentous algae within the food web, total resource consumption did not increase as predicted. The neutral net effect of warming on total energy flow through the food web was a result of taxon-level variation in responses to warming, a neutral effect on total invertebrate production, and strong trophic redundancy within the invertebrate assemblage. Thus, food webs characterized by a high degree of trophic redundancy may be more resistant to the effects of climate warming than those with more diverse and specialized consumers.

Global patterns and drivers of ecosystem functioning in rivers and riparian zones.

River ecosystems receive and process vast quantities of terrestrial organic carbon, the fate of which depends strongly on microbial activity. Variation in and controls of processing rates, however, are poorly characterized at the global scale. In response, we used a peer-sourced research network and a highly standardized carbon processing assay to conduct a global-scale field experiment in greater than 1000 river and riparian sites. We found that Earth's biomes have distinct carbon processing signatures. Slow processing is evident across latitudes, whereas rapid rates are restricted to lower latitudes. Both the mean rate and variability decline with latitude, suggesting temperature constraints toward the poles and greater roles for other environmental drivers (e.g., nutrient loading) toward the equator. These results and data set the stage for unprecedented "next-generation biomonitoring" by establishing baselines to help quantify environmental impacts to the functioning of ecosystems at a global scale.

Increased resource use efficiency amplifies positive response of aquatic primary production to experimental warming.

Climate warming is affecting the structure and function of river ecosystems, including their role in transforming and transporting carbon (C), nitrogen (N), and phosphorus (P). Predicting how river ecosystems respond to warming has been hindered by a dearth of information about how otherwise well-studied physiological responses to temperature scale from organismal to ecosystem levels. We conducted an ecosystem-level temperature manipulation to quantify how coupling of stream ecosystem metabolism and nutrient uptake responded to a realistic warming scenario. A ~3.3°C increase in mean water temperature altered coupling of C, N, and P fluxes in ways inconsistent with single-species laboratory experiments. Net primary production tripled during the year of experimental warming, while whole-stream N and P uptake rates did not change, resulting in 289% and 281% increases in autotrophic dissolved inorganic N and P use efficiency (UE), respectively. Increased ecosystem production was a product of unexpectedly large increases in mass-specific net primary production and autotroph biomass, supported by (i) combined increases in resource availability (via N mineralization and N2 fixation) and (ii) elevated resource use efficiency, the latter associated with changes in community structure. These large changes in C and nutrient cycling could not have been predicted from the physiological effects of temperature alone. Our experiment provides clear ecosystem-level evidence that warming can shift the balance between C and nutrient cycling in rivers, demonstrating that warming will alter the important role of in-stream processes in C, N, and P transformations. Moreover, our results reveal a key role for nutrient supply and use efficiency in mediating responses of primary producers to climate warming.

Bridging Food Webs, Ecosystem Metabolism, and Biogeochemistry Using Ecological Stoichiometry Theory.

Although aquatic ecologists and biogeochemists are well aware of the crucial importance of ecosystem functions, i.e., how biota drive biogeochemical processes and vice-versa, linking these fields in conceptual models is still uncommon. Attempts to explain the variability in elemental cycling consequently miss an important biological component and thereby impede a comprehensive understanding of the underlying processes governing energy and matter flow and transformation. The fate of multiple chemical elements in ecosystems is strongly linked by biotic demand and uptake; thus, considering elemental stoichiometry is important for both biogeochemical and ecological research. Nonetheless, assessments of ecological stoichiometry (ES) often focus on the elemental content of biota rather than taking a more holistic view by examining both elemental pools and fluxes (e.g., organismal stoichiometry and ecosystem process rates). ES theory holds the promise to be a unifying concept to link across hierarchical scales of patterns and processes in ecology, but this has not been fully achieved. Therefore, we propose connecting the expertise of aquatic ecologists and biogeochemists with ES theory as a common currency to connect food webs, ecosystem metabolism, and biogeochemistry, as they are inherently concatenated by the transfer of carbon, nitrogen, and phosphorous through biotic and abiotic nutrient transformation and fluxes. Several new studies exist that demonstrate the connections between food web ecology, biogeochemistry, and ecosystem metabolism. In addition to a general introduction into the topic, this paper presents examples of how these fields can be combined with a focus on ES. In this review, a series of concepts have guided the discussion: (1) changing biogeochemistry affects trophic interactions and ecosystem processes by altering the elemental ratios of key species and assemblages; (2) changing trophic dynamics influences the transformation and fluxes of matter across environmental boundaries; (3) changing ecosystem metabolism will alter the chemical diversity of the non-living environment. Finally, we propose that using ES to link nutrient cycling, trophic dynamics, and ecosystem metabolism would allow for a more holistic understanding of ecosystem functions in a changing environment.

Shifts in community size structure drive temperature invariance of secondary production in a stream-warming experiment.

A central question at the interface of food-web and climate change research is how secondary production, or the formation of heterotroph biomass over time, will respond to rising temperatures. The metabolic theory of ecology (MTE) hypothesizes the temperature-invariance of secondary production, driven by matched and opposed forces that reduce biomass of heterotrophs while increasing their biomass turnover rate (production : biomass, or P:B) with warming. To test this prediction at the whole community level, we used a geothermal heat exchanger to experimentally warm a stream in southwest Iceland by 3.8°C for two years. We quantified invertebrate community biomass, production, and P : B in the experimental stream and a reference stream for one year prior to warming and two years during warming. As predicted, warming had a neutral effect on community production, but this result was not driven by opposing effects on community biomass and P:B. Instead, warming had a positive effect on both the biomass and production of larger-bodied, slower-growing taxa (e.g., larval black flies, dipteran predators, snails) and a negative effect on small-bodied taxa with relatively high growth rates (e.g., ostracods, larval chironomids). We attribute these divergent responses to differences in thermal preference between small- vs. large-bodied taxa. Although metabolic demand vs. resource supply must ultimately constrain community production, our results highlight the potential for idiosyncratic community responses to warming, driven by variation in thermal preference and body size within regional species pools.

A global database of nitrogen and phosphorus excretion rates of aquatic animals.

Animals can be important in modulating ecosystem-level nutrient cycling, although their importance varies greatly among species and ecosystems. Nutrient cycling rates of individual animals represent valuable data for testing the predictions of important frameworks such as the Metabolic Theory of Ecology (MTE) and ecological stoichiometry (ES). They also represent an important set of functional traits that may reflect both environmental and phylogenetic influences. Over the past two decades, studies of animal-mediated nutrient cycling have increased dramatically, especially in aquatic ecosystems. Here we present a global compilation of aquatic animal nutrient excretion rates. The dataset includes 10,534 observations from freshwater and marine animals of N and/or P excretion rates. These observations represent 491 species, including most aquatic phyla. Coverage varies greatly among phyla and other taxonomic levels. The dataset includes information on animal body size, ambient temperature, taxonomic affiliations, and animal body N:P. This data set was used to test predictions of MTE and ES, as described in Vanni and McIntyre (2016; Ecology DOI: 10.1002/ecy.1582).

Experimental whole-stream warming alters community size structure.

How ecological communities respond to predicted increases in temperature will determine the extent to which Earth's biodiversity and ecosystem functioning can be maintained into a warmer future. Warming is predicted to alter the structure of natural communities, but robust tests of such predictions require appropriate large-scale manipulations of intact, natural habitat that is open to dispersal processes via exchange with regional species pools. Here, we report results of a two-year whole-stream warming experiment that shifted invertebrate assemblage structure via unanticipated mechanisms, while still conforming to community-level metabolic theory. While warming by 3.8 °C decreased invertebrate abundance in the experimental stream by 60% relative to a reference stream, total invertebrate biomass was unchanged. Associated shifts in invertebrate assemblage structure were driven by the arrival of new taxa and a higher proportion of large, warm-adapted species (i.e., snails and predatory dipterans) relative to small-bodied, cold-adapted taxa (e.g., chironomids and oligochaetes). Experimental warming consequently shifted assemblage size spectra in ways that were unexpected, but consistent with thermal optima of taxa in the regional species pool. Higher temperatures increased community-level energy demand, which was presumably satisfied by higher primary production after warming. Our experiment demonstrates how warming reassembles communities within the constraints of energy supply via regional exchange of species that differ in thermal physiological traits. Similar responses will likely mediate impacts of anthropogenic warming on biodiversity and ecosystem function across all ecological communities.

Seasonal Change in Trophic Niche of Adfluvial Arctic Grayling (Thymallus arcticus) and Coexisting Fishes in a High-Elevation Lake System.

Introduction of non-native species is a leading threat to global aquatic biodiversity. Competition between native and non-native species is often influenced by changes in suitable habitat or food availability. We investigated diet breadth and degree of trophic niche overlap for a fish assemblage of native and non-native species inhabiting a shallow, high elevation lake system. This assemblage includes one of the last remaining post-glacial endemic populations of adfluvial Arctic grayling (Thymallus arcticus) in the contiguous United States. We examined gut contents and stable isotope values of fish taxa in fall and spring to assess both short- (days) and long-term (few months) changes in trophic niches. We incorporate these short-term (gut contents) data into a secondary isotope analysis using a Bayesian statistical framework to estimate long-term trophic niche. Our data suggest that in this system, Arctic grayling share both a short- and long-term common food base with non-native trout of cutthroat x rainbow hybrid species (Oncorhynchus clarkia bouvieri x Oncorhynchus mykiss) and brook trout (Salvelinus fontinalis). In addition, trophic niche overlap among Arctic grayling, hybrid trout, and brook trout appeared to be stronger during spring than fall. In contrast, the native species of Arctic grayling, burbot (Lota lota), and suckers (Catostomus spp.) largely consumed different prey items. Our results suggest strong seasonal differences in trophic niche overlap among Arctic grayling and non-native trout, with a potential for greatest competition for food during spring. We suggest that conservation of endemic Arctic grayling in high-elevation lakes will require recognition of the potential for coexisting non-native taxa to impede well-intentioned recovery efforts.

Warming alters coupled carbon and nutrient cycles in experimental streams.

Although much effort has been devoted to quantifying how warming alters carbon cycling across diverse ecosystems, less is known about how these changes are linked to the cycling of bioavailable nitrogen and phosphorus. In freshwater ecosystems, benthic biofilms (i.e. thin films of algae, bacteria, fungi, and detrital matter) act as biogeochemical hotspots by controlling important fluxes of energy and material. Understanding how biofilms respond to warming is thus critical for predicting responses of coupled elemental cycles in freshwater systems. We developed biofilm communities in experimental streamside channels along a gradient of mean water temperatures (7.5-23.6 °C), while closely maintaining natural diel and seasonal temperature variation with a common water and propagule source. Both structural (i.e. biomass, stoichiometry, assemblage structure) and functional (i.e. metabolism, N2 -fixation, nutrient uptake) attributes of biofilms were measured on multiple dates to link changes in carbon flow explicitly to the dynamics of nitrogen and phosphorus. Temperature had strong positive effects on biofilm biomass (2.8- to 24-fold variation) and net ecosystem productivity (44- to 317-fold variation), despite extremely low concentrations of limiting dissolved nitrogen. Temperature had surprisingly minimal effects on biofilm stoichiometry: carbon:nitrogen (C:N) ratios were temperature-invariant, while carbon:phosphorus (C:P) ratios declined slightly with increasing temperature. Biofilm communities were dominated by cyanobacteria at all temperatures (>91% of total biovolume) and N2 -fixation rates increased up to 120-fold between the coldest and warmest treatments. Although ammonium-N uptake increased with temperature (2.8- to 6.8-fold variation), the much higher N2 -fixation rates supplied the majority of N to the ecosystem at higher temperatures. Our results demonstrate that temperature can alter how carbon is cycled and coupled to nitrogen and phosphorus. The uncoupling of C fixation from dissolved inorganic nitrogen supply produced large unexpected changes in biofilm development, elemental cycling, and likely downstream exports of nutrients and organic matter.

Does N2 fixation amplify the temperature dependence of ecosystem metabolism?

Variation in resource supply can cause variation in temperature dependences of metabolic processes (e.g., photosynthesis and respiration). Understanding such divergence is particularly important when using metabolic theory to predict ecosystem responses to climate warming. Few studies, however, have assessed the effect of temperature-resource interactions on metabolic processes, particularly in cases where the supply of limiting resources exhibits temperature dependence. We investigated the responses of biomass accrual, gross primary production (GPP), community respiration (CR), and N2 fixation to warming during biofilm development in a streamside channel experiment. Areal rates of GPP, CR, biomass accrual, and N2 fixation scaled positively with temperature, showing a 32- to 71-fold range across the temperature gradient (approximately 7 degrees-24 degrees C). Areal N2-fixation rates exhibited apparent activation energies (1.5-2.0 eV; 1 eV = approximately 1.6 x 10(-19) J) approximating the activation energy of the nitrogenase reaction. In contrast, mean apparent activation energies for areal rates of GPP (2.1-2.2 eV) and CR (1.6-1.9 eV) were 6.5- and 2.7-fold higher than estimates based on metabolic theory predictions (i.e., 0.32 and 0.65 eV, respectively) and did not significantly differ from the apparent activation energy observed for N2 fixation. Mass-specific activation energies for N2 fixation (1.4-1.6 eV), GPP (0.3-0.5 eV), and CR (no observed temperature relationship) were near or lower than theoretical predictions. We attribute the divergence of areal activation energies from those predicted by metabolic theory to increases in N2 fixation with temperature, leading to amplified temperature dependences of biomass accrual and areal rates of GPP and R. Such interactions between temperature dependences must be incorporated into metabolic models to improve predictions of ecosystem responses to climate change.

Mercury and selenium accumulation in the Colorado River food web, Grand Canyon, USA.

Mercury (Hg) and selenium (Se) biomagnify in aquatic food webs and are toxic to fish and wildlife. The authors measured Hg and Se in organic matter, invertebrates, and fishes in the Colorado River food web at sites spanning 387 river km downstream of Glen Canyon Dam (AZ, USA). Concentrations were relatively high among sites compared with other large rivers (mean wet wt for 6 fishes was 0.17-1.59 µg g(-1) Hg and 1.35-2.65 µg g(-1) Se), but consistent longitudinal patterns in Hg or Se concentrations relative to the dam were lacking. Mercury increased (slope = 0.147) with δ(15) N, a metric of trophic position, indicating biomagnification similar to that observed in other freshwater systems. Organisms regularly exceeded exposure risk thresholds for wildlife and humans (6-100% and 56-100% of samples for Hg and Se, respectfully, among risk thresholds). In the Colorado River, Grand Canyon, Hg and Se concentrations pose exposure risks for fish, wildlife, and humans, and the findings of the present study add to a growing body of evidence showing that remote ecosystems are vulnerable to long-range transport and subsequent bioaccumulation of contaminants. Management of exposure risks in Grand Canyon will remain a challenge, as sources and transport mechanisms of Hg and Se extend far beyond park boundaries.

Interactions between temperature and nutrients across levels of ecological organization.

Temperature and nutrient availability play key roles in controlling the pathways and rates at which energy and materials move through ecosystems. These factors have also changed dramatically on Earth over the past century as human activities have intensified. Although significant effort has been devoted to understanding the role of temperature and nutrients in isolation, less is known about how these two factors interact to influence ecological processes. Recent advances in ecological stoichiometry and metabolic ecology provide a useful framework for making progress in this area, but conceptual synthesis and review are needed to help catalyze additional research. Here, we examine known and potential interactions between temperature and nutrients from a variety of physiological, community, and ecosystem perspectives. We first review patterns at the level of the individual, focusing on four traits--growth, respiration, body size, and elemental content--that should theoretically govern how temperature and nutrients interact to influence higher levels of biological organization. We next explore the interactive effects of temperature and nutrients on populations, communities, and food webs by synthesizing information related to community size spectra, biomass distributions, and elemental composition. We use metabolic theory to make predictions about how population-level secondary production should respond to interactions between temperature and resource supply, setting up qualitative predictions about the flows of energy and materials through metazoan food webs. Last, we examine how temperature-nutrient interactions influence processes at the whole-ecosystem level, focusing on apparent vs. intrinsic activation energies of ecosystem processes, how to represent temperature-nutrient interactions in ecosystem models, and patterns with respect to nutrient uptake and organic matter decomposition. We conclude that a better understanding of interactions between temperature and nutrients will be critical for developing realistic predictions about ecological responses to multiple, simultaneous drivers of global change, including climate warming and elevated nutrient supply.