{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T08:51:12Z","timestamp":1791276672983,"version":"4.1.0"},"reference-count":101,"publisher":"Wiley","issue":"7","license":[{"start":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T00:00:00Z","timestamp":1791244800000},"content-version":"vor","delay-in-days":5,"URL":"http:\/\/onlinelibrary.wiley.com\/termsAndConditions#vor"},{"start":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T00:00:00Z","timestamp":1790812800000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/100006103","name":"Southern SARE","doi-asserted-by":"publisher","award":["GS23\u2010279"],"award-info":[{"award-number":["GS23\u2010279"]}],"id":[{"id":"10.13039\/100006103","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["esajournals.onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Ecological Applications"],"published-print":{"date-parts":[[2026,10]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>\n                    Grasslands cover nearly 40% of the terrestrial surface and support essential ecosystem services, yet their biodiversity, productivity, and stability are increasingly threatened by climate extremes, land use change, and disruption of biogeochemical cycles in the Anthropocene. Nitrogen (N) limitation is a primary constraint on grassland functioning, and incorporation of N\u2010fixing legumes can alleviate this limitation. However, system outcomes depend less on legume presence alone than on the balance achieved among grass and legume functional groups. Here, we present a conceptual synthesis integrating ecological theory with empirical evidence to examine how continuous variation in functional\u2010group balance regulates N cycling, primary production, forage nutritive value, livestock performance and ecosystem resilience. Our synthesis advances four main points: (1) Ecosystem responses along species gradients are frequently nonlinear, with intermediate optima emerging from facilitative N niche complementarity interacting with physiological and competitive constraints. (2) These optima vary systematically with climatic context, species physiology and management intensity, with C\n                    <jats:sub>4<\/jats:sub>\n                    grass\u2010based systems in warm climates typically reaching peak performance at lower legume proportions than cooler C\n                    <jats:sub>3<\/jats:sub>\n                    grass\u2010dominated systems. (3) Climate change is likely to decouple vegetation dynamics from functional optima by disproportionately stimulating legume growth and altering water and nutrient efficiencies, increasing the risk of mismatches between botanical composition and ecosystem performance. (4) Integrating resource\u2010ratio and state\u2010and\u2010transition frameworks, we discuss how grazing management, N inputs, and herbivore diet selectivity can shift systems toward target species proportions that enhance production efficiency while strengthening resistance and recovery capacity in mixed grassland. We conclude by outlining research and management agendas emphasizing gradient\u2010based experimental and observational designs to better resolve continuous agroecological response functions, improve cross\u2010system synthesis, and inform context\u2010specific strategies for enhancing productivity, efficiency, and resilience of grasslands under increasing climatic variability.\n                  <\/jats:p>","DOI":"10.1002\/eap.70324","type":"journal-article","created":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T07:57:46Z","timestamp":1791273466000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Facilitative N niche complementarity through grass\u2013legume balance in the Anthropocene: A conceptual synthesis"],"prefix":"10.1002","volume":"36","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-3258-6272","authenticated-orcid":false,"given":"Nicolas","family":"Caram","sequence":"first","affiliation":[{"name":"Agronomy Department University of Florida  Gainesville Florida USA"},{"name":"Departamento de Producci\u00f3n Animal y Pasturas, Facultad de Agronom\u00eda Universidad de la Rep\u00fablica  Paysand\u00fa Uruguay"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lynn E.","family":"Sollenberger","sequence":"additional","affiliation":[{"name":"Agronomy Department University of Florida  Gainesville Florida USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,10,6]]},"reference":[{"key":"e_1_2_12_2_1","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1016\/j.envexpbot.2005.10.008","article-title":"Effect of Elevated Temperature and Water Availability on CO2 Exchange and Nitrogen Fixation of Nodulated Alfalfa Plants","volume":"59","author":"Aranjuelo I.","year":"2007","journal-title":"Environmental and Experimental Botany"},{"key":"e_1_2_12_3_1","doi-asserted-by":"crossref","first-page":"377","DOI":"10.1007\/s10584-019-02464-z","article-title":"Global and Regional Impacts of Climate Change at Different Levels of Global Temperature Increase","volume":"155","author":"Arnell N. W.","year":"2019","journal-title":"Climatic Change"},{"issue":"3","key":"e_1_2_12_4_1","doi-asserted-by":"crossref","first-page":"721","DOI":"10.1002\/eap.1680","article-title":"Elevated CO2 Induces Substantial and Persistent Declines in Forage Quality Irrespective of Warming in Mixed Grass Prairie","volume":"28","author":"Augustine D. J.","year":"2018","journal-title":"Ecological Applications"},{"issue":"10","key":"e_1_2_12_5_1","doi-asserted-by":"crossref","first-page":"720","DOI":"10.1038\/s43017-021-00207-2","article-title":"Combatting Global Grassland Degradation","volume":"2","author":"Bardgett R. D.","year":"2021","journal-title":"Nature Reviews Earth & Environment"},{"issue":"2","key":"e_1_2_12_6_1","doi-asserted-by":"crossref","first-page":"167","DOI":"10.1016\/j.tree.2018.10.013","article-title":"The Future of Complementarity: Disentangling Causes from Consequences","volume":"34","author":"Barry K. E.","year":"2019","journal-title":"Trends in Ecology & Evolution"},{"issue":"3","key":"e_1_2_12_7_1","doi-asserted-by":"crossref","first-page":"311","DOI":"10.2134\/jpa1994.0311","article-title":"Improving Legume Persistence in Forage Crop Systems","volume":"7","author":"Beuselinck P. R.","year":"1994","journal-title":"Journal of Production Agriculture"},{"issue":"5","key":"e_1_2_12_8_1","doi-asserted-by":"crossref","first-page":"748","DOI":"10.1017\/S1751731112000304","article-title":"The Importance of Grasslands for Animal Production and Other Functions: A Review on Management and Methodological Progress in the Tropics","volume":"6","author":"Boval M.","year":"2012","journal-title":"Animal"},{"key":"e_1_2_12_9_1","volume-title":"Model Selection and Multimodel Inference: A Practical Information\u2010Theoretic Approach","author":"Burnham K. P.","year":"2002"},{"key":"e_1_2_12_10_1","doi-asserted-by":"crossref","first-page":"1251","DOI":"10.1093\/jxb\/erp326","article-title":"Water\u2010Tolerant Legume Nodulation","volume":"61","author":"Capoen W.","year":"2010","journal-title":"Journal of Experimental Botany"},{"key":"e_1_2_12_11_1","unstructured":"Caram N.2026.\u201cFacilitative N Niche Complementarity through Grass\u2010Legume Balance in the Anthropocene: A Conceptual Synthesis [Dataset].\u201dZenodo.https:\/\/doi.org\/10.5281\/zenodo.21925852."},{"key":"e_1_2_12_12_1","doi-asserted-by":"crossref","first-page":"152","DOI":"10.1016\/j.rama.2024.06.006","article-title":"Grazing Intensity Controls the Seasonal Utilization of Functionally Diverse Patches by Mediating Herbivore Selectivity","volume":"96","author":"Caram N.","year":"2024","journal-title":"Rangeland Ecology & Management"},{"key":"e_1_2_12_13_1","doi-asserted-by":"crossref","DOI":"10.1016\/j.agsy.2022.103582","article-title":"Studying Beef Production Evolution to Plan for Ecological Intensification of Grazing Ecosystems","volume":"205","author":"Caram N.","year":"2023","journal-title":"Agricultural Systems"},{"key":"e_1_2_12_14_1","doi-asserted-by":"crossref","first-page":"1085","DOI":"10.1007\/s11104-025-07234-8","article-title":"Nitrogen Niche Differentiation and Overyielding Along a Legume Proportion Gradient in a Subtropical Grass\u2013Legume Mixture","volume":"513","author":"Caram N.","year":"2025","journal-title":"Plant and Soil"},{"key":"e_1_2_12_15_1","doi-asserted-by":"crossref","DOI":"10.1002\/csc2.21426","article-title":"Optimum Legume Proportion for Plant and Animal Production in a Subtropical Grassland","volume":"65","author":"Caram N.","year":"2025","journal-title":"Crop Science"},{"issue":"4","key":"e_1_2_12_16_1","doi-asserted-by":"crossref","DOI":"10.1093\/aobpla\/plaf036","article-title":"Nutritional and Physiological Limitations Shape the Radiation\u2010Use Efficiency Response of Grass\u2013Legume Mixtures in Warm Climates","volume":"17","author":"Caram N.","year":"2025","journal-title":"AoB Plants"},{"key":"e_1_2_12_17_1","doi-asserted-by":"crossref","DOI":"10.1016\/j.eja.2026.128154","article-title":"Nitrogen Limitation and Species Balance Trade\u2010Offs in C4 Grass\u2010Legume Mixtures: Evidence from the Nitrogen Nutrition Index","volume":"178","author":"Caram N.","year":"2026","journal-title":"European Journal of Agronomy"},{"issue":"2","key":"e_1_2_12_18_1","doi-asserted-by":"crossref","first-page":"353","DOI":"10.1023\/A:1024847017371","article-title":"Nitrogen Fixation in Perennial Forage Legumes in the Field","volume":"253","author":"Carlsson G.","year":"2003","journal-title":"Plant and Soil"},{"issue":"2","key":"e_1_2_12_19_1","doi-asserted-by":"crossref","first-page":"311","DOI":"10.1080\/01904167.2012.739250","article-title":"Nitrogen Limitation of Rhizoma Peanut Growth","volume":"36","author":"Cathey S. 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Delayed Complementarity, Overyielding, and Improved Residue Composition in Ageing Alfalfa\u2013Fescue Mixtures","volume":"495","author":"Louarn G.","year":"2024","journal-title":"Plant and Soil"},{"key":"e_1_2_12_48_1","doi-asserted-by":"crossref","DOI":"10.1016\/j.fcr.2024.109642","article-title":"Complementarity and Competitive Trade\u2010Offs Enhance Forage Productivity, Nutritive Balance, Land and Water Use, and Economics in Legume\u2013Grass Intercropping","volume":"319","author":"Luo C. 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E.","year":"2022","journal-title":"Agricultural & Environmental Letters"},{"key":"e_1_2_12_54_1","volume-title":"Forage in Ruminant Nutrition","author":"Minson D. J.","year":"1990"},{"issue":"3","key":"e_1_2_12_55_1","doi-asserted-by":"crossref","first-page":"747","DOI":"10.2307\/2401901","article-title":"Solar Radiation and Productivity in Tropical Ecosystems","volume":"9","author":"Monteith J. L.","year":"1972","journal-title":"Journal of Applied Ecology"},{"issue":"3","key":"e_1_2_12_56_1","doi-asserted-by":"crossref","first-page":"301","DOI":"10.1111\/j.1365-2494.2011.00806.x","article-title":"Sustainable, Low\u2010Input, Warm\u2010Season Grass\u2013Legume Grassland Mixtures: Mission (Nearly) Impossible?","volume":"66","author":"Muir J. 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