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.

Maximum likelihood outperforms binning methods for detecting differences in abundance size spectra across environmental gradients.

Individual body size distributions (ISD) within communities are remarkably consistent across habitats and spatiotemporal scales and can be represented by size spectra, which are described by a power law. The focus of size spectra analysis is to estimate the exponent ( λ ) of the power law. A common application of size spectra studies is to detect anthropogenic pressures. Many methods have been proposed for estimating λ most of which involve binning the data, counting the abundance within bins, and then fitting an ordinary least squares regression in log-log space. However, recent work has shown that binning procedures return biased estimates of λ compared to procedures that directly estimate λ using maximum likelihood estimation (MLE). While it is clear that MLE produces less biased estimates of site-specific λ's, it is less clear how this bias affects the ability to test for changes in λ across space and time, a common question in the ecological literature. Here, we used simulation to compare the ability of two normalised binning methods (equal logarithmic and log2 bins) and MLE to (1) recapture known values of λ , and (2) recapture parameters in a linear regression measuring the change in λ across a hypothetical environmental gradient. We also compared the methods using two previously published body size datasets across a natural temperature gradient and an anthropogenic pollution gradient. Maximum likelihood methods always performed better than common binning methods, which demonstrated consistent bias depending on the simulated values of λ . This bias carried over to the regressions, which were more accurate when λ was estimated using MLE compared to the binning procedures. Additionally, the variance in estimates using MLE methods is markedly reduced when compared to binning methods. The error induced by binning methods can be of similar magnitudes as the variation previously published in experimental and observational studies, bringing into question the effect sizes of previously published results. However, while the methods produced different regression slope estimates, they were in qualitative agreement on the sign of those slopes (i.e. all negative or all positive). Our results provide further support for the direct estimation of λ and its relative variation across environmental gradients using MLE over the more common methods of binning.

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.

Individual size distributions across North American streams vary with local temperature.

Parameters describing the negative relationship between abundance and body size within ecological communities provide a summary of many important biological processes. While it is considered to be one of the few consistent patterns in ecology, spatiotemporal variation of this relationship across continental scale temperature gradients is unknown. Using a database of stream communities collected across North America (18-68°N latitude, -4 to 25°C mean annual air temperature) over 3 years, we constructed 160 individual size distribution (ISD) relationships (i.e. abundance size spectra). The exponent parameter describing ISD's decreased (became steeper) with increasing mean annual temperature, with median slopes varying by ~0.2 units across the 29°C temperature gradient. In addition, total community biomass increased with increasing temperatures, contrary with theoretical predictions. Our study suggests conservation of ISD relationships in streams across broad natural environmental gradients. This supports the emerging use of size-spectra deviations as indicators of fundamental changes to the structure and function of ecological communities.

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.

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.

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.

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.