Coupled energy pathways and the resilience of size-structured food webs
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Abstract
Size-based food-web models, which focus on body size rather than species identity, capture the generalist and transient feeding interactions in most marine ecosystems and are well-supported by data. Here, we develop a size-based model that incorporates dynamic interactions between marine benthic (detritus-based) and pelagic (primary producer based) pathways to investigate how the coupling of these pathways affects food web stability and resilience. All model configurations produced stable steady-state size spectra. Resilience was measured by the return speed obtained from local stability analysis. Return times following large perturbations away from steady-state were also measured. Resilience varied nonlinearly with both predator and detrital coupling, and high resilience came from predators (1) feeding entirely in the slow benthic zone or (2) feeding across the two energy pathways, with most food coming from the fast pelagic pathway. When most of the energy flowed through the pelagic pathway, resilience was positively related to turnover rate. When most of the energy flowed through the benthic pathway, resilience was negatively related to turnover rate. Analysis of the effects of large perturbations revealed that resilience for pelagic ecosystems depended on the nature of the perturbation and the degree of benthic–pelagic coupling. Areas with very little or no benthic–pelagic coupling (e.g. deep seas or highly stratified water columns) may return more quickly following pulses of detrital fallout or primary production but could be much less resilient to the effects of human-induced mortality (harvesting).
Keywords
Benthic–pelagic links Food web dynamics Predator–prey Size spectrum Stability Trophic interactionsNotes
Acknowledgements
This research was funded by the UK Department of Environment, Food and Rural Affairs project M10-01, Cefas Seedcorn project DP222 and EU IMAGE (FP6 contract-044227). JLB was supported by the Visitors Programme at the NERC Centre for Population Biology, Imperial College, Silwood Park Campus, UK. RL was supported by a Killam Visiting Professorship at the University of Calgary Canada and an Erskine Fellowship at the University of Canterbury, New Zealand. We thank participants of the European Science Foundation Network “Body-size and Ecosystem Dynamics: Integrating pure and applied approaches from aquatic and terrestrial ecology to support an ecosystem approach (SIZEMIC)” for helpful discussions especially Anje-Margriet Neutel. We are grateful for the insightful comments provided by two anonymous referees.
Supplementary material
The effects of different levels of plankton density and detrital input on turnover rates and dominant eigenvalue results for comparison (GIF 42 kb)
The effects of preferred predator–prey mass ratio and the width of the predator–prey mass ratio on turnover rates and dominant eigenvalue results (GIF 41 kb)
Bottom-up primary producer perturbations. Relative biomass trajectories over time and corresponding resilience as measured by the reciprocal of return time 1/T R for each level of predator preference for benthic prey ω B and each component of the ecosystem: a, b pelagic predators, c, d benthic detritivores and e, f detritus following a 30-day 50% increase in primary producer density. In all plots, the black line corresponds to predators feeding only within the pelagic community (no coupling, ω B = 0). The grey lines show varying the strengths of coupling towards the benthic pathway (the lightest line corresponds to predators only feeding on benthic prey, ω B = 1) (GIF 48 kb)
Top-down predator harvesting perturbations. Relative biomass trajectories over time and corresponding resilience as measured by the reciprocal of return time 1/T R for each level of predator preference for benthic prey ω B and each component of the ecosystem: a, b pelagic predators, c, d benthic detritivores and e, f detritus following a 30-day 50% decrease in pelagic predators >10 g. In all plots, the black line corresponds to predators feeding within the pelagic community (no coupling, ω B = 0). The grey lines show varying the strengths of coupling towards the benthic pathway (the lightest line corresponds to predators only feeding on benthic prey, ω B = 1) (GIF 52 kb)
Bottom-up detritus perturbations. Relative biomass trajectories over time and corresponding resilience as measured by the reciprocal of return time 1/T R for each level of predator preference for benthic prey ω B and each component of the ecosystem: a, b pelagic predators, c, d benthic detritivores and e, f detritus following a 30-day 50% increase in detritus density. In all plots, the black line corresponds to predators feeding only within the pelagic community (no coupling, ω B = 0). The grey lines show varying the strengths of coupling towards the benthic pathway (the lightest line corresponds to predators only feeding on benthic prey, ω B = 1) (GIF 45 kb)
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