Abstract
Ecological communities and their ecosystem functions are sensitive to temperature, and aquatic habitats worldwide continue to experience unprecedented warming. Understanding ecological effects of warming requires linking empirical evidence to theories that allow projection to unobserved conditions. Metabolic scaling theory and its tests suggest that warming accelerates ecosystem functions (e.g., oxygen flux), yet this prediction apparently contradicts community-level studies suggesting warming is a stressor that can reduce ecosystem function. We sought to reconcile these predictions with an experimental test of the hypothesis that cascading trophic interactions modify the temperature-dependence of community structure and ecosystem fluxes. In a series of independent freshwater ecosystems exposed to a thermal gradient, we found that warmer temperatures strengthened the trophic cascade increased and indirectly changed community structure by altering grazer species composition and phytoplankton biomass. Temperature-driven community shifts only modestly affected the temperature dependence of net ecosystem oxygen fluxes. Over the 10 °C thermal gradient, NPP and ER increased ∼2.7-fold among ecosystems, while standing phytoplankton biomass declined by 85-95%. The exponential increase in oxygen flux over the thermal gradient, as well as monotonic declines in phytoplankton standing stock, suggested no threshold effects of warming across systems. We also observed temperature variation over time, within ecosystems. For phytoplankton biomass, temporal variation had the opposite effect to spatial variation, suggesting that within-community temporal change in community structure was not predicted by space-for-time substitution. We conclude that food chain length can modify effects of temperature on ecosystem fluxes, but that temperature can still have continuous and positive effects on ecosystem fluxes, consistent with patterns based on large-scale, macroecological comparisons. Changes in community structure, including temperature dependent trophic cascades, may be compatible with prevailing and predictable effects of temperature on ecosystem functions related to fundamental effects of temperature on metabolism.
Statement of authorship JG & MIO designed the study, MIO & US provided materials, JG & SJC performed research and collected data, JG performed zooplankton analysis, SJC performed phytoplankton analysis, JG & MIO performed modeling work, analyzed data output, and wrote the first draft, and all authors contributed substantially to reviews