Abstract
Large, complex networks of ecological interactions with random structure tend invariably to instability. This mathematical relationship between complexity and local stability ignited a debate that has populated ecological literature for more than three decades. Here we show that, when species interact as predators and prey, systems as complex as the ones observed in nature can still be stable. Moreover, stability is highly robust to perturbations of interaction strength, and is largely a property of structure driven by predator–prey loops with the stability of these small modules cascading into that of the whole network. These results apply to empirical food webs and models that mimic the structure of natural systems as well. These findings are also robust to the inclusion of other types of ecological links, such as mutualism and interference competition, as long as consumer–resource interactions predominate. These considerations underscore the influence of food web structure on ecological dynamics and challenge the current view of interaction strength and long cycles as main drivers of stability in natural communities.
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Acknowledgements
We thank D. Alonso, A. Bodini, J.M. Dambacher and A. P. Dobson for stimulating discussions. This work was supported by a Centennial Fellowship by the J. S. McDonnell Foundation to M.P.
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Supplementary Fig. 1
Number of right-signed terms in the characteristic polynomial of the system represented in Figure 3. (a) percentage of terms with the right sign for each coefficient of the characteristic polynomial. (b) Sign difference: number of terms with the right sign - number of terms with the wrong sign (EPS 5.85 kb)
Supplementary Fig. 2a
Percentage of food webs, built with the “cascade model” (left) and the “niche” model (right), that are stable given C and S (EPS 29.6 kb)
Supplementary Fig. 2b
(EPS 26.2 kb)
Supplementary Fig. 3
Comparison between random matrices (left) and random predator-prey matrices (right) when interaction strengths are drawn from a lognormal (μ = 0, σ = 1) distribution. (a-b) Percentage of stable systems for each pair (C, S). (c-d) Average percentage of eigenvalues with negative real part. (e-f) Percentage of stable matrices produced by randomizing 100 times the coefficients of each stable matrix in (a-b). The number of matrices concurring to form this average is not fixed, as in the previous graphs, depending on the number of stable matrices (a-b). The dots represent the values of 10 empirical food webs (EPS 111 kb)
Supplementary Fig. 4
Comparison between random matrices (left) and random predator-prey matrices (right) when interaction strengths are drawn from an uniform (U[0,1]) distribution. (a-b) Percentage of stable systems for each pair (C, S). (c-d) Average percentage of eigenvalues with negative real part. (e-f) Percentage of stable matrices produced by randomizing 100 times the coefficients of each stable matrix in (a-b). The number of matrices concurring to form this average is not fixed, as in the previous graphs, depending on the number of stable matrices (a-b). The dots represent the values of 10 empirical food webs (EPS 117 kb)
Supplementary Fig. 5
Percentage of stable systems when 10% of the connectance is represented by double (mutualism or interference) connections (top). In the bottom graph we see the effects on stability of increasing the percentage of double interactions. The solid line represents the effects on predator-prey systems and the dashed line the effects on random networks (EPS 38.4 kb)
Supplementary Fig. 6
Effects on stability of increasing the percentage of null (left) or positive (right) coefficients on the diagonal. The solid line represents the effects on predator-prey systems and the dashed line the effects on random networks. In the right part of the graph, red dashed and solid lines represent the fraction of systems with negative trace (EPS 6.76 kb)
Supplementary Fig. 7
Comparison between random matrices (left) and random predator-prey matrices (right) when diagonal elements are positive with 5% probability and null with 5% probability. The other diagonal coefficients are negative. (a-b) Percentage of stable systems for each pair (C, S). (c-d) Average percentage of eigenvalues with negative real part. (e-f) Percentage of stable matrices produced by randomizing 100 times the coefficients of each stable matrix in (a-b). The number of matrices concurring to form this average is not fixed, as in the previous graphs, depending on the number of stable matrices (a-b). The dots represent the values of 10 empirical food webs (EPS 110 kb)
Supplementary Fig. 8
Effects of the distribution range on the probability of stability illustrating that the fraction of weak interactions has no effect on the probability of stability. We built 500 networks of the random (red) and predator-prey (blue) type. Each network contains 25 species and has connectance 0.1. The coefficient strengths are taken from a χ 2 distribution with k degrees of freedom. The coefficient signs are chosen as in the main text. Changing the degree of freedom reduces the fraction of weak interactions and moves the mean of the distribution toward higher values (EPS 8.71 kb)
Supplementary Table 1
Transformation of food webs into community matrices (PDF 39.5 kb)
Supplementary Table 2
Numerical computation of the number of terms with the right sign in a predator-prey matrix that is completely connected (PDF 33.6 kb)
12080_2007_7_MOESM11_ESM.pdf
Network structure, predator-prey modules, and stability in large food webs: Electronic Supplementary Material (ESM) (PDF 116 kb)
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Allesina, S., Pascual, M. Network structure, predator–prey modules, and stability in large food webs. Theor Ecol 1, 55–64 (2008). https://doi.org/10.1007/s12080-007-0007-8
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DOI: https://doi.org/10.1007/s12080-007-0007-8