Stoichiometry and Life-History Interact to Determine the Magnitude of Cross-Ecosystem Element and Biomass Fluxes

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Abstract
Ecosystems are linked through the transfer of materials and energy. Studies examining material fluxes across habitat boundaries frequently quantify unidirectional flows of nutrients and energy. However, material fluxes can be multidirectional, and we lack a conceptual framework to describe how their quantity and stoichiometry influence the net transfer of individual elements between ecosystems. Here we develop a zero net transfer isocline (ZNTI) framework that integrates the relative mass and stoichiometry of fluxes into and out of an ecosystem. We then use case studies with amphibians and salmon to elucidate how life history, ontogenetic shifts in stoichiometry, and trophic interactions shape relative fluxes of nutrients between aquatic and terrestrial ecosystems. Because they increase in both size and Ca content from ova to metamorphs, amphibian life histories strongly bias them towards net Ca export into the terrestrial environment. Because amphibian biomass, C, P, and Ca zero net transfer isoclines do not overlap, there is no value of survivorship where the net flux of biomass, C, P, and Ca are simultaneously balanced between terrestrial and aquatic habitats. The degree of iteroparity and semelparity in salmon strongly affects both the magnitude of net biomass and P flux between riverine and marine environments. While the net direction of biomass flux generally remains strongly biased towards import into the riverine system, net P flux can reach net export into the marine environment because of increasing adult breeding survival leading to reduced mass and %P of what they deposit in rivers (e.g., ova versus whole carcasses). These examples highlight how ontogenetic shifts in body size and stoichiometry result in asymmetric fluxes of elements and biomass that can lead to simultaneous net imports and exports of different elements within the same system. Furthermore, they demonstrate how changes in life-history characteristics and stage-specific survivorship can lead to changes in net elemental transport between ecosystems.

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