Abstract
Consumptive interactions across trophic levels underpin ecosystem stability. Although theoretical and experimental studies have examined the diversity–stability relationship, empirical evidence linking trophic interactions to ecosystem stability remains limited. Here we evaluate how trophic interactions relate to stability across 430 plant–herbivorous insect networks from temperate and tropical ecosystems. We quantified interaction indicators alongside ecosystem resistance and resilience under drought and wet events. Higher plant diversity was associated with drought resistance but with reduced resilience, and these relationships weakened after accounting for sampling and environmental covariates. In contrast, drought resistance increased with network modularity, whereas resilience decreased with robustness, although these patterns were absent in temperate forests. Diversity and interaction indicators did not predict stability under wet events. Although both directions of trophic interaction–stability relationships were significant, the pathway from trophic interactions to stability (top-down) was stronger than the reverse. Our findings provide large-scale empirical evidence that top-down effects probably dominate the relationship between plant–herbivorous insect interactions and ecosystem stability under climate change.
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Data availability
The raw data associated with this study are available from figshare (https://doi.org/10.6084/m9.figshare.32305317; ref. 95). This study also used previously published data available from the Harmonized World Soil Database (https://gaez.fao.org/pages/hwsd), Global Land Evaporation Amsterdam Model (https://www.gleam.eu) and CHELSA database (https://chelsa-climate.org). Source data are provided with this paper.
Code availability
All R scripts for statistical analyses are available from figshare (https://doi.org/10.6084/m9.figshare.32305317; ref. 95).
References
Forzieri, G., Dakos, V., McDowell, N. G., Ramdane, A. & Cescatti, A. Emerging signals of declining forest resilience under climate change. Nature 608, 534–540 (2022).
Google Scholar
Bennett, A. C. et al. Sensitivity of South American tropical forests to an extreme climate anomaly. Nat. Clim. Change 13, 967–974 (2023).
Google Scholar
Smith, M. D. The ecological role of climate extremes: current understanding and future prospects. J. Ecol. 99, 651–655 (2011).
Google Scholar
De Boeck, H. J. et al. Patterns and drivers of biodiversity–stability relationships under climate extremes. J. Ecol. 106, 890–902 (2018).
Google Scholar
Pimm, S. L. The complexity and stability of ecosystems. Nature 307, 321–326 (1984).
Google Scholar
Tilman, D., Reich, P. B. & Knops, J. M. H. Biodiversity and ecosystem stability in a decade-long grassland experiment. Nature 441, 629–632 (2006).
Google Scholar
Arnoldi, J. F., Loreau, M. & Haegeman, B. The inherent multidimensionality of temporal variability: how common and rare species shape stability patterns. Ecol. Lett. 22, 1557–1567 (2019).
Google Scholar
Chen, Q. et al. Multidimensional responses of grassland stability to eutrophication. Nat. Commun. 14, 6375 (2023).
Google Scholar
Donohue, I. et al. On the dimensionality of ecological stability. Ecol. Lett. 16, 421–429 (2013).
Google Scholar
Pennekamp, F. et al. Biodiversity increases and decreases ecosystem stability. Nature 563, 109–112 (2018).
Google Scholar
Kéfi, S. et al. Advancing our understanding of ecological stability. Ecol. Lett. 22, 1349–1356 (2019).
Google Scholar
Oliver, T. H. et al. Biodiversity and resilience of ecosystem functions. Trends Ecol. Evol. 30, 673–684 (2015).
Google Scholar
Mahecha, M. D. et al. Biodiversity loss and climate extremes—study the feedbacks. Nature 612, 30–32 (2022).
Google Scholar
Hector, A. et al. General stabilizing effects of plant diversity on grassland productivity through population asynchrony and overyielding. Ecology 91, 2213–2220 (2010).
Google Scholar
Schnabel, F. et al. Species richness stabilizes productivity via asynchrony and drought-tolerance diversity in a large-scale tree biodiversity experiment. Sci. Adv. 7, eabk1643 (2021).
Google Scholar
Lhomme, J. P. & Winkel, T. Diversity–stability relationships in community ecology: re-examination of the portfolio effect. Theor. Popul. Biol. 62, 271–279 (2002).
Google Scholar
Yachi, S. & Loreau, M. Biodiversity and ecosystem productivity in a fluctuating environment: the insurance hypothesis. Proc. Natl Acad. Sci. USA 96, 1463–1468 (1999).
Google Scholar
Loreau, M. et al. Biodiversity as insurance: from concept to measurement and application. Biol. Rev. 96, 2333–2354 (2021).
Google Scholar
Gross, K. et al. Species richness and the temporal stability of biomass production: a new analysis of recent biodiversity experiments. Am. Nat. 183, 1–12 (2014).
Google Scholar
Haddad, N. M., Crutsinger, G. M., Gross, K., Haarstad, J. & Tilman, D. Plant diversity and the stability of foodwebs. Ecol. Lett. 14, 42–46 (2011).
Google Scholar
Wang, M. Q. et al. Asynchrony and functional diversity couple herbivore community dynamics to host plant diversity. Nat. Commun. 17, 762 (2026).
Google Scholar
Liang, M., Cappelli, S. L., Borer, E. T., Tilman, D. & Seabloom, E. W. Consumers modulate effects of plant diversity on community stability. Ecol. Lett. 28, e70103 (2025).
Google Scholar
Eschenbrenner, J. & Thébault, É Diversity, food web structure and the temporal stability of total plant and animal biomasses. Oikos 2023, e08769 (2023).
Google Scholar
Thébault, E. & Loreau, M. Trophic interactions and the relationship between species diversity and ecosystem stability. Am. Nat. 166, E95–E114 (2005).
Google Scholar
Zhao, Q. et al. Horizontal and vertical diversity jointly shape food web stability against small and large perturbations. Ecol. Lett. 22, 1152–1162 (2019).
Google Scholar
Halpern, B. S., Borer, E. T., Seabloom, E. W. & Shurin, J. B. Predator effects on herbivore and plant stability. Ecol. Lett. 8, 189–194 (2005).
Google Scholar
O’Gorman, E. J. & Emmerson, M. C. Perturbations to trophic interactions and the stability of complex food webs. Proc. Natl Acad. Sci. USA 106, 13393–13398 (2009).
Google Scholar
Gottschall, F. et al. Spatiotemporal dynamics of abiotic and biotic properties explain biodiversity–ecosystem-functioning relationships. Ecol. Monogr. 92, e01490 (2022).
Google Scholar
Thébault, E. & Fontaine, C. Stability of ecological communities and the architecture of mutualistic and trophic networks. Science 329, 853–856 (2010).
Google Scholar
Robinson, M. L. & Strauss, S. Y. Generalists are more specialized in low-resource habitats, increasing stability of ecological network structure. Proc. Natl Acad. Sci. USA 117, 2043–2048 (2020).
Google Scholar
Worm, B. & Duffy, J. E. Biodiversity, productivity and stability in real food webs. Trends Ecol. Evol. 18, 628–632 (2003).
Google Scholar
Li, Y. et al. Multitrophic arthropod diversity mediates tree diversity effects on primary productivity. Nat. Ecol. Evol. 7, 832–840 (2023).
Google Scholar
Popkin, G. Forest fight. Science 374, 1184–1189 (2021).
Google Scholar
Delmas, E. et al. Analysing ecological networks of species interactions. Biol. Rev. 94, 16–36 (2019).
Google Scholar
Newman, M. E. J. Modularity and community structure in networks. Proc. Natl Acad. Sci. USA 103, 8577–8582 (2006).
Google Scholar
Bascompte, J., Jordano, P., Melián, C. J. & Olesen, J. M. The nested assembly of plant–animal mutualistic networks. Proc. Natl Acad. Sci. USA 100, 9383–9387 (2003).
Google Scholar
Martinez, N. D. Constant connectance in community food webs. Am. Nat. 139, 1208–1218 (1992).
Google Scholar
Dunne, J. A., Williams, R. J. & Martinez, N. D. Network structure and biodiversity loss in food webs: robustness increases with connectance. Ecol. Lett. 5, 558–567 (2002).
Google Scholar
May, R. M. Will a large complex system be stable? Nature 238, 413–414 (1972).
Google Scholar
Kohli, M., Borer, E. T., Kinkel, L. & Seabloom, E. W. Stability of grassland production is robust to changes in the consumer food web. Ecol. Lett. 22, 707–716 (2019).
Google Scholar
Scherber, C. et al. Bottom-up effects of plant diversity on multitrophic interactions in a biodiversity experiment. Nature 468, 553–556 (2010).
Google Scholar
Lu, X. et al. Bottom-up effects of plant quantity and quality on arthropod diversity across multiple trophic levels in a semi-arid grassland. J. Ecol. 110, 2717–2730 (2022).
Google Scholar
Song, C., Rohr, R. P. & Saavedra, S. Why are some plant–pollinator networks more nested than others? J. Anim. Ecol. 86, 1417–1424 (2017).
Google Scholar
Isbell, F. et al. Biodiversity increases the resistance of ecosystem productivity to climate extremes. Nature 526, 574–577 (2015).
Google Scholar
Luo, Y. et al. Multitrophic diversity and biotic associations influence subalpine forest ecosystem multifunctionality. Ecology 103, e3745 (2022).
Google Scholar
Bai, Y. & Tang, Z. Enhanced effects of species richness on resistance and resilience of global tree growth to prolonged drought. Proc. Natl Acad. Sci. USA 121, e2410467121 (2024).
Google Scholar
Hafner, B. D. et al. Hydraulic redistribution under moderate drought among English oak, European beech and Norway spruce determined by deuterium isotope labeling in a split-root experiment. Tree Physiol. 37, 950–960 (2017).
Google Scholar
Jactel, H. & Brockerhoff, E. G. Tree diversity reduces herbivory by forest insects. Ecol. Lett. 10, 835–848 (2007).
Google Scholar
Forrester, D. I. The spatial and temporal dynamics of species interactions in mixed-species forests: from pattern to process. Forest Ecol. Manag. 312, 282–292 (2014).
Google Scholar
Haberstroh, S. & Werner, C. The role of species interactions for forest resilience to drought. Plant Biol. J. 24, 1098–1107 (2022).
Google Scholar
Plaut, J. A. et al. Reduced transpiration response to precipitation pulses precedes mortality in a piñon–juniper woodland subject to prolonged drought. New Phytol. 200, 375–387 (2013).
Google Scholar
Barbeta, A., Ogaya, R. & Peñuelas, J. Dampening effects of long-term experimental drought on growth and mortality rates of a Holm oak forest. Glob. Change Biol. 19, 3133–3144 (2013).
Google Scholar
Hillebrand, H. et al. Decomposing multiple dimensions of stability in global change experiments. Ecol. Lett. 21, 21–30 (2018).
Google Scholar
Liu, Y., Hu, J. & Gore, J. Ecosystem stability relies on diversity difference between trophic levels. Proc. Natl Acad. Sci. USA 121, e2416740121 (2024).
Google Scholar
Blüthgen, N. & Staab, M. A critical evaluation of network approaches for studying species interactions. Annu. Rev. Ecol. Evol. Syst. 55, e1904 (2024).
Google Scholar
Wang, M. Multidimensionality of tree communities structure host–parasitoid networks and their phylogenetic composition. Elife 13, RP100202 (2025).
Google Scholar
López-Carretero, A., Díaz-Castelazo, C., Boege, K. & Rico-Gray, V. Temporal variation in structural properties of tropical plant–herbivore networks: the role of climatic factors. Acta Oecol. 92, 59–66 (2018).
Google Scholar
Santos-Neto, P. E., Arnan, X., Ribeiro-Neto, J. D., Wirth, R. & Leal, I. R. Aridity, but not disturbance, reduces the specialization and modularity of plant–insect herbivore interaction networks in Caatinga dry forest. J. Insect Conserv. 26, 175–189 (2022).
Google Scholar
Weissflog, A., Markesteijn, L., Lewis, O. T., Comita, L. S. & Engelbrecht, B. M. J. Contrasting patterns of insect herbivory and predation pressure across a tropical rainfall gradient. Biotropica 50, 302–311 (2018).
Google Scholar
Coley, P. D., Barone, J. A. & Barone, C. Herbivory and plant defenses in tropical forests. Annu. Rev. Ecol. Syst. 27, 305–340 (1996).
Google Scholar
Givnish, T. J. in Physiological Ecology of Plants of the Wet Tropics 12 (eds Medina, E. et al.) 51–84 (Springer, 1984).
Azevedo-Schmidt, L., Swain, A., Shoemaker, L. G. & Currano, E. D. Landscape-level variability and insect herbivore outbreak captured within modern forests provides a framework for interpreting the fossil record. Sci. Rep. 13, 9701 (2023).
Google Scholar
Gross, T., Rudolf, L., Levin, S. A. & Dieckmann, U. Generalized models reveal stabilizing factors in food webs. Science 325, 747–750 (2009).
Google Scholar
Nie, S. et al. Will a large complex system be productive? Ecol. Lett. 26, 1325–1335 (2023).
Google Scholar
Keyes, A. A., Barner, A. K. & Dee, L. E. Synthesising the relationships between food web structure and robustness. Ecol. Lett. 27, e14533 (2024).
Google Scholar
Blüthgen, N., Menzel, F. & Blüthgen, N. Measuring specialization in species interaction networks. BMC Ecol. 6, 9 (2006).
Google Scholar
Allesina, S. & Tang, S. Stability criteria for complex ecosystems. Nature 483, 205–208 (2012).
Google Scholar
Feng, J. et al. Mediating role of food web structure in linking diversity to multidimensional stability: evidence from global marine ecosystems. Sci. Adv. 41, eadv3841 (2025).
Google Scholar
Phillips, R. D., Peakall, R., van der Niet, T. & Johnson, S. D. Niche perspectives on plant–pollinator interactions. Trends Plant Sci. 25, 779–793 (2020).
Google Scholar
Krasnov, B. R. et al. Phylogenetic signal in module composition and species connectivity in compartmentalized host–parasite networks. Am. Nat. 179, 501–511 (2012).
Google Scholar
Mello, M. A. R. et al. Insights into the assembly rules of a continent-wide multilayer network. Nat. Ecol. Evol. 3, 1525–1532 (2019).
Google Scholar
Vidal, M. C. & Murphy, S. M. Bottom-up vs. top-down effects on terrestrial insect herbivores: a meta-analysis. Ecol. Lett. 21, 138–150 (2018).
Google Scholar
Smilanich, A. M., Fincher, R. M. & Dyer, L. A. Does plant apparency matter? Thirty years of data provide limited support but reveal clear patterns of the effects of plant chemistry on herbivores. New Phytol. 210, 1044–1057 (2016).
Google Scholar
Arif, S. & MacNeil, M. A. Predictive models aren’t for causal inference. Ecol. Lett. 25, 1741–1745 (2022).
Google Scholar
Craven, D. et al. A cross-scale assessment of productivity–diversity relationships. Glob. Ecol. Biogeogr. 29, 1940–1955 (2020).
Google Scholar
O’Dea, R. E. et al. Preferred reporting items for systematic reviews and meta-analyses in ecology and evolutionary biology: a PRISMA extension. Biol. Rev. 96, 1695–1722 (2021).
Google Scholar
Miralles, D. G. et al. GLEAM4: global land evaporation and soil moisture dataset at 0.1° resolution from 1980 to near present. Sci. Data 12, 416 (2025).
Google Scholar
Camps-Valls, G. et al. A unified vegetation index for quantifying the terrestrial biosphere. Sci. Adv. 7, eabc7447 (2021).
Google Scholar
Zeng, Y. et al. Optical vegetation indices for monitoring terrestrial ecosystems globally. Nat. Rev. Earth Environ. 3, 477–493 (2022).
Google Scholar
Zhang, Y., Keenan, T. F. & Zhou, S. Exacerbated drought impacts on global ecosystems due to structural overshoot. Nat. Ecol. Evol. 5, 1490–1498 (2021).
Google Scholar
Yao, Y., Liu, Y., Zhou, S., Song, J. & Fu, B. Soil moisture determines the recovery time of ecosystems from drought. Glob. Change Biol. 29, 3562–3574 (2023).
Google Scholar
Vicente-Serrano, S. M. et al. Response of vegetation to drought time-scales across global land biomes. Proc. Natl Acad. Sci. USA 110, 52–57 (2013).
Google Scholar
Van Passel, J. et al. Climatic legacy effects on the drought response of the Amazon rainforest. Glob. Change Biol. 28, 5808–5819 (2022).
Google Scholar
Yang, H. et al. The detection and attribution of extreme reductions in vegetation growth across the global land surface. Glob. Change Biol. 29, 2351–2362 (2023).
Google Scholar
Bai, Y., Chen, J., Zhang, Y. & Tang, Z. Response modes of global vegetation to extreme drought. Glob. Change Biol. 31, e70488 (2025).
Google Scholar
Ingrisch, J. & Bahn, M. Towards a comparable quantification of resilience. Trends Ecol. Evol. 33, 251–259 (2018).
Google Scholar
Yao, Y. et al. Declining tradeoff between resistance and resilience of ecosystems to drought. Earths Future 12, e2024EF004665 (2024).
Google Scholar
Arrhenius, O. Species and area. J. Ecol. 9, 95–99 (1921).
Google Scholar
McCain, C. M. Area and mammalian elevational diversity. Ecology 88, 76–86 (2007).
Google Scholar
Bates, D., Mächler, M., Bolker, B. & Walker, S. Fitting linear mixed-effects models using lme4. J. Stat. Softw. 67, 1–48 (2015).
Google Scholar
Le Bagousse-Pinguet, Y. et al. Testing the environmental filtering concept in global drylands. J. Ecol. 105, 1058–1069 (2017).
Google Scholar
Westreich, D. & Greenland, S. The table 2 fallacy: presenting and interpreting confounder and modifier coefficients. Am. J. Epidemiol. 177, 292–298 (2013).
Google Scholar
Lai, J., Zou, Y., Zhang, S., Zhang, X. & Mao, L. Glmm.Hp: an R package for computing individual effect of predictors in generalized linear mixed models. J. Plant Ecol. 15, 1302–1307 (2022).
Google Scholar
Lefcheck, J. S. piecewiseSEM: Piecewise structural equation modelling in R for ecology, evolution, and systematics. Methods Ecol. Evol. 7, 573–579 (2016).
Google Scholar
Wang, Y., Qin, Q., Yu, Z. & Liu, J. Dataset & R codes-Top-down effects dominate the relationship between plant–herbivorous insect interactions and ecosystem stability. figshare https://doi.org/10.6084/m9.figshare.32305317 (2026).
Peres-Neto, P. R. et al. Variation partitioning of species data matrices: estimation and comparison of fractions. Ecology 87, 2614–2625 (2006).
Google Scholar
Funding
This work was supported by grants from the National Natural Science Foundation of China (32501439), Hebei Provincial Natural Science Foundation (C2024205002) and Science Foundation of Hebei Normal University (L2024B18).
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Y.W., Q.Q. and J.L. conceived of the study idea. Y.W., Q.Q., Z.W. and Y.Z. collected the data. Y.W., Q.Q. and J.F. performed the analyses. The manuscript was drafted by Y.W. and Q.Q. with contributions from J.L., Z.Y. and C.W. All authors contributed critically to the manuscript and gave final approval for publication.
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Extended data
Extended Data Fig. 1 Relationships between plant diversity, trophic interaction indicators and wet resistance or resilience across ecosystem types.
Circles and error bars represent average parameter estimates (standardized coefficients) and 95% confidence intervals based on linear mixed-effects models: “lmer(stability ~ (1|study ID) + sampling effects + trophic interactions or plant diversity)”. P values represent the probability of type I error based on analysis of variance (two-sided F-test). Values in bold indicate significant (P < 0.05) or marginally significant (0.05 ≤ P < 0.1) effects. The complete results are presented in Supplementary Tables 3 and 6. The numbers in parentheses show sample sizes.
Extended Data Fig. 2 Relative importance of plant diversity and trophic interactions on wet resistance and resilience.
a–f, Relative importance of plant diversity and trophic interactions on wet resistance (a–c) or resilience (d–f) across ecosystem types (temperate forest (a and d), tropical forest (b and e) and savanna (c and f)). The bar plots indicate percentages of explained variance (relative importance) for plant diversity, trophic interactions, covariates and random effects. The marginal R2 (R2m) and conditional R2 (R2c) values represent the proportion of variance explained by all factors without accounting for random effects and accounting for random effects, respectively. It is important to note that variable importance based on hierarchical partitioning can be negative due to multicollinearity96. The numbers in parentheses show sample sizes.
Extended Data Fig. 3 Relationships of plant diversity and trophic interaction indicators across ecosystem types.
a–c, Results from piecewise SEM on the relationships of plant diversity and trophic interaction indicators in temperate forests (a), tropical forests (b) and savannas (c). Alternative SEMs were tested to elucidate the relationships between plant diversity and interaction indicators (Supplementary Fig. 3). The values represent standardized path coefficients. Solid arrows denote significant positive (blue) or negative (red) relationships (P < 0.05; two-sided test). Dashed arrows (grey) denote non-significant paths (P > 0.05; two-sided test). *P < 0.05; **P < 0.01; ***P < 0.001. The numbers in parentheses show sample sizes.
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Wang, Y., Qin, Q., Feng, J. et al. Top-down effects dominate the relationship between plant–herbivorous insect interactions and ecosystem stability.
Nat Ecol Evol (2026). https://doi.org/10.1038/s41559-026-03114-1
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DOI: https://doi.org/10.1038/s41559-026-03114-1
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