Testing ecological models
Which ecological models remain compatible with observations when much of the system is hidden? We develop tests that can reject candidate models without observing every species or driver.
Covariance criteria link observable population gains and losses through necessary conditions for a model. In a selected set of 172 population time series, the classic logistic model passed the covariance test in only 10 cases. Comparing 18 predator–prey series without major rapid evolution with 13 where it was observed revealed deviations from the baseline Lotka–Volterra form in predator dynamics. Under the test’s assumptions, a violated condition rules out the candidate model; passing does not prove it correct. Observation quality and the sampling needed to estimate the quantities remain important.
A complementary preprint on label invariance asks whether predictions change when identical individuals are split into arbitrary groups. Together with our work comparing theoretical frameworks, these projects ask both how to test a model and which scientific job it is built to do.
Key references
9
- Rigorous validation of ecological models against empirical time series. Nature Ecology & Evolution (2025).
- Toward a Conceptual Commons for Theory in Community Ecology. Annual Review of Ecology, Evolution, and Systematics (2026).
- What Have We Learned From Empirical Applications of Modern Coexistence Theory? Ecology Letters (2026).
- The Equilibrium Conundrum. Ecology Letters (2025).
- Bridging parametric and nonparametric measures of species interactions unveils new insights of non‐equilibrium dynamics. Oikos (2021).
- Disentangling the effects of external perturbations on coexistence and priority effects. Journal of Ecology (2020).
- On the Consequences of the Interdependence of Stabilizing and Equalizing Mechanisms. The American Naturalist (2019).
- Beware z‐scores. Journal of Animal Ecology (2019).
- Towards a Probabilistic Understanding About the Context-Dependency of Species Interactions. Trends in Ecology & Evolution (2020).
In the pipeline
4
- Label invariance: a diagnostic for ecological models. Submitted; public preprint. Gibbs T et al.
- When does heterogeneity average out in coarse-grained ecological communities?. Submitted. Luo M et al.
- A trilemma of ecological complexity. In preparation. Metz T et al.
- Lotka–Volterra at 100: a more honest starting point. In preparation. Song C
Coexistence of many species
How do many species coexist, and how does environmental change alter their opportunities to persist? We connect multispecies theory with interactions across trophic levels, environmental variation and empirical estimates of species interactions.
Our structural approaches measure the range of model conditions compatible with coexistence. This is a way to assess coexistence opportunities under specified dynamics; it is distinct from directly observing indefinite persistence in a field community. Published studies examine network structure and species abundance, phenology, individual variation, soil microbes and multitrophic interactions.
A submitted manuscript investigates whether finite within-species budgets can support coexistence without between-species trade-offs. A second examines constraints on annual-plant interactions across variable rainfall. Work on fluctuating environments asks how the timing of conditions changes the conclusions drawn from averages.
Key references
16
- Trophic tug‐of‐war: Coexistence mechanisms within and across trophic levels. Ecology Letters (2024).
- What Have We Learned From Empirical Applications of Modern Coexistence Theory? Ecology Letters (2026).
- On the Consequences of the Interdependence of Stabilizing and Equalizing Mechanisms. The American Naturalist (2019).
- Disentangling the effects of external perturbations on coexistence and priority effects. Journal of Ecology (2020).
- Generalism drives abundance: A computational causal discovery approach. PLOS Computational Biology (2022).
- Telling ecological networks apart by their structure: An environment-dependent approach. PLOS Computational Biology (2020).
- Mean species responses predict effects of environmental change on coexistence. Ecology Letters (2023).
- Soil microbial influences over coexistence potential in multispecies plant communities in a subtropical forest. Ecology (2024).
- Synthesizing the effects of individual‐level variation on coexistence. Ecological Monographs (2022).
- Understanding the emergence of contingent and deterministic exclusion in multispecies communities. Ecology Letters (2021).
- Structural stability: Concepts, methods, and applications. Biodiversity Science (2020).
- Rethinking the importance of the structure of ecological networks under an environment‐dependent framework. Ecology and Evolution (2018).
- A guideline to study the feasibility domain of multi-trophic and changing ecological communities. Journal of Theoretical Biology (2018).
- Will a small randomly assembled community be feasible and stable? Ecology (2018).
- Structural stability as a consistent predictor of phenological events. Proceedings of the Royal Society B: Biological Sciences (2018).
- Why are some plant–pollinator networks more nested than others? Journal of Animal Ecology (2017).
In the pipeline
4
- Multispecies coexistence does not require trade-offs between species. Submitted. Max D et al.
- The standard plant–soil feedback model underpredicts coexistence. Submitted. Max D et al.
- Species interactions are internally constrained despite large climatic variability. Submitted; public preprint. Picó S et al.
- Community response and the temporal niche in variable environments. In preparation. Barabás G et al.
Assembly and priority effects
Does the order in which species arrive determine the community that forms? We use topology, graph theory and information theory to distinguish history-dependent assembly outcomes from outcomes fixed by the system and its conditions.
The possible priority effects multiply super-exponentially with species number, yet our framework reduces them to four basic types. Coexistence holes, gaps among possible communities, make assembly and disassembly discontinuous. Assembly graphs connect assemblages through species introductions and the transitions that follow. These representations help make questions about arrival order explicit and identify which outcomes can be reached.
For experimental use, an introduction sequence and the community it produces define a candidate transition. Repeating sequences can reveal variation in outcomes; the representation must be interpreted alongside the observation timescale and community definition. Our published assembly-graph review connects theoretical approaches with measurable components. Ongoing microbial work examines how antibiotic-resistance evolution affects assembly predictability.
Key references
6
- Assembly Graph as the Rosetta Stone of Ecological Assembly. Environmental Microbiology (2025).
- Coexistence holes characterize the assembly and disassembly of multispecies systems. Nature Ecology & Evolution (2021).
- Untangling the complexity of priority effects in multispecies communities. Ecology Letters (2021).
- Understanding the emergence of contingent and deterministic exclusion in multispecies communities. Ecology Letters (2021).
- Structural changes within trophic levels are constrained by within‐family assembly rules at lower trophic levels. Ecology Letters (2018).
- Disentangling the effects of external perturbations on coexistence and priority effects. Journal of Ecology (2020).
In the pipeline
1
- Antibiotic resistance evolution and the predictability of bacterial community assembly. In preparation. Hogle SL et al.
Geometry and physics of communities
What do geometry and physics reveal about community responses to change? We use these tools to pose ecological questions about sustained pressures, temporal dynamics and differences among assemblages.
A submitted manuscript on structural sensitivity studies which directions of sustained environmental change a community amplifies or absorbs. A public preprint on time irreversibility measures how population time series differ when played forward and backward, with applications involving evolution, nutrient supply and temperature.
Our published geometric approach to beta diversity measures assemblage structure from a different perspective than conventional pairwise dissimilarity. The useful choice depends on the ecological question: a complementary measure is not a universal replacement. Related published work connects dynamical and structural indicators of resilience and examines swarming systems.
Key references
4
- A geometric approach to beta diversity. Ecological Monographs (2025).
- A geometric approach to beta diversity: Reply. Ecological Monographs (2026).
- Merging dynamical and structural indicators to measure resilience in multispecies systems. Journal of Animal Ecology (2021).
- ACCELERATING THE EMERGENCE OF ORDER IN SWARMING SYSTEMS. Advances in Complex Systems (2020).
In the pipeline
2
- Structural sensitivity: a geometric method for community responses to sustained environmental change. Submitted. Luo M et al.
- Quantifying irreversibility of ecological systems. Submitted; public preprint. Li J et al.
Monitoring and global change
What can partial surveys reliably tell us about persistence, and how are communities responding to invasions, land use and fire? We connect limited observations to specific ecological inferences, rather than treating every kind of missing data as the same problem.
Our published monitoring work uses small interaction subnetworks as probabilistic indicators of larger-network persistence. In the Seychelles pollination-network application, we estimated that monitoring five plant species could reduce combined plant-observation and insect-identification costs by 42%. That estimate depends on the study system and monitoring target. Related work studies what incomplete patch surveys can reveal about metapopulation persistence. Imperfect detection, missing species and missing interactions require attention to the assumptions of each method.
Applied studies examine habitat fragmentation and mutualistic assemblages, and land consolidation, rodent communities and disease transmission. Current manuscripts investigate introduced species in seed-dispersal communities, functional-richness measures and the relationship between fuel-treatment hazard reduction and ecological restoration.
Key references
4
- Rapid monitoring of ecological persistence. Proceedings of the National Academy of Sciences (2023).
- Metapopulation persistence can be inferred from incomplete surveys. Proceedings of the Royal Society B: Biological Sciences (2022).
- Anthropogenic land consolidation intensifies zoonotic host diversity loss and disease transmission in human habitats. Nature Ecology & Evolution (2025).
- Larger habitat islands maintain more persistent mutualistic assemblages. Nature Communications (2026).
In the pipeline
3
- Introduced species erode the stability of seed-dispersal communities worldwide. Submitted. Song C et al.
- A mismatch in hazard reduction and ecological restoration in western US fuel treatments. Submitted. Metz T et al.
- Using alpha shapes to measure functional richness: a novel methodology. Submitted. Guoli Z et al.
Ecology meets evolution
How do ecological and evolutionary change shape each other? This newer direction has two connected strands: feedbacks between organisms and communities, and the evolution of trait distributions.
Ecological and evolutionary feedbacks. Published model-testing work examines predator–prey systems with rapid evolution. Submitted manuscripts develop a theory of host–microbiome speciation and investigate predators and antibiotic-resistant microbial strains.
Trait distributions and adaptation. Work in preparation asks how reproductive transmission and selection shape trait distributions, when approximately Gaussian distributions arise, how skewness affects changes in trait variance, and how geometry constrains adaptation in changing environments.
Key references
1
- Rigorous validation of ecological models against empirical time series. Nature Ecology & Evolution (2025).
In the pipeline
6
- A theory of holobiont speciation. Submitted. Yang H et al.
- Microfaunal predators maintain antibiotic resistant strains in microbial communities. Submitted. Hogle SL et al.
- Geometric constraints on adaptation in changing environments. In preparation. Song C
- Skewness shields trait variance from natural selection. In preparation. Song C et al.
- Whence Gaussian: a reproductive renormalization group in quantitative genetics. In preparation. Song C
- Conditional chaos limits for prospective Mendelian transmission under selection. In preparation. Song C