Abstract
The metaecosystem framework has been proposed to conceptualize the interactive effects of dispersal and resource flows on the structure and functioning of communities in a heterogeneous environment. Here, we model a two-patch metaecosystem where two species with a trade-off in resource requirements compete for two limiting resources—generalizing the so-called gradostat experimental setup. We study the competition outcome in dependence of resource heterogeneity and between-patch diffusion for different combinations of resource supply ratios. Our numerical simulations show that community composition and local and regional diversity are determined by the interplay of resource heterogeneity, resource supply stoichiometry, and diffusion rate. High resource heterogeneity increases regional diversity, with species coexisting due to spatial segregation, whereas low resource heterogeneity favors local diversity, as species coexist mainly by local resource partitioning. Regional diversity averaged across a gradient of resource ratios decreases monotonically with diffusion rate, while local diversity follows a unimodal dependency. However, these dependencies become bimodal for high resource heterogeneity because various bistable states occur at intermediate diffusion rates. We identify three kinds of bistable states with species priority effect: (i) bistability between the dominance of one or the other competitor, (ii) bistability between one species dominance or species coexistence, and (iii) two alternative coexistence regimes differing in species-relative abundances. Most bistable states appear at high resource levels when biomass fluxes strongly interact with resource fluxes. Our analysis provides new insights for the potential effects of metaecosystem dynamics on biodiversity patterns.
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Acknowledgments
I.T. acknowledges support from Deutsche Forschungsgemeinschaft (DFG), project GZ: RY 143/1-1. ABR acknowledges the Lower Saxony Ministry for Science and Culture, project POSER.
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Fig. S1
Representation of the competitors by their R∗ values in the resource supplies plane. The dashed lines are the zero net growth isoclines (ZNGI) of species 1 (blue) and species 2 (red). The diagonal dotted line from the top-left to the bottom-right is the set of 60 resource supply ratios used to calculate species 2 relative biomass in Fig. 4, and aggregated regional and local diversity in Fig. 6. The red circles indicate the three specific resource supply ratios used for calculation of the biodiversity in Fig. 7. (PDF 100 kb)
Figure S2
The full diagram of competition outcomes under strong heterogeneous conditions without between-patch diffusion. Numbers indicate the competition outcomes in each patch (patch A/patch B): 0 means no species survives, 1 means species 1 wins, 2 means that species 2 wins, and 1,2 means local coexistence. The regional coexistence occurs when species are present at different patches, e.g. 2/1. The black dashed rectangular indicates the part of the full diagram shown in Fig. 2B. (PDF 117 kb)
Figure S3
Species biomass distribution for various competition outcomes. (Left panels) The competition outcomes under strongly heterogeneous conditions for low (k = 0.02), intermediate (k = 0.1) and high (k = 0.45) diffusion rates (same as Fig. 3B, D, F). (Right panels) the species biomasses in each patch for the various resource combinations, marked by roman numeral in the left panes, and different initial conditions (1st column: when species 1 is the resident and species 2 invades, 2nd column: the opposite invasion sequence). (PDF 183 kb)
Figure S4
Effects of resource heterogeneity and diffusion rate on (A) Community R∗ values and (B) total average biomass. The community R∗ was calculated as the minimal resource supply levels of resources (at the balanced resource ratio) needed for survival of at least one of the competitors. The total biomass was the averaged along a gradient of resource supply ratios along the diagonal from the top-left to down-right of resource planes (Fig. S1). (PDF 161 kb)
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Tsakalakis, I., Blasius, B. & Ryabov, A. Resource competition and species coexistence in a two-patch metaecosystem model. Theor Ecol 13, 209–221 (2020). https://doi.org/10.1007/s12080-019-00442-w
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DOI: https://doi.org/10.1007/s12080-019-00442-w