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.

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.

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.

Temperature effects on fish production across a natural thermal gradient.

Global warming is widely predicted to reduce the biomass production of top predators, or even result in species loss. Several exceptions to this expectation have been identified, however, and it is vital that we understand the underlying mechanisms if we are to improve our ability to predict future trends. Here, we used a natural warming experiment in Iceland and quantitative theoretical predictions to investigate the success of brown trout as top predators across a stream temperature gradient (4-25 °C). Brown trout are at the northern limit of their geographic distribution in this system, with ambient stream temperatures below their optimum for maximal growth, and above it in the warmest streams. A five-month mark-recapture study revealed that population abundance, biomass, growth rate, and production of trout all increased with stream temperature. We identified two mechanisms that contributed to these responses: (1) trout became more selective in their diet as stream temperature increased, feeding higher in the food web and increasing in trophic position; and (2) trophic transfer through the food web was more efficient in the warmer streams. We found little evidence to support a third potential mechanism: that external subsidies would play a more important role in the diet of trout with increasing stream temperature. Resource availability was also amplified through the trophic levels with warming, as predicted by metabolic theory in nutrient-replete systems. These results highlight circumstances in which top predators can thrive in warmer environments and contribute to our knowledge of warming impacts on natural communities and ecosystem functioning.

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.

Regional impacts of warming on biodiversity and biomass in high latitude stream ecosystems across the Northern Hemisphere.

Warming can have profound impacts on ecological communities. However, explorations of how differences in biogeography and productivity might reshape the effect of warming have been limited to theoretical or proxy-based approaches: for instance, studies of latitudinal temperature gradients are often conflated with other drivers (e.g., species richness). Here, we overcome these limitations by using local geothermal temperature gradients across multiple high-latitude stream ecosystems. Each suite of streams (6-11 warmed by 1-15°C above ambient) is set within one of five regions (37 streams total); because the heating comes from the bedrock and is not confounded by changes in chemistry, we can isolate the effect of temperature. We found a negative overall relationship between diatom and invertebrate species richness and temperature, but the strength of the relationship varied regionally, declining more strongly in regions with low terrestrial productivity. Total invertebrate biomass increased with temperature in all regions. The latter pattern combined with the former suggests that the increased biomass of tolerant species might compensate for the loss of sensitive species. Our results show that the impact of warming can be dependent on regional conditions, demonstrating that local variation should be included in future climate projections rather than simply assuming universal relationships.

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.

Functional diversity and community assembly of river invertebrates show globally consistent responses to decreasing glacier cover.

Global change threatens invertebrate biodiversity and its central role in numerous ecosystem functions and services. Functional trait analyses have been advocated to uncover global mechanisms behind biodiversity responses to environmental change, but the application of this approach for invertebrates is underdeveloped relative to other organism groups. From an evaluation of 363 records comprising >1.23 million invertebrates collected from rivers across nine biogeographic regions on three continents, consistent responses of community trait composition and diversity to replicated gradients of reduced glacier cover are demonstrated. After accounting for a systematic regional effect of latitude, the processes shaping river invertebrate functional diversity are globally consistent. Analyses nested within individual regions identified an increase in functional diversity as glacier cover decreases. Community assembly models demonstrated that dispersal limitation was the dominant process underlying these patterns, although environmental filtering was also evident in highly glacierized basins. These findings indicate that predictable mechanisms govern river invertebrate community responses to decreasing glacier cover globally.