Habitat engineering by an apex predator generates spatial trophic dynamics across a temporal environmental stress gradient.

Abstract

Ecosystem engineering is a facilitative interaction that generates bottom-up extrinsic variability that may increase species coexistence, particularly along a stress/disturbance gradient. American alligators (Alligator mississippiensis) create and maintain 'alligator ponds' that serve as dry-season refuges for other animals. During seasonal water recession, these ponds present an opportunity to examine predictions of the stress-gradient (SGH) and intermediate disturbance hypotheses (IDH). To test the assumption that engineering would facilitate species coexistence in ponds along a stress gradient (seasonal drying), we modelled fish catch-per-unit-effort (CPUE) in ponds and marshes using a long-term dataset (1997-2022). Stomach contents (n = 1677 from 46 species) and stable isotopes of carbon and nitrogen (n = 3978 representing 91 taxa) from 2018 to 2019 were used to evaluate effects of engineering on trophic dynamics. We quantified diets, trophic niche areas, trophic positions and basal-resource use among habitats and between seasons. As environmental stress increases, we used seasonal changes in trophic niche areas as a proxy for competition to examine SGH and IDH. Across long-term data, fish CPUE increased by a factor of 12 in alligator ponds as the marsh dried. This validates the assumption that ponds are an important dry-season refuge. We found that 73% of diet shifts occurred during the dry season but that diets differed among habitats in only 11% of comparisons. From wet season to dry season, both stomach contents and stable isotopes revealed changes in niche areas. Direction of change depended on trophic guild but was opposite between stable-isotope and stomach-content niches, except for detritivores. Stomach-content niches generally increased suggesting decreased competition in the dry season consistent with existing theory, but stable-isotope niches yielded the opposite. This may result from a temporal mismatch with stomach contents reflecting diets over hours, while stable isotopes integrate diet over weeks. Consumptive effects may have a stronger effect than competition on niche areas over longer time intervals. Overall, our results demonstrated that alligators ameliorated dry-season stress by engineering deep-water habitats and altering food-web dynamics. We propose that ecosystem engineers facilitate coexistence at intermediate values of stress/disturbance consistent with predictions of both the SGH and IDH.