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Ecological changes with minor effect initiate evolution to delayed regime shifts


Regime shifts have been documented in a variety of natural and social systems. These abrupt transitions produce dramatic shifts in the composition and functioning of socioecological systems. Existing theory on ecosystem resilience has only considered regime shifts to be caused by changes in external conditions beyond a tipping point and therefore lacks an evolutionary perspective. In this study, we show how a change in external conditions has little ecological effect and does not push the system beyond a tipping point. The change therefore does not cause an immediate regime shift but instead triggers an evolutionary process that drives a phenotypic trait beyond a tipping point, thereby resulting (after a substantial delay) in a selection-induced regime shift. Our finding draws attention to the fact that regime shifts observed in the present may result from changes in the distant past, and highlights the need for integrating evolutionary dynamics into the theoretical foundation for ecosystem resilience.

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Fig. 1: Potential ecological and evolutionary effects of changing conditions in ecosystems with ASSs.
Fig. 2: Eco-evolutionary dynamics of a population with a habitat switch experiencing a decrease in mortality in the adult habitat.
Fig. 3: Ecological and eco-evolutionary consequences of a decrease in mortality risk.
Fig. 4: Eco-evolutionary dynamics of a population with a habitat switch experiencing a decrease in mortality in the adult habitat for two different regimes of mortality in the nursery habitat.

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Data availability

No data were collected or used in this study.

Code availability

The implementation of the Escalator Boxcar Train numerical method and the PSPM package used to analyse the model can be found in and, respectively.


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This research was supported by the European Research Council under the European Union’s Seventh Framework Programme (FP/2007-2013)/ ERC grant no. 322814.

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Authors and Affiliations



P.C.C.-P. and A.M.deR. designed methodology and gave final approval for publication. P.C.C.-P. conceived the ideas, analysed the results and led the writing of the manuscript. A.M.deR. developed the model formulation and contributed to later versions of the manuscript.

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Correspondence to P. Catalina Chaparro-Pedraza.

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Extended data

Extended Data Fig. 1 Population compositions in the ASSs when the average body size at habitat switch equals 0.25.

The two ASSs correspond to low (solid line) and high biomass levels (dashed line) in the nursery habitat, and to high (solid line) and low biomass levels (dashed line) in the adult habitat. These alternative stable population compositions represent the population structure approximately at time 260 in Fig. 2 (before the regime shift in Fig. 2) and approximately at time 340 in Fig. 2 (after the regime shift in Fig. 2). The regime shift observed in Fig. 2 leads to a decrease in population density in the nursery habitat (green region) and an increase in population density in the adult habitat (blue region), mainly as a consequence of an increase in the density of immature individuals (smaller than the size at maturation). This increase in density of immature individuals in the adult habitat results in increased competition in this habitat that produces a reduction of 32% in the maximum asymptotic body size after the regime shift (reduction from 3.71 to 2.52). Parameter values as in Fig. 2.

Extended Data Fig. 2 Eco-evolutionary effects of trait variation in the population.

a) Ecological and b) evolutionary dynamics before and after a reduction of mortality in the adult habitat (vertical dotted line, from 2 to 1.5). When trait variation is represented with a truncated normal distribution with a minimum and maximum value equal to 80% and 120% (black lines) the regime shift occurs at time 390, whereas when the minimum and maximum value equal to 90% and 110% (grey lines) the regime shift occurs at time 940. Mortality in habitat 1 is 0.8, other parameters as in Table 1 (see Methods).

Extended Data Fig. 3 Long-term stability of the system.

Population biomass and food resource densities in the nursery and adult habitat and selection gradient as a function of body size at habitat switch after a decrease in mortality when the evolutionary endpoint occurs a) in one of the alternative stable ecological equilibrium resulting in a single regime shift (dynamics shown in Fig. 4a) and b) in the unstable equilibrium resulting in repeated delayed regime shifts (dynamics shown in Fig. 4b). Ecologically stable (solid lines) and unstable (dashed lines) equilibrium values are indicated with black lines as well as minimum and maximum densities during oscillatory dynamics (dotted lines). The direction of selection is indicated with thick arrows (orange when negative and blue when positive) and ecological dynamics with double vertical arrows (yellow). The evolutionary endpoint is indicated with a circle (open circle in case it corresponds to an unstable ecological equilibrium, filled circle if it correspond to a stable ecological equilibrium). The direction of selection (bottom plots) is positive at low values of the trait (blue shaded area), negative at high values (pink shaded area) and either negative or positive at intermediate values of the trait (mixed shaded area), depending on which of the two ASSs the population is in. Parameter values as in Fig. 4.

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Chaparro-Pedraza, P.C., de Roos, A.M. Ecological changes with minor effect initiate evolution to delayed regime shifts. Nat Ecol Evol 4, 412–418 (2020).

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