bioRxivEcologyPreprint
Ecological networks balance connectivity with flexibility to perturbations
Àlex Giménez‐RomeroiD, Daniel OróiD, Jordi BascompteiD, Meritxell GenovartiD
Abstract
Food webs become progressively sparser as species richness increases, yet the number of interactions grows faster than linearly, placing communities between the limiting expectations of constant degree and constant connectance. Similar scaling has also been documented in other interaction networks, but whether they share a common architectural regime, and what general principle could generate it, remain unresolved. Here, we show that 1,533 ecological networks spanning different interaction types broadly converge on an intermediate link--species scaling regime, with an overall exponent of 1.43. We interpret this pattern through a thermodynamic trade-off in which increasing network connectivity enhances signal propagation while reducing the diversity of independent modes contributing to the network's response to perturbations. Optimizing this trade-off yields a scaling exponent of 1.45, closely matching the empirical estimate, while randomly adding or removing interactions generally reduces thermodynamic efficiency in empirical networks. These results support a propagation--response-diversity trade-off as a plausible systems-level principle underlying how ecological network complexity grows with diversity.
In-depth summary
Claude Opus 5.5 · xhigh effort · 8 Nov 2026 · from the full text of version 2 (31 Oct 2026) · CC BY 4.0
Across 1,569 food-web, mutualistic and host–parasite networks, connectance falls as S^−0.65 while links per species rise; a diffusion-efficiency metric places networks near a local optimum.
Evidence that connectance declines with species richness, a structural prediction of May-type stability theory, has come mostly from food webs, and it sits uneasily beside the finding that total and per-species interactions grow with diversity. This study tests the scaling across interaction types and reads the combined pattern through a thermodynamic framework for diffusion on networks that weighs signal propagation against response diversity.
Key results
ShownConnectance falls as a power of species richness in the pooled data and within each interaction class. The decline is shallower than the S^−1 bound derived from random-matrix stability and feasibility theory.
Pooled fit C ∼ S^−0.65 (95% CI [−0.67, −0.64], R^2 = 0.77, p < 0.001) on the full dataset (1,569 networks in S3; Fig. 2b legend says ∼1300); food web, host–parasite, pollination and seed-dispersal fits agree within 95% CIs, as do bipartite vs non-bipartite subsets; random-matrix baseline exponent is −1. · A broad architectural tendency across ecological…; Figure legends
ShownA fixed number of partners per species does not explain the sparsity. Mean degree rises with richness, so total links grow superlinearly, between the constant-degree (L ∼ S) and fully connected (L ∼ S^2) limits.
⟨k⟩ ∼ S^0.43 (R^2 = 0.54, p < 0.001), with the same conclusion within interaction types; L ∼ S^γ with γ = 1.43 [1.41, 1.45] (R^2 = 0.93, p < 0.001); degree distributions are heavy-tailed and fitted by truncated power laws. · A broad architectural tendency across ecological…; Figure legends
SuggestedThe efficiency metric weighs propagation gain against loss of response diversity. By this metric, observed networks sit near a local maximum: randomly adding or deleting links mostly lowers efficiency, and a mean-field estimate of the optimal link-scaling exponent is close to the fitted one.
Δη = ηperturbed − ηemp after adding or removing 1% of links is mostly negative (Fig. 3d, at τdiff = 1/λ2); changes of 1% to 10% in either direction also give negative Δη (100 perturbed networks per case); mean ⟨γopt⟩ = 1.42 vs fitted γ = 1.43 [1.41, 1.45]. The authors call this compatibility, not prediction. · Diffusion dynamics and the structural balance of…; Interaction perturbation experiments; Optimal scaling from thermodynamic efficiency; Figure legends
ShownDegree heterogeneity explains a substantial fraction of the diffusion behaviour. Random graphs matched on size and density fall short of empirical efficiency, whereas degree-preserving rewiring closely tracks it over most of the diffusion range.
Erdős–Rényi nulls preserving size and density show systematically lower efficiency at intermediate and large τ in every interaction type; configuration-model nulls preserving degree sequence follow empirical η(τ) closely over most of the range, with small but systematic deviations in several interaction types, particularly mutualistic systems (higher empirical efficiency at intermediate τ); 100 nulls per network. · Diffusion dynamics and the structural balance of…; Thermodynamic efficiency of signal propagation; Null network models
ShownMutualistic networks, especially plant–pollinator and plant–ant, beat degree-preserving null efficiency more often than antagonistic networks. Networks with this excess also tend to have shorter path lengths.
A higher fraction of mutualistic networks have ΔηCM > 0 (binomial 95% CIs, Fig. 4a), while antagonistic networks stay near CM expectations; several mutualistic classes keep positive deviations under a fixed-marginal Curveball null; ΔηCM correlates with the global-efficiency deviation ΔεCM (R^2 = 0.41, p < 0.001). The text gives no fractions. · Mutualistic networks further exploit network…; Null network models; Global efficiency; Figure legends
Methods, data and reproducibility
Compiled 1,800 interaction networks from Web of Life and Mangal (1,569 after filtering), binarized them and fitted power laws of connectance, mean degree and link count against richness. For 1,304 networks with S > 5, computed Laplacian-diffusion thermodynamic efficiency η(τ) and compared it with Erdős–Rényi, configuration-model and Curveball nulls and with networks having 1–10% of links added or removed.
- Ecological interaction networks (food webs, host–parasite, pollination, seed dispersal, plant–ant, anemone–fish) from the Web of Life and Mangal databases, converted to unweighted adjacency matrices · multiple species · n = 1,569 after filtering (of 1,800); 1,304 with S > 5 for efficiency
- Species richness and connectance values for communities analysed by Jacquet et al., used only for comparison in (S, C) space · n = 116 communities
- Processed networks in GraphML format with derived structural and thermodynamic-efficiency metrics, deposited on Zenodo · multiple species · 10.5281/zenodo.20670601
- Methods & tools
- Web of Life database, Mangal database, Mangal.jl, NetworkX, graph Laplacian diffusion / thermodynamic efficiency (free energy, von Neumann entropy), Erdős–Rényi null model, configuration model, Curveball algorithm
- Code
- https://github.com/agimenezromero/Ecological-networks-balance-connectivity-with-flexibility-to-perturbationshttps://doi.org/10.5281/zenodo.20670601
- Data & accessions
- https://www.web-of-life.es/https://mangal.io/doc/https://mangal.io/doc/jl/https://doi.org/10.5281/zenodo.20670601
- Reproducibility gaps
- The text states no software versions, dependency files or compute requirements. The raw networks come from the third-party Web of Life and Mangal databases.
Limitations (6)
- The analysis uses unweighted topology only, so it leaves out interaction strengths, which affect ecological dynamics and stability. · stated by the authors
- Directed networks (e.g., trophic) were symmetrized, so the efficiency results describe an undirected structural backbone and lose interaction direction and sign. · stated by the authors
- Diffusion is a minimal, roughly linear stand-in for how perturbations spread. It does not test dynamical stability directly and misses nonlinear community dynamics. · stated by the authors
- The networks were built with different methods, sampling effort and completeness. Coverage of regions, habitats, interaction types and microbial vs multicellular communities is uneven, and most networks are snapshots without temporal context. · stated by the authors
- The optimal-exponent expression is a heuristic mean-field approximation. Agreement between γopt and the fitted exponent shows compatibility, not a uniquely identified mechanism. · stated by the authors
- The paper gives different network counts: about 1,500 in the abstract, 1,569 in the results, ∼1300 in the Fig. 2b legend and 1,304 for efficiency. It is unclear exactly which set underlies each fit. · noted in this summary
Read this if you study ecological network structure or complexity–stability theory and want a test of connectance scaling across interaction types, with a diffusion-based efficiency interpretation and open code.
- Citations
- 1
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