Opinion
Emergent Properties Delineate Marine Ecosystem Perturbation and Recovery

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Despite ongoing debate, we assert that there are consistent, fundamental features of marine ecosystems.

Key fundamental features are emergent properties of marine ecosystems that include sigmoidal cumulative biomass–trophic-level and ‘hockey-stick’ cumulative production–biomass curves.

These consistent, theoretically supported patterns occur in over 120 marine ecosystems.

A developing cumulative trophic theory builds upon past advances, widespread empirical support, and confirmation of simple predictions with observed responses.

Cumulative curve parameters can help delineate when marine ecosystems are perturbed or recovered, integrating across a range of stressors and response mechanisms.

Whether there are common and emergent patterns from marine ecosystems remains an important question because marine ecosystems provide billions of dollars of ecosystem services to the global community, but face many perturbations with significant consequences. Here, we develop cumulative trophic patterns for marine ecosystems, featuring sigmoidal cumulative biomass (cumB)–trophic level (TL) and ‘hockey-stick’ production (cumP)–cumB curves. The patterns have a trophodynamic theoretical basis and capitalize on emergent, fundamental, and invariant features of marine ecosystems. These patterns have strong global support, being observed in over 120 marine ecosystems. Parameters from these curves elucidate the direction and magnitude of marine ecosystem perturbation or recovery; if biomass and productivity can be monitored effectively over time, such relations may prove to be broadly useful. Curve parameters are proposed as possible ecosystem thresholds, perhaps to better manage the marine ecosystems of the world.

Section snippets

Are there Fundamental Patterns of Marine Ecosystems?

The question remains whether there are common patterns from ecosystems that arise from underlying, fundamental ecosystem processes [1]. Several have been proposed [2] and often such proposed patterns are emergent properties of ecosystems 3, 4. Yet, whether such emergent patterns are germane for marine ecosystems remains unclear due to the distinctiveness of the marine environment 5, 6 (Box 1). Detecting any such common emergent patterns in marine ecosystems is useful to understand the effects

Indicators and Patterns

Indicators are needed to assess the degree of perturbation or recovery of marine ecosystems 11, 16. Addressing elements of marine ecosystem response to perturbations has typically emphasized a population or habitat level that disregards community- or systemic-level compensation and dynamics. However, aggregative or system-level patterns tend to be more robust and integrative 17, 18. Further, if changes in aggregative or system-level patterns are persistent, they represent bigger concerns than

Pattern Detection

The cumB–TL plots exhibit the sigmoidal shape as expected (Figure 2). Even the widest percentiles from empirical data encompassing at least 95% of all upwelling, continental shelf, reef, and open ocean ecosystems show relatively limited variance from the median pattern. Estuaries also exhibit this pattern, but have a wider range of variance near the base of the curve (Figure 2E). There is a distinct and repeatable sigmoidal pattern across TL for cumB, with a noticeable increase usually between

Consistency of Perturbation Effects and Thresholds

Time series from over 20 marine ecosystems demonstrate that curve parameters exhibit notable responses to perturbation (Figure 4A–D; Box 2). For instance, the Black Sea is a system known to have experienced significant and major perturbations related to fishing pressure and an invasive jellyfish outburst 36, 37, and these are reflected in the time series of inflection points (Figure 4A,C) and slopes (Figure 4B,D), which exhibit high variability and an overall decline in both parameters. Other

Explanation Based on Trophic Theory

Basic ecosystem theory clearly emphasizes an energetic context within which ecological systems function. Thermodynamic constraints limit the overall productivity of a system, reflected directly in the observation that transfer efficiencies, or the efficiency that the energy in an ecosystem is transferred from TL to succeeding TL, are always <1, such that there is a decline in production rate with increasing TLs 40, 41, 42. The TL concept posited by Lindeman and others, which summarizes the

Delineating Thresholds of Perturbation and Recovery

From this theoretical background, widespread empirical support, and common values of curve parameters, can we delineate when a marine ecosystem is in an abnormal state, indicative of probable perturbation (Box 4)? Or, conversely, can we delineate when a marine ecosystem has ‘recovered’? As noted, changes in the slope and inflection points of the cumB–TL and cumP–cumB curves can be tracked over time and related to particular pressures (Figure 4A–D) [50]. Similar to population- or habitat-level

Acknowledgments

This work was conceptualized and conducted as part of visiting scientist exchanges between F.P. and J.L. via support from the University of Ca’Foscari and NOAA, a visiting scientist exchange via a Frohlich Fellowship (J.L.) and support from CSIRO and NOAA between J.L. and E.F., NOAA support for J.L. to visit S.L. and C.S., CAMEO support for MENU2, and in conjunction with numerous ICES working groups in which several of the authors participated. We thank Selina Heppel, Eva Plaganyi-Lloyd, Tim

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