A unified meta‐ecosystem dynamics model: Integrating herbivore‐plant subwebs with the intermittent upwelling hypothesis

Abstract

AbstractDetermining the relative influence of biotic and abiotic processes in structuring communities at local to large spatial scales is best understood using a biogeographic comparative‐experimental approach. Using this approach, previous work suggests that intertidal community dynamics (top‐down and bottom‐up effects) vary unimodally along an upwelling‐based productivity gradient, termed the Intermittent Upwelling Hypothesis (IUH). Evidence consistent with the IUH comes from the sessile invertebrate/predator (SIP) subweb in certain rocky intertidal communities, but whether this pattern extends to macrophyte/herbivore (MH) subwebs is unknown. Here we ask: Are MH subwebs also structured as predicted by the IUH? What is the relative importance of herbivory and predation in structuring these communities? Under what conditions do ecological subsidies like nutrients or propagule production drive community dynamics? And are omnivorous interactions important? We hypothesize that MH subwebs are driven by a new construct, the Grazing‐Weakening Hypothesis (GWH), which states that MH interactions weaken monotonically with increasing nutrients, with strong (weak) herbivory and low (high) macrophyte productivity at low (high) nutrients. We explored local‐to‐large spatial scale dynamics of both subwebs using a biogeographic comparative‐experimental factorial field experiment testing joint and separate effects of herbivores and predators between two continents. Experiments at 10 sites ranging from persistent upwelling to persistent downwelling regimes ran for 26–29 months in Oregon and California, and New Zealand (NZ) South Island. For the MH subweb, results were consistent with the GWH: herbivory declined and macrophytes increased with increasing nutrients. As expected, results for the SIP subweb were consistent with the IUH: predator effect size was unimodally related to upwelling. Overall, herbivory explained more variation in community structure than did predation, especially in NZ. Omnivory was weak, sessile invertebrates outcompeted macrophytes, and ocean‐driven subsidies provided the basic template driving ecosystem dynamics. We propose a unified meta‐ecosystem dynamics model combining MH and SIP results: with increased upwelling, sessile invertebrates and underlying dynamics vary unimodally (as in the IUH), while herbivory decreases and macrophytes generally increase. While this model was based on research in temperate ecosystems varying in upwelling regime, its wider applicability remains to be tested.