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Habitat heterogeneity and dispersal network structure as drivers of metacommunity dynamics

Davide Bernardi, Alice Doimo, Giorgio Nicoletti, Prajwal Padmanabha, Andrea Rinaldo, Samir Suweis, Sandro Azaele, Amos Maritan

TL;DR

The paper develops a bottom-up framework that derives metacommunity dynamics on arbitrary dispersal networks from detailed individual-based processes, producing an effective dispersal kernel $\hat{K}$ that couples local demography to landscape topology. By exploiting a fast-exploration regime, it reduces to tractable one- or few-dimensional descriptions, yielding exact persistence criteria via the leading Perron–Frobenius eigenvalue $\lambda_{\max}$ and a generalized metapopulation capacity. The authors further incorporate stochasticity to reveal universal finite-size scaling of extinction times and extend the theory to multispecies communities, showing how habitat heterogeneity and network structure enable niche formation and stable coexistence above a critical heterogeneity. The framework also accommodates extensions to diverse ecological processes, providing a versatile and interpretable route to spatially explicit ecological theory with analytical benchmarks and potential data-guided applications.

Abstract

Spatial structure and species interactions jointly shape the dynamics and biodiversity of ecological systems, yet most theoretical models either neglect spatial heterogeneity or sacrifice analytical tractability. Here, we provide a unified microscopic, mechanistic framework for deriving effective metapopulation and metacommunity models from individual-based ecological dynamics on arbitrary dispersal networks. The resulting coarse-grained description features an effective dispersal kernel that encodes both microscopic dynamical parameters and network topology. Based on this framework, we demonstrate exact analytical results for species persistence in both homogeneous and heterogeneous landscapes, including a generalization of the classical concept of metapopulation capacity to non-uniform local extinction rates. Incorporating stochasticity arising from finite carrying capacities, we obtain a reduced one-dimensional description that reveals universal finite-size scaling laws for extinction times and fluctuations. Extending the approach to multiple competing species, we prove that in homogeneous environments monodominance can be avoided only in a fine-tuned, marginally stable coexistence state, and that the classic metapopulation capacity gives only a necessary but not sufficient condition for persistence. We demonstrate that heterogeneous habitats can support stable coexistence, but only above a critical level of heterogeneity. Finally, we outline how additional ecological processes can be systematically incorporated within the same formalism. Together, these results provide analytical benchmarks and a general route for constructing spatially explicit ecological theories based on an interpretable underlying mechanistic foundation.

Habitat heterogeneity and dispersal network structure as drivers of metacommunity dynamics

TL;DR

The paper develops a bottom-up framework that derives metacommunity dynamics on arbitrary dispersal networks from detailed individual-based processes, producing an effective dispersal kernel that couples local demography to landscape topology. By exploiting a fast-exploration regime, it reduces to tractable one- or few-dimensional descriptions, yielding exact persistence criteria via the leading Perron–Frobenius eigenvalue and a generalized metapopulation capacity. The authors further incorporate stochasticity to reveal universal finite-size scaling of extinction times and extend the theory to multispecies communities, showing how habitat heterogeneity and network structure enable niche formation and stable coexistence above a critical heterogeneity. The framework also accommodates extensions to diverse ecological processes, providing a versatile and interpretable route to spatially explicit ecological theory with analytical benchmarks and potential data-guided applications.

Abstract

Spatial structure and species interactions jointly shape the dynamics and biodiversity of ecological systems, yet most theoretical models either neglect spatial heterogeneity or sacrifice analytical tractability. Here, we provide a unified microscopic, mechanistic framework for deriving effective metapopulation and metacommunity models from individual-based ecological dynamics on arbitrary dispersal networks. The resulting coarse-grained description features an effective dispersal kernel that encodes both microscopic dynamical parameters and network topology. Based on this framework, we demonstrate exact analytical results for species persistence in both homogeneous and heterogeneous landscapes, including a generalization of the classical concept of metapopulation capacity to non-uniform local extinction rates. Incorporating stochasticity arising from finite carrying capacities, we obtain a reduced one-dimensional description that reveals universal finite-size scaling laws for extinction times and fluctuations. Extending the approach to multiple competing species, we prove that in homogeneous environments monodominance can be avoided only in a fine-tuned, marginally stable coexistence state, and that the classic metapopulation capacity gives only a necessary but not sufficient condition for persistence. We demonstrate that heterogeneous habitats can support stable coexistence, but only above a critical level of heterogeneity. Finally, we outline how additional ecological processes can be systematically incorporated within the same formalism. Together, these results provide analytical benchmarks and a general route for constructing spatially explicit ecological theories based on an interpretable underlying mechanistic foundation.
Paper Structure (23 sections, 6 theorems, 135 equations, 6 figures, 1 table)

This paper contains 23 sections, 6 theorems, 135 equations, 6 figures, 1 table.

Key Result

Theorem 1

Given the following metapopulation dynamics where $e_i>0$ and the kernel matrix $\hat{K}$ is irreducible. Let $\hat{\mathcal{E}}$ be the matrix with elements $\mathcal{E}_{ij} = e_i \delta_{ij}$, and $\hat{\mathcal{C}}$ be the matrix with elements $\mathcal{C}_{ij} = c_i \delta_{ij}$. Then, the generalized landscape matrix is also irreducible, and has a Perron-Frobenius positive eigenvalue $\lam

Figures (6)

  • Figure 1: (a) Sketch of the model. We consider the dynamics of a settled population, which resides in habitat patches, and of explorers that move along a dispersal network and attempt colonization. In the fast-exploration regime, this leads to an effective metapopulation model with an explicit colonization kernel. (b) The colonization kernel depends manifestly on the topology of the dispersal network. In general, it is not a function of the network distance $d_{ij}$ alone, but rather depends on all paths connecting two habitat patches. (c-e) The metapopulation capacity, which determines a species' survival ability, depends on both the exploration efficiency $f$ and the network topology. In particular, more heterogeneous dispersal networks are more advantageous at intermediate and low $f$, affecting the stationary population and explorers (panels (d) and (e)). Figure adapted from NicPad2023.
  • Figure 2: (a–b) Comparison of stochastic trajectories from the effective quasi-stationary SDE \ref{['eq:stochSDE']} and the full Gillespie simulation in the survival and extinction regimes. (c–d) Finite-size scaling of the survival probability at criticality: (c) raw curves of survival probability as functions of time for different carrying capacities $M$, (d) collapse under the scaling ansatz \ref{['eq:StScaling']}. The initial condition is fixed at $p_0=1/2$. Averages are obtained over $10^7$ numerical realizations of the dynamics in \ref{['eq:stochSDE']}. (e–f) Coefficient of variation of extinction times as a function of initial condition $p_0$ and deviation from criticality $\Delta$ for different values of the local carrying capacity $M$: (e) unscaled, (f) rescaled using finite-size scaling of the first two moments \ref{['eq:scaling']}. Averages are obtained over $10^7$ numerical realizations of the dynamics in \ref{['eq:stochSDE']}. (g–h) Fluctuations in the metastable regime: (g) distribution of settled densities obtained from 350 independent realizations with Gaussian fit; (h) standard deviation $\sigma$ vs. $e/c$, compared with the analytical estimate based on the Ornstein--Uhlenbeck (OU) approximation. The numerical values of $\sigma$ are obtained from Gaussian fits to distributions built from 250 independent realizations. Adapted from ref. DoiNic2025.
  • Figure 3: Visualization of the microscopic model with multiple species competing for space or a common resource leads to an effective metacommunity model. Assuming fast exploration, time-scale separation leads to an effective kernel description (\ref{['eqn:kernel-multispec']}) combined with the effective metacommunity equations reported in \ref{['eqn:model-general']}.
  • Figure 4: (a) Fine-tuned fragile coexistence in homogeneous environments. Time evolution of $p_{\alpha i}$ (the light colored lines correspond to different patches while the dark colored lines correspond to the averages across all patches, $\langle p_\alpha\rangle$), $i=1,\dots, N=25$ and $\alpha=1,2=S$ in a translationally invariant homogeneous environment (the kernel is computed from a 2D regular grid). The two species differ in exploration efficiency, with $f_1 = 1$ and $f_2 = 3$. a) Species’ (average) extinction rates are chosen such that $r_1 = e_1 / z_1 = e_2 / z_2 = r_2=0.002$, with $e_1 = 0.1, e_2=0.15$. These conditions satisfy the hypotheses of \ref{['th:theorem-TI-multispec']}, and the metacommunity dynamics converge to the center manifold. (b) Here, $r_2$ is $1\%$ smaller than in (a), leading to the extinction of species 2. The transient dynamics remain close to the center manifold, which acts as a slow manifold. Since the point on the center manifold reached by the trajectory depends on the initial condition, species 2 exhibits higher abundance during the transient phase, even though it ultimately declines to zero. (c) Convergence of the dynamical evolution of the average $\ev{p_\alpha}$ to the fixed-point solution, with three species (dotted lines) given by \ref{['eqn:Self-consistent-stationary']} for the kernel given by \ref{['eqn:MF-scaling']}. (d) Same, but for the average abundance of each species in each patch, given by \ref{['eqn:SS-general']}. Here, $r_{\alpha i}$ were chosen from a lognormal distribution with mean 1.5 and standard deviation equal to 2.
  • Figure 5: Analytical phase diagram of coexistence regions with mean-field dispersion in heterogeneous habitats. (a) Fraction of coexisting species as a function of the baseline metacommunity fitness $R$ and habitat disorder variance $v^2$. (b) Species localization as a function of the same parameters shows increased localization in the coexisting phase and a strong increase near the global extinction boundary $R=1$. (c) Horizontal slice of panel (b) showing localization as a function of $R$, at a fixed disorder value. A strong increase in the localization near the extinction boundary is observed. (d) Vertical slice of panel (b), localization as a function of the disorder variance for a fixed $R$, which shows the increase of localization when transitioning from monodominance to coexistence, as an indication of niche formation. Numerical simulations are obtained using a log-normal distribution for the disorder. Here, $\Delta_\alpha$ is evenly spaced between $\pm 5/2$, so that $\gamma_\Delta = 2.5$. The theory is computed from numerical integration of \ref{['eqn:moments-p-alpha']}. An expansion for large $S$ only gives an accurate prediction for very large $S$. Adapted from ref. PadNic2024.
  • ...and 1 more figures

Theorems & Definitions (13)

  • Theorem 1
  • Corollary 1
  • Corollary 2
  • Theorem 2
  • Theorem 3
  • Theorem 4
  • proof : Proof of \ref{['th:theorem1']}
  • proof : Proof of \ref{['th:firstcorollary']}
  • proof : Proof of \ref{['th:HO-assumptions']}
  • proof : Proof of \ref{['th:theorem-TI-singlespecies']}
  • ...and 3 more