Abstract
Freshwater and terrestrial ecosystems are linked by reciprocal flows of energy and nutrients, including the emergence of aquatic insects that provide abundant, high-quality prey for riparian birds. Local studies show that emergent aquatic insects can enhance bird reproduction and survival, yet it remains unclear whether such associations scale up across landscapes and persist under human disturbance. Here we show that the richness of mayflies, stoneflies and caddisflies (Ephemeroptera, Plecoptera and Trichoptera; EPT), an indicator of freshwater integrity and insect emergence, predicts the prevalence of 288 bird species across 14,177 rivers in the contiguous USA, even after accounting for human land use. Aerial insectivorous birds exhibit more than threefold higher prevalence at sites with high compared with low EPT richness. This linkage is consistent across river types describing hydrology (including intermittent systems), temperature and geomorphology, with the strongest gains occurring in rivers with low baseline bird prevalence. The associations are weaker in urbanized sites but persist in highly agricultural landscapes provided that EPT richness remains relatively high. Our findings provide continental-scale evidence that emergent aquatic insects sustain riparian birds, underscoring that freshwater conservation offers considerable co-benefits to terrestrial ecosystems.
This is a preview of subscription content, access via your institution
Access options
Access through your institution
Access Nature and 54 other Nature Portfolio journals
Get Nature+, our best-value online-access subscription
$32.99 / 30 days
cancel any time
Subscribe to this journal
Receive 12 digital issues and online access to articles
$119.00 per year
only $9.92 per issue
Buy this article
- Purchase on SpringerLink
- Instant access to the full article PDF.
USD 39.95
Prices may be subject to local taxes which are calculated during checkout
Similar content being viewed by others
Fluctuation in the diversity of mayflies (Insecta, Ephemerida) as documented in the fossil record
The ecological benefits of more room for rivers
Standardized diversity estimation uncovers global distribution patterns and drivers of stream insects
Data availability
Bird survey data were obtained from the eBird Basic Dataset40 (via Cornell Lab of Ornithology at https://ebird.org/science/download-ebird-data-products) and are available under a data-use agreement from the Cornell Lab of Ornithology. Aquatic invertebrate data were compiled from US federal and state biomonitoring programmes and are subject to the data-sharing policies of the respective agencies. As these datasets are owned by the contributing agencies, the original raw data cannot be redistributed by the authors. Information on data sources, spatial coverage and contact details for access requests is provided in Supplementary Table 6. The minimum dataset with processed data required to reproduce the analyses and figures presented in this study, together with the analysis code, are publicly available via Zenodo at https://doi.org/10.5281/zenodo.17643668 (ref. 81).
Code availability
All code used for data processing, statistical analyses and figure generation is available via Zenodo at https://doi.org/10.5281/zenodo.17643668 (ref. 81).
References
Reid, A. J. et al. Emerging threats and persistent conservation challenges for freshwater biodiversity. Biol. Rev. 94, 849–873 (2019).
Google Scholar
Tickner, D. et al. Bending the curve of global freshwater biodiversity loss: an emergency recovery plan. BioScience 70, 330–342 (2020).
Google Scholar
Bowler, D. E., Heldbjerg, H., Fox, A. D., De Jong, M. & Böhning-Gaese, K. Long-term declines of European insectivorous bird populations and potential causes. Conserv. Biol. 33, 1120–1130 (2019).
Google Scholar
Johnston, A. et al. North American bird declines are greatest where species are most abundant. Science 388, 532–537 (2025).
Google Scholar
Both, C., Bouwhuis, S., Lessells, C. M. & Visser, M. E. Climate change and population declines in a long-distance migratory bird. Nature 441, 81–83 (2006).
Google Scholar
Møller, A. P. Parallel declines in abundance of insects and insectivorous birds in Denmark over 22 years. Ecol. Evol. 9, 6581–6587 (2019).
Google Scholar
Tallamy, D. W. & Shriver, W. G. Are declines in insects and insectivorous birds related?. Ornithol. Appl. 123, duaa059 (2021).
Adams, V. M. et al. Planning across freshwater and terrestrial realms: cobenefits and tradeoffs between conservation actions. Conserv. Lett. 7, 425–440 (2014).
Google Scholar
Leal, C. G. et al. Integrated terrestrial–freshwater planning doubles conservation of tropical aquatic species. Science 370, 117–121 (2020).
Google Scholar
Wegscheider, B. et al. Modeling nature-based restoration potential across aquatic–terrestrial boundaries. Conserv. Biol. 39, e70046 (2025).
Google Scholar
Kowarik, C., Martin-Creuzburg, D., Mathers, K. L., Weber, C. & Robinson, C. T. Stream degradation affects aquatic resource subsidies to riparian ground-dwelling spiders. Sci. Total Environ. 855, 158658 (2023).
Google Scholar
Nakano, S. & Murakami, M. Reciprocal subsidies: dynamic interdependence between terrestrial and aquatic food webs. Proc. Natl Acad. Sci. USA 98, 166–170 (2001).
Google Scholar
Twining, C. W., Shipley, J. R. & Winkler, D. W. Aquatic insects rich in omega-3 fatty acids drive breeding success in a widespread bird. Ecol. Lett. 21, 1812–1820 (2018).
Google Scholar
Baxter, C. V., Fausch, K. D. & Carl Saunders, W. Tangled webs: reciprocal flows of invertebrate prey link streams and riparian zones. Freshw. Biol. 50, 201–220 (2005).
Google Scholar
Jackson, J. K. & Fisher, S. G. Secondary production, emergence, and export of aquatic insects of a Sonoran Desert stream. Ecology 67, 629–638 (1986).
Google Scholar
Twining, C. W. et al. Aquatic and terrestrial resources are not nutritionally reciprocal for consumers. Funct. Ecol. 33, 2042–2052 (2019).
Google Scholar
Schilke, P. R. et al. Modeling a cross-ecosystem subsidy: forest songbird response to emergent aquatic insects. Landsc. Ecol. 35, 1587–1604 (2020).
Google Scholar
Marczak, L. B., Thompson, R. M. & Richardson, J. S. Meta-analysis: trophic level, habitat, and productivity shape the food web effects of resource subsidies. Ecology 88, 140–148 (2007).
Google Scholar
Lafage, D. et al. Local and landscape drivers of aquatic-to-terrestrial subsidies in riparian ecosystems: a worldwide meta-analysis. Ecosphere 10, e02697 (2019).
Google Scholar
Subalusky, A. L. & Post, D. M. Context dependency of animal resource subsidies. Biol. Rev. 94, 517–538 (2019).
Google Scholar
Muehlbauer, J. D., Collins, S. F., Doyle, M. W. & Tockner, K. How wide is a stream? Spatial extent of the potential “stream signature” in terrestrial food webs using meta-analysis. Ecology 95, 44–55 (2014).
Google Scholar
Nash, L. N. et al. Aquatic–terrestrial linkages drive contrasting biodiversity patterns in tropical and temperate forests. Proc. R. Soc. B 292, 20242423 (2025).
Google Scholar
Brooks, J. M., Baxter, C. V., Warren, D. R. & MacNeill, K. L. Enter the mosaic: aquatic–terrestrial reciprocal fluxes and food webs are dynamically interdependent across space and through time. Freshw. Biol. 70, e70094 (2025).
Google Scholar
Martin-Creuzburg, D., Kowarik, C. & Straile, D. Cross-ecosystem fluxes: export of polyunsaturated fatty acids from aquatic to terrestrial ecosystems via emerging insects. Sci. Total Environ. 577, 174–182 (2017).
Google Scholar
Shipley, J. R. et al. Consumer biodiversity increases organic nutrient availability across aquatic and terrestrial ecosystems. Science 386, 335–340 (2024).
Google Scholar
Stenroth, K., Polvi, L. E., Fältström, E. & Jonsson, M. Land-use effects on terrestrial consumers through changed size structure of aquatic insects. Freshw. Biol. 60, 136–149 (2015).
Google Scholar
Eriksen, T. E. et al. A global perspective on the application of riverine macroinvertebrates as biological indicators in Africa, South-Central America, Mexico and Southern Asia. Ecol. Indic. 126, 107609 (2021).
Google Scholar
Statzner, B. & Resh, V. H. Multiple-site and-year analyses of stream insect emergence: a test of ecological theory. Oecologia 96, 65–79 (1993).
Google Scholar
Larsen, S., Muehlbauer, J. D. & Marti, E. Resource subsidies between stream and terrestrial ecosystems under global change. Glob. Change Biol. 22, 2489–2504 (2016).
Google Scholar
Schulz, R. et al. A synthesis of anthropogenic stress effects on emergence-mediated aquatic–terrestrial linkages and riparian food webs. Sci. Total Environ. 908, 168186 (2024).
Google Scholar
Schürings, C., Feld, C. K., Kail, J. & Hering, D. Effects of agricultural land use on river biota: a meta-analysis. Environ. Sci. Eur. 34, 124 (2022).
Google Scholar
Peredo Arce, A., Palt, M., Schletterer, M. & Kail, J. Has riparian woody vegetation a positive effect on dispersal and distribution of mayfly, stonefly and caddisfly species?. Sci. Total Environ. 879, 163137 (2023).
Google Scholar
Perkin, E. K. et al. The influence of artificial light on stream and riparian ecosystems: questions, challenges, and perspectives. Ecosphere 2, art122 (2011).
Google Scholar
Walsh, C. J. et al. The urban stream syndrome: current knowledge and the search for a cure. J. N. Am. Benthol. Soc. 24, 706–723 (2005).
Google Scholar
Ohler, K., Schreiner, V. C., Link, M., Liess, M. & Schäfer, R. B. Land use changes biomass and temporal patterns of insect cross-ecosystem flows. Glob. Change Biol. 29, 81–96 (2023).
Google Scholar
Kraus, J. M., Walters, D. M. & Mills, M. A. (eds) Contaminants and Ecological Subsidies: The Land–Water Interface. (Springer, 2020); https://doi.org/10.1007/978-3-030-49480-3
Anderson, H. E., Albertson, L. K. & Walters, D. M. Thermal variability drives synchronicity of an aquatic insect resource pulse. Ecosphere 10, e02852 (2019).
Google Scholar
Goetz, S., Steinberg, D., Dubayah, R. & Blair, B. Laser remote sensing of canopy habitat heterogeneity as a predictor of bird species richness in an eastern temperate forest, USA. Remote Sens. Environ. 108, 254–263 (2007).
Google Scholar
Progar, R. & Moldenke, A. R. Aquatic insect emergence from headwater streams flowing through regeneration and mature forests in Western Oregon. J. Freshw. Ecol. 24, 53–66 (2009).
Google Scholar
Sullivan, B. L. et al. The eBird enterprise: an integrated approach to development and application of citizen science. Biol. Conserv. 169, 31–40 (2014).
Google Scholar
Birds of the world. Cornell Lab of Ornithology https://doi.org/10.2173/bow (2020).
Wilman, H. et al. EltonTraits 1.0: species-level foraging attributes of the world’s birds and mammals: ecological archives E095-178. Ecology 95, 2027 (2014).
Google Scholar
Aronson, M. F. J. et al. A global analysis of the impacts of urbanization on bird and plant diversity reveals key anthropogenic drivers. Proc. R. Soc. B 281, 20133330 (2014).
Google Scholar
Chace, J. F. & Walsh, J. J. Urban effects on native avifauna: a review. Landsc. Urban Plann. 74, 46–69 (2006).
Google Scholar
Benton, T. G., Vickery, J. A. & Wilson, J. D. Farmland biodiversity: is habitat heterogeneity the key?. Trends Ecol. Evol. 18, 182–188 (2003).
Google Scholar
Johnston, A., Matechou, E. & Dennis, E. B. Outstanding challenges and future directions for biodiversity monitoring using citizen science data. Methods Ecol. Evol. 14, 103–116 (2023).
Google Scholar
Isaac, N. J. B., Van Strien, A. J., August, T. A., De Zeeuw, M. P. & Roy, D. B. Statistics for citizen science: extracting signals of change from noisy ecological data. Methods Ecol. Evol. 5, 1052–1060 (2014).
Google Scholar
Messager, M. L. et al. Global prevalence of non-perennial rivers and streams. Nature 594, 391–397 (2021).
Google Scholar
Datry, T. et al. Causes, responses, and implications of anthropogenic versus natural flow intermittence in river networks. BioScience 73, 9–22 (2023).
Google Scholar
Manning, D. W. P. & Sullivan, S. M. P. Conservation across aquatic–terrestrial boundaries: linking continental-scale water quality to emergent aquatic insects and declining aerial insectivorous birds. Front. Ecol. Evol. 9, 633160 (2021).
Google Scholar
Leathers, K., Herbst, D., De Mendoza, G., Doerschlag, G. & Ruhi, A. Climate change is poised to alter mountain stream ecosystem processes via organismal phenological shifts. Proc. Natl Acad. Sci. USA 121, e2310513121 (2024).
Google Scholar
Lovász, L., Korner-Nievergelt, F. & Amrhein, V. Natural grazing by horses and cattle promotes bird diversity in a restored European alluvial grassland. PeerJ 12, e17777 (2024).
Google Scholar
Herzon, I. & Helenius, J. Agricultural drainage ditches, their biological importance and functioning. Biol. Conserv. 141, 1171–1183 (2008).
Google Scholar
Basile, M., Augustinus, B. A. & Brockerhoff, E. G. Urban tree canopy cover over 30% and native trees enhance bird insectivory and tree biosecurity. Biol. Conserv. 310, 111387 (2025).
Google Scholar
MacDade, L. S., Rodewald, P. G. & Hatch, K. A. Contribution of emergent aquatic insects to refueling in spring migrant songbirds. Auk 128, 127–137 (2011).
Google Scholar
Imlay, T. L., Mann, H. A. R. & Leonard, M. L. No effect of insect abundance on nestling survival or mass for three aerial insectivores. Avian Conserv. Ecol. 12, 19 (2017).
Google Scholar
Gratton, C. & Zanden, M. J. V. Flux of aquatic insect productivity to land: comparison of lentic and lotic ecosystems. Ecology 90, 2689–2699 (2009).
Google Scholar
Walter, J. A., Fleck, R., Kastens, J. H., Pace, M. L. & Wilkinson, G. M. Temporal coherence between lake and landscape primary productivity. Ecosystems 24, 502–515 (2021).
Google Scholar
Acreman, M., Hughes, K. A., Arthington, A. H., Tickner, D. & Dueñas, M. Protected areas and freshwater biodiversity: a novel systematic review distils eight lessons for effective conservation. Conserv. Lett. 13, e12684 (2020).
Google Scholar
Skagen, S. K. et al. Geography of spring landbird migration through riparian habitats in southwestern North America. Condor 107, 212–227 (2005).
Google Scholar
Peller, T. & Altermatt, F. Invasive species drive cross-ecosystem effects worldwide. Nat. Ecol. Evol. 8, 1087–1097 (2024).
Google Scholar
Mason, R. J. et al. Rebalancing river lateral connectivity: an interdisciplinary focus for research and management. WIREs Water 12, e1766 (2025).
Google Scholar
McCabe, C. L., Matthaei, C. D. & Tonkin, J. D. The ecological benefits of more room for rivers. Nat. Water 3, 260–270 (2025).
Google Scholar
Strimas-Mackey, M. et al. Best practices for using eBird data. Version 2.0. Zenodo https://doi.org/10.5281/zenodo.3620739 (2023).
Fink, D. et al. eBird status and trends, data version 2023. Cornell Lab of Ornithology https://doi.org/10.2173/WZTW8903 (2025).
McKay, L. et al. NHDPlus Version 2: User Guide (Data Model Version 2.1) (US Environmental Protection Agency & US Geological Survey, 2012).
Sundermann, A., Stoll, S. & Haase, P. River restoration success depends on the species pool of the immediate surroundings. Ecol. Appl. 21, 1962–1971 (2011).
Google Scholar
Lorenz, A. W. & Feld, C. K. Upstream river morphology and riparian land use overrule local restoration effects on ecological status assessment. Hydrobiologia 704, 489–501 (2013).
Google Scholar
Abood, S. A., Wieczorek, M. & Spencer, L. US Forest Service national riparian areas base map for the conterminous United States in 2019. US Department of Agriculture https://doi.org/10.2737/RDS-2019-0030 (2022).
Dewitz, J. National Land Cover Database (NLCD) 2019 (US Geological Survey, 2021).
Multi-Resolution Land Characteristics (MRLC) Consortium. NLCD Tree Cover (US Geological Survey, 2021).
McManamay, R. A. & DeRolph, C. R. A stream classification system for the conterminous United States. Sci. Data 6, 190017 (2019).
Google Scholar
Bradt, P., Urban, M., Goodman, N., Bissell, S. & Spiegel, I. Stability and resilience in benthic macroinvertebrate assemblages. Hydrobiologia 403, 123–133 (1999).
Google Scholar
Johnston, A. et al. Analytical guidelines to increase the value of community science data: an example using eBird data to estimate species distributions. Divers. Distrib. 27, 1265–1277 (2021).
Google Scholar
Bonada, N., Rieradevall, M., Prat, N. & Resh, V. H. Benthic macroinvertebrate assemblages and macrohabitat connectivity in Mediterranean-climate streams of northern California. J. N. Am. Benthol. Soc. 25, 32–43 (2006).
Google Scholar
Martins, R. T. et al. Congruence and responsiveness in the taxonomic compositions of Amazonian aquatic macroinvertebrate and fish assemblages. Hydrobiologia 849, 2281–2298 (2022).
Google Scholar
Brooks, M. E. et al. glmmTMB balances speed and flexibility among packages for zero-inflated generalized linear mixed modeling. R J. 9, 378 (2017).
Google Scholar
Smithson, M. & Verkuilen, J. A better lemon squeezer? Maximum-likelihood regression with beta-distributed dependent variables. Psychol. Methods 11, 54–71 (2006).
Google Scholar
Hartig, F. DHARMa: residual diagnostics for hierarchical (multi-level/mixed) regression models. R Package Version 0.4.7 https://doi.org/10.32614/CRAN.package.DHARMa (2022).
Nagelkerke, N. J. D. A note on a general definition of the coefficient of determination. Biometrika 78, 691–692 (1991).
Google Scholar
Schürings et al. Freshwater insect richness predicts riparian bird prevalence across the conterminous United States (version 1). Zenodo https://doi.org/10.5281/zenodo.17643668 (2026).
Google Scholar
Acknowledgements
This work was supported by the Deutsche Forschungsgemeinschaft (DFG, German Research Foundation) through a Walter Benjamin Fellowship (project number SCHU 3827/1-1 to C.S.) and by the Richard C. and Lois M. Worthington Endowed Professor in Fisheries Management (University of Washington) to J.D.O. We thank the US Environmental Protection Agency, US Geological Survey, Bureau of Land Management, US Forest Service, US Fish and Wildlife Service, National Park Service and numerous state agencies for conducting and providing macroinvertebrate monitoring data, and the Cornell Lab of Ornithology for access to eBird data. We thank D. Fink for discussions on the use of eBird data and acknowledge the many citizen scientists whose observations made this study possible. We thank G. Jacuzzi for constructive feedback.
Author information
Authors and Affiliations
Contributions
C.S. and J.D.O. conceptualized the study. C.S. carried out the analyses, and C.S. and J.D.O. wrote the paper.
Corresponding author
Ethics declarations
Competing interests
The authors declare no competing interests.
Peer review
Peer review information
Nature Ecology & Evolution thanks Federico Morelli and the other, anonymous, reviewer(s) for their contribution to the peer review of this work. Peer reviewer reports are available.
Additional information
Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Extended data
Extended Data Fig. 1 Spatial distribution of all aquatic insect and bird survey locations prior to filtering.
(A) Red points show all aquatic insect survey sites that fall within the map extent (total dataset: 141,150 sites). (B) Blue points show all eBird survey locations within the displayed area (total dataset: 11,962,137 surveys).
Extended Data Fig. 2 Overview of data integration and analytical workflow.
Schematic representation of all processing steps linking aquatic insect surveys with riparian bird communities across the conterminous United States. Federal and state biomonitoring data were filtered for recent (≤ 25 years) spring–summer samples, and eBird surveys were standardized for effort and trait-annotated. Riparian interaction zones were delineated around rivers (50–550 m buffers) and intersected with bird surveys within 5 km of insect sites. Proximity weighting accounted for spatial and temporal correspondence, followed by 5-km rarefaction to ensure independence. Insect and bird metrics (EPT richness and weighted bird prevalence) were related to riparian land-cover variables using hierarchical hurdle models, with subsequent validation, diagnostics, and marginal-effects analyses.
Supplementary information
Supplementary Information (download PDF )
Supplementary Tables 1–9.
Reporting Summary (download PDF )
Peer Review File (download PDF )
Rights and permissions
Springer Nature or its licensor (e.g. a society or other partner) holds exclusive rights to this article under a publishing agreement with the author(s) or other rightsholder(s); author self-archiving of the accepted manuscript version of this article is solely governed by the terms of such publishing agreement and applicable law.
Reprints and permissions
About this article
Cite this article
Schürings, C., Olden, J.D. Cross-ecosystem linkages between freshwater insects and riparian birds across the USA.
Nat Ecol Evol (2026). https://doi.org/10.1038/s41559-026-03041-1
Received:
Accepted:
Published:
Version of record:
DOI: https://doi.org/10.1038/s41559-026-03041-1
Source: Ecology - nature.com

