Department Aquatic Ecology

Nonliving resources frequently flow across ecosystem boundaries, which can yield networks of spatially coupled ecosystems. Yet the significance of resource flows for ecosystem function has predominantly been understood by studying two or a few coupled ecosystems, overlooking the broader resource flow network and its spatial structure. Here, we investigate how the spatial structure of larger resource flow networks influences ecosystem function at metaecosystem scales by analyzing metaecosystem models with homogeneously versus heterogeneously distributed resource flow networks but otherwise identical characteristics. We show that metaecosystem function can differ strongly between metaecosystems with contrasting resource flow networks. Differences in function generally arise through the scaling up of nonlinear local processes interacting with spatial variation in local dynamics, the latter of which is influenced by network structure. However, we find that neither network structure guarantees the greatest metaecosystem function. Rather, biotic (organism traits) and abiotic (resource flow rates) properties interact with network structure to determine which yields greater metaecosystem function. Our findings suggest that the spatial structure of resource flow networks coupling ecosystems can be a driver of ecosystem function at landscape scales. Furthermore, our study demonstrates how modifications to the structural, biotic, or abiotic properties of metaecosystem networks can have nontrivial large-scale effects on ecosystem function.

Ecosystems are widely interconnected by spatial flows of material, but the overall importance of these flows relative to local ecosystem functioning remains unclear. Here we provide a quantitative synthesis on spatial flows of carbon connecting ecosystems worldwide. Cross-ecosystem flows range over eight orders of magnitude, bringing between 10⁻³ and 10⁵ gC m⁻² year⁻¹ to recipient ecosystems. Magnitudes are similar to local fluxes in freshwater and benthic ecosystems, but two to three orders of magnitude lower in terrestrial systems, demonstrating different dependencies on spatial flows among ecosystem types. The strong spatial couplings also indicate that ecosystems are vulnerable to alterations of cross-ecosystem flows. Thus, a reconsideration of ecosystem functioning, including a spatial perspective, is urgently needed.

The meta-ecosystem framework demonstrates the significance of among-ecosystem spatial flows for ecosystem dynamics and has fostered a rich body of theory. The high level of abstraction of the models, however, impedes applications to empirical systems. We argue that further understanding of spatial dynamics in natural systems strongly depends on dense exchanges between field and theory. From empiricists, more and specific quantifications of spatial flows are needed, defined by the major categories of organismal movement (dispersal, foraging, life-cycle, and migration). In parallel, the theoretical framework must account for the distinct spatial scales at which these naturally common spatial flows occur. Integrating all levels of spatial connections among landscape elements will upgrade and unify landscape and meta-ecosystem ecology into a single framework for spatial ecology.