Biological nitrogen fixation is a key driver of global primary production and climate. Decades of effort have repeatedly updated nitrogen fixation estimates for terrestrial and open ocean systems, yet other aquatic systems in between have largely been ignored. Here we present an evaluation of nitrogen fixation for inland and coastal waters. We demonstrate that water column and sediment nitrogen fixation is ubiquitous across these diverse aquatic habitats, with rates ranging six orders of magnitude. We conservatively estimate that, despite accounting for less than 10% of the global surface area, inland and coastal aquatic systems fix 40 (30 to 54) teragrams of nitrogen per year, equivalent to 15% of the nitrogen fixed on land and in the open ocean. Inland systems contribute more than half of this biological nitrogen fixation.

Aix Marseille Univ Université de Toulon CNRS IRD MIO UM 110 Marseille France

Atlantic Coastal Environmental Sciences Division United States Environmental Protection Agency Narragansett RI USA

Cary Institute of Ecosystem Studies Millbrook NY USA

Department of Aquatic Sciences and Assessment Swedish University of Agricultural Sciences Uppsala Sweden

Department of Biodiversity Earth and Environmental Science Drexel University Philadelphia PA USA

Department of Biological Sciences Michigan Technological University Houghton MI USA

Department of Biology Baylor University Waco TX USA

Department of Biology Boston University Boston MA USA

Department of Earth and Environment Boston University Boston MA USA

Department of Natural Science Hawai'i Pacific University Honolulu HI USA

Earth and Environmental Science CUNY Advanced Science Research Center Brooklyn College New York NY USA

Flathead Lake Biological Station University of Montana Polson MT USA

Institute of Environmental Studies Charles University Prague Czech Republic

National Oceanography Centre European Way Southampton UK

School for Environment and Sustainability University of Michigan Ann Arbor MI USA

School of Marine Science and Policy University of Delaware Lewes DE USA

School of Natural and Environmental Sciences Newcastle University Newcastle upon Tyne UK

School of the Environment Washington State University Richland WA USA

Turing Center for Living Systems Aix Marseille University Marseille France

Water Quality and Ecology Research Unit National Sedimentation Laboratory Agricultural Research Service United States Department of Agriculture Oxford MS USA

Water Resources Science University of Minnesota St Paul MN USA

Komentář v

Science. 2025 Jun 12;388(6752):1130-1131. doi: 10.1126/science.ady3520. PubMed

Zobrazit více v PubMed

Rucker HR, Kaçar B, Enigmatic evolution of microbial nitrogen fixation: insights from Earth’s past. [Preprint] (2024). 10.1016/j.tim.2023.03.011. PubMed DOI

Pi H-W, Lin J-J, Chen C-A, Wang P-H, Chiang Y-R, Huang C-C, Young C-C, Li W-H, Origin and evolution of nitrogen fixation in prokaryotes. Mol Biol Evol 39, msac181 (2022). PubMed PMC

Coale TH, Loconte V, Turk-Kubo KA, Vanslembrouck B, Mak WKE, Cheung S, Ekman A, Chen J-H, Hagino K, Takano Y, Nitrogen-fixing organelle in a marine alga. Science (1979) 384, 217–222 (2024). PubMed

Wen Z, Browning TJ, Cai Y, Dai R, Zhang R, Du C, Jiang R, Lin W, Liu X, Cao Z, Nutrient regulation of biological nitrogen fixation across the tropical western North Pacific. Sci Adv 8, eabl7564 (2022). PubMed PMC

Peng J, Dan L, Wang Y-P, Tang X, Yang X, Yang F, Lu X, Pak B, Role contribution of biological nitrogen fixation to future terrestrial net land carbon accumulation under warming condition at centennial scale. J Clean Prod 202, 1158–1166 (2018).

Gruber N, “The dynamics of the marine nitrogen cycle and its influence on atmospheric CO 2 variations” in The Ocean Carbon Cycle and Climate (Springer, 2004), pp. 97–148.

Zhang X, Ward BB, Sigman DM, Global nitrogen cycle: critical enzymes, organisms, and processes for nitrogen budgets and dynamics. Chem Rev 120, 5308–5351 (2020). PubMed

Tranvik L, Cole JJ, Prairie YT, The study of carbon in inland waters-from isolated ecosystems to players in the global carbon cycle. Limnol Oceanogr Lett 3, 41–48 (2018).

Rocher-Ros G, Stanley EH, Loken LC, Casson NJ, Raymond PA, Liu S, Amatulli G, Sponseller RA, Global methane emissions from rivers and streams. Nature 621, 530–535 (2023). PubMed PMC

Regnier P, Resplandy L, Najjar RG, Ciais P, The land-to-ocean loops of the global carbon cycle. Nature 603, 401–410 (2022). PubMed

Rosentreter JA, Laruelle GG, Bange HW, Bianchi TS, Busecke JJM, Cai W-J, Eyre BD, Forbrich I, Kwon EY, Maavara T, Coastal vegetation and estuaries are collectively a greenhouse gas sink. Nat Clim Chang 13, 579–587 (2023).

Cole JJ, Prairie YT, Caraco NF, McDowell WH, Tranvik LJ, Striegl RG, Duarte CM, Kortelainen P, Downing JA, Middelburg JJ, Plumbing the global carbon cycle: integrating inland waters into the terrestrial carbon budget. Ecosystems 10, 172–185 (2007).

Fowler D, Coyle M, Skiba U, Sutton MA, Cape JN, Reis S, Sheppard LJ, Jenkins A, Grizzetti B, Galloway N, Vitousek P, Leach A, Bouwman AF, Butterbach-bahl K, Dentener F, Stevenson D, Amann M, Voss M, Fowler D, The global nitrogen cycle in the twenty-first century. (2013). PubMed PMC

Voss M, Bange HW, Dippner JW, Middelburg JJ, Montoya JP, Ward B, The marine nitrogen cycle: recent discoveries, uncertainties and the potential relevance of climate change. Philosophical Transactions of the Royal Society B: Biological Sciences 368, 20130121 (2013). PubMed PMC

Marcarelli AM, Fulweiler RW, Scott JT, Nitrogen fixation: A poorly understood process along the freshwater‐marine continuum. Limnol Oceanogr Lett 7, 1–10 (2022). PubMed PMC

Brezonik PL, Harper CL, Nitrogen fixation in some anoxic lacustrine environments. Science (1979) 164, 1277–1279 (1969). PubMed

Keirn MA, Brezonik PL, Nitrogen fixation by bacteria in Lake Mize, Florida and in some lacustrine sediments. Limnol Oceanogr 16, 720–731 (1971).

Dugdale R, Dugdale V, Neess J, Goering J, Nitrogen fixation in lakes. Science (1979) 130, 859–860 (1959). PubMed

Goering JJ, Neess JC, Nitrogen fixation in two Wisconsin Lakes. Limnol Oceanogr 9, 530–539 (1964).

Howarth RW, Marino R, Lane J, Cole JJ, Nitrogen fixation in freshwater, estuarine, and marine ecosystems. 1. Rates and importance 1. Limnol Oceanogr 33, 669–687 (1988).

Scott JT, McCarthy MJ, Nitrogen fixation may not balance the nitrogen pool in lakes over timescales relevant to eutrophication management. Limnol Oceanogr 55, 1265–1270 (2010).

Grantz EM, Kogo A, Scott JT, Partitioning whole‐lake denitrification using in situ dinitrogen gas accumulation and intact sediment core experiments. Limnol Oceanogr 57, 925–935 (2012).

Eberhard EK, Techtmann SM, V Baxter C, Marcarelli AM, Spatial Heterogeneity of Nitrogen Fixation and Denitrification in Streams. Ecosystems 28, 18 (2025).

Wang S, Jiang L, Zhao Z, Chen Z, Wang J, Alain K, Cui L, Zhong Y, Peng Y, Lai Q, Chemolithoautotrophic diazotrophs dominate dark nitrogen fixation in mangrove sediments. ISME J 18, wrae119 (2024). PubMed PMC

Xin Y, Gao Q, Chen X, Sun S, Liu J, Gao H, Zhou J, Xia X, High biological N fixation potential dominated by heterotrophic diazotrophs in alpine permafrost rivers on the Qinghai‒Tibet Plateau. Water Res 264, 122239 (2024). PubMed

Grimm NB, Petrone KC, Nitrogen fixation in a desert stream ecosystem. Biogeochemistry 37, 33–61 (1997).

Patoine A, Graham MD, Leavitt PR, Spatial variation of nitrogen fixation in lakes of the northern Great Plains. Limnol Oceanogr 51, 1665–1677 (2006).

Fulweiler RW, More foxes than hedgehogs: the case for nitrogen fixation in coastal marine sediments. Global Biogeochem Cycles 37, e2023GB007777 (2023).

Bertics VJ, Löscher CR, Salonen I, Dale AW, Gier J, Schmitz RA, Treude T, Occurrence of benthic microbial nitrogen fixation coupled to sulfate reduction in the seasonally hypoxic Eckernförde Bay, Baltic Sea. Biogeosciences 10, 1243–1258 (2013).

Joshi HM, Tabita FR, A global two component signal transduction system that integrates the control of photosynthesis, carbon dioxide assimilation, and nitrogen fixation. Proc Natl Acad Sci U S A 93, 14515–20 (1996). PubMed PMC

Nixon SW, Eutrophication and the macroscope. Hydrobiologia 629, 5–19 (2009).

Fulweiler RW, Heiss EM, (Nearly) A Decade of Directly Measured Sediment N2 Fluxes: What Can Narragansett Bay Tell Us About the Global Ocean Nitrogen Budget? Oceanography, doi: 10.5670/oceanog.2014.22 (2014). DOI

Newell SE, McCarthy MJ, Gardner WS, Fulweiler RW, Sediment Nitrogen Fixation: a Call for Re-evaluating Coastal N Budgets. Estuaries and Coasts, doi: 10.1007/s12237-016-0116-y (2016). DOI

Larsson U, Hajdu S, Walve J, Elmgren R, Baltic Sea nitrogen fixation estimated from the summer increase in upper mixed layer total nitrogen. Limnol Oceanogr 46, 811–820 (2001).

Fulweiler RWW, Nixon SWW, Buckley BAA, Granger SLL, Reversal of the net dinitrogen gas flux in coastal marine sediments. Nature 448, 180–182 (2007). PubMed

Capone DG, Carpenter EJ, Nitrogen fixation in the marine environment. Science (1979) 217, 1140–1142 (1982). PubMed

Schlesinger WH,BES, Biogeochemistry An Analysis of Global Change (ed. 4th, 2020).

Fulweiler RW, Berberich ME, Rinehart SA, Taylor JM, Kelly MC, Ray NE, Oczkowski A, Benavides M, Church MJ, Loeks B, Newell S, Olofsson M, Oppong J, Roley SS, Vizza C, Wilson ST, Scott JT, Marcarelli AM, Global dataset of nitrogen fixation rates across inland and coastal waters based on a coordinated synthesis effort ver 2‥ [Preprint] (2025). PubMed PMC

Fulweiler RW, Berberich ME, Rinehart SA, Taylor JM, Kelly MC, Ray NE, Oczkowski A, Balint SJ, Geisser AH, Mahoney CR, A global dataset of nitrogen fixation rates across inland and coastal waters. Limnol Oceanogr Lett (2025). PubMed PMC

Vieillard AM, Newell SE, Thrush SF, Recovering from bias: A call for further study of underrepresented tropical and low‐nutrient estuaries. J Geophys Res Biogeosci 125, e2020JG005766 (2020).

Cheung HLS, Hillman JR, Pilditch CA, Savage C, Santos IR, Glud RN, Nascimento FJA, Thrush SF, Bonaglia S, Denitrification, anammox, and DNRA in oligotrophic continental shelf sediments. Limnol Oceanogr (2024).

Boyero L, Ramírez A, Dudgeon D, Pearson RG, Are tropical streams really different? J North Am Benthol Soc 28, 397–403 (2009).

Prentice IC, Modelling the demand for new nitrogen fixation by terrestrial ecosystems. Biogeosciences 14, 2003–2017 (2017).

Von Friesen LW, Riemann L, Nitrogen fixation in a changing Arctic Ocean: an overlooked source of nitrogen? Front Microbiol 11, 596426 (2020). PubMed PMC

Stewart WD, Fitzgerald GP, Burris RH, In situ studies on N2 fixation using the acetylene reduction technique. Proc Natl Acad Sci U S A 58 (1967). PubMed PMC

Flett RJ, Hamilton RD, Campbell NER, Aquatic acetylene reduction techniques: solutions to several problems. Can J Microbiol 22 (1976). PubMed

Sprott GD, Jarrell KF, Shaw KM, Knowles R, Acetylene as an inhibitor of methanogenic bacteria. Microbiology (N Y) 128, 2453–2462 (1982).

Fulweiler RW, Heiss EM, Rogener MK, Newell SE, LeCleir GR, Kortebein SM, Wilhelm SW, Examining the impact of acetylene on N-fixation and the active sediment microbial community. Front Microbiol 6, 1–9 (2015). PubMed PMC

Seitzinger SP, Garber JH, Nitrogen fixation and calibration of the. Mar. Ecol. Prog. Ser 37, 65–73 (1987).

Großkopf T, Mohr W, Baustian T, Schunck H, Gill D, Kuypers MMM, Lavik G, Schmitz R. a., Wallace DWR, LaRoche J, Doubling of marine dinitrogen-fixation rates based on direct measurements. Nature 488, 361–364 (2012). PubMed

Wilson ST, Böttjer D, Church MJ, Karl DM, Comparative assessment of nitrogen fixation methodologies, conducted in the oligotrophic north pacific ocean. Appl Environ Microbiol 78, 6516–6523 (2012). PubMed PMC

Gardner WS, McCarthy MJ, An S, Sobolev D, Sell KS, Brock D, Nitrogen fixation and dissimilatory nitrate reduction to ammonium (DNRA) support nitrogen dynamics in Texas estuaries. Limnol Oceanogr 51, 558–568 (2006).

Taylor JM, Andersen IM, Hoke AK, Kelly PT, Scott JT, In-situ N2: Ar ratios describe the balance between nitrogen fixation and denitrification in shallow eutrophic experimental lakes. Biogeochemistry 166, 283–301 (2023).

Tucker J, Giblin AE, Hopkinson CS, Kelsey SW, Howes BL, Response of benthic metabolism and nutrient cycling to reductions in wastewater loading to Boston Harbor, USA. Estuar Coast Shelf Sci 151, 54–68 (2014).

Boynton WR, Ceballos MAC, Hodgkins CLS, Liang D, Testa JM, Large-Scale Spatial and Temporal Patterns and Importance of Sediment–Water Oxygen and Nutrient Fluxes in the Chesapeake Bay Region. Estuaries and Coasts 46, 356–375 (2023).

Roberts BJ, Mulholland PJ, In-stream biotic control on nutrient biogeochemistry in a forested stream, West Fork of Walker Branch. J Geophys Res Biogeosci 112 (2007).

Newbold JW, Elwood JW, O’Neill RV, Sheldon AL, Phosphorus dynamics in a woodland stream ecosystem: a study of nutrient spiralling. Ecology 64 (1983).

Mulholland PJ, Newbold JD, Elwood JW, Ferren LA, Webster JR, Phosphorus spiralling in a woodland stream: seasonal variations. Ecology 66 (1985). PubMed

Chua EJ, Fulweiler RW, Capturing the rapid response of sediments to low-oxygen conditions with high temporal resolution gas concentration measurements. Front Environ Sci 10, 1028405 (2023).

Billaud VA, Nitrogen Fixation and the Utilization of Other Inorganic Nitrogen Sources in a Subarctic Lake. Journal of the Fisheries Research Board of Canada 25 (1968).

Hayes NM, Patoine A, Haig HA, Simpson GL, Swarbrick VJ, Wiik E, Leavitt PR, Spatial and temporal variation in nitrogen fixation and its importance to phytoplankton in phosphorus‐rich lakes. Freshw Biol 64, 269–283 (2019).

Marcarelli AM, Wurtsbaugh WA, Nitrogen fixation varies spatially and seasonally in linked stream-lake ecosystems. Biogeochemistry 94, 95–110 (2009).

Tilstra A, Van Hoytema N, Cardini U, Bednarz VN, Rix L, Naumann MS, Al-Horani FA, Wild C, Effects of water column mixing and stratification on planktonic primary production and dinitrogen fixation on a northern Red Sea coral reef. Front Microbiol 9, 2351 (2018). PubMed PMC

Wurtsbaugh WA, Horne AJ, Iron in eutrophic Clear Lake, California: its importance for algal nitrogen fixation and growth. Canadian Journal of Fisheries and Aquatic Sciences 40, 1419–1429 (1983).

Pedersen JN, Bombar D, Paerl RW, Riemann L, Diazotrophs and N2-fixation associated with particles in coastal estuarine waters. Front Microbiol 9, 2759 (2018). PubMed PMC

Hamersley MR, Turk KA, Leinweber A, Gruber N, Zehr JP, Gunderson T, Capone DG, Nitrogen fixation within the water column associated with two hypoxic basins in the Southern California Bight. Aquatic Microbial Ecology 63, 193–205 (2011).

Triska FJ, Sedell JR, Cromack K Jr, V Gregory S, McCorison FM, Nitrogen budget for a small coniferous forest stream. Ecol Monogr 54, 119–140 (1984).

Marcarelli AM, Wurtsbaugh WA, Nitrogen fixation varies spatially and seasonally in linked stream-lake ecosystems. Biogeochemistry 94, 95–110 (2009).

Rolando JL, Kolton M, Song T, Liu Y, Pinamang P, Conrad R, Morris JT, Konstantinidis KT, Kostka JE, Sulfur oxidation and reduction are coupled to nitrogen fixation in the roots of the salt marsh foundation plant Spartina alterniflora. Nat Commun 15, 3607 (2024). PubMed PMC

Cui J, Zhang M, Chen L, Zhang S, Luo Y, Cao W, Zhao J, Wang L, Jia Z, Bao Z, Methanotrophs contribute to nitrogen fixation in emergent macrophytes. Front Microbiol 13, 851424 (2022). PubMed PMC

Marino R, Hayn M, Howarth RW, Giblin AE, McGlathery KJ, Berg P, Nitrogen fixation associated with epiphytes on the seagrass Zostera marina in a temperate lagoon with moderate to high nitrogen loads. Biogeochemistry 166, 211–226 (2023).

Mohr W, Lehnen N, Ahmerkamp S, Marchant HK, Graf JS, Tschitschko B, Yilmaz P, Littmann S, Gruber-Vodicka H, Leisch N, Terrestrial-type nitrogen-fixing symbiosis between seagrass and a marine bacterium. Nature 600, 105–109 (2021). PubMed PMC

Finke LR, Seeley HW Jr, Nitrogen fixation (acetylene reduction) by epiphytes of freshwater macrophytes. Appl Environ Microbiol 36, 129–138 (1978). PubMed PMC

Šantrůčková H, Rejmánková E, Pivničková B, Snyder JM, Nutrient enrichment in tropical wetlands: shifts from autotrophic to heterotrophic nitrogen fixation. Biogeochemistry 101, 295–310 (2010).

McGlathery KJ, Risgaard-Petersen N, Christensen PB, Temporal and spatial variation in nitrogen fixation activity in the eelgrass Zostera marina rhizosphere. Mar Ecol Prog Ser 168, 245–258 (1998).

Welsh DT, Nitrogen fixation in seagrass meadows: regulation, plant–bacteria interactions and significance to primary productivity. Ecol Lett 3, 58–71 (2000).

Head WD, Carpenter EJ, Nitrogen fixation associated with the marine macroalga Codium fragile 1. Limnol Oceanogr 20, 815–823 (1975).

Raut Y, Capone DG, Macroalgal detrital systems: an overlooked ecological niche for heterotrophic nitrogen fixation. Environ Microbiol 23, 4372–4388 (2021). PubMed

Hamersley MR, Sohm JA, Burns JA, Capone DG, Nitrogen fixation associated with the decomposition of the giant kelp Macrocystis pyrifera. Aquat Bot 125, 57–63 (2015).

Zilius M, Bonaglia S, Broman E, Chiozzini VG, Samuiloviene A, Nascimento FJA, Cardini U, Bartoli M, N2 fixation dominates nitrogen cycling in a mangrove fiddler crab holobiont. Sci Rep 10, 13966 (2020). PubMed PMC

König S, Gros O, Heiden SE, Hinzke T, Thürmer A, Poehlein A, Meyer S, Vatin M, Tocny J, Ponnudurai R, Nitrogen fixation in a chemoautotrophic lucinid symbiosis. Nat Microbiol 2, 1–10 (2016). PubMed

Knapp AN, The sensitivity of marine N2 fixation to dissolved inorganic nitrogen. Front Microbiol 3, 1–14 (2012). PubMed PMC

Gier J, Sommer S, Löscher CR, Dale AW, Schmitz RA, Treude T, Nitrogen fixation in sediments along a depth transect through the Peruvian oxygen minimum zone. 4065–4080 (2016).

Vitousek PM, Menge DNL, Reed SC, Cleveland CC, Biological nitrogen fixation: rates, patterns and ecological controls in terrestrial ecosystems. Philosophical Transactions of the Royal Society B: Biological Sciences 368, 20130119 (2013). PubMed PMC

Cleveland CC, Townsend AR, Schimel DS, Fisher H, Howarth RW, Hedin LO, Perakis SS, Latty EF, Von Fischer JC, Elseroad A, Global patterns of terrestrial biological nitrogen (N2) fixation in natural ecosystems. Global Biogeochem Cycles 13, 623–645 (1999).

Zehr JP, Capone DG, Zehr JP, Capone DG, N 2 Fixation in Ocean Basins. Marine Nitrogen Fixation, 143–156 (2021).

Shao Z, Xu Y, Wang H, Luo W, Wang L, Huang Y, Agawin NSR, Ahmed A, Benavides M, Bentzon-Tilia M, Global oceanic diazotroph database version 2 and elevated estimate of global oceanic N 2 fixation. Earth Syst Sci Data 15, 3673–3709 (2023).

Tang W, Wang S, Fonseca-Batista D, Dehairs F, Gifford S, Gonzalez AG, Gallinari M, Planquette H, Sarthou G, Cassar N, Revisiting the distribution of oceanic N2 fixation and estimating diazotrophic contribution to marine production. Nat Commun 10, 831 (2019). PubMed PMC

Wang W-L, Moore JK, Martiny AC, Primeau FW, Convergent estimates of marine nitrogen fixation. Nature 566, 205–211 (2019). PubMed

Fulweiler R, Rinehart S, Taylor J, Kelly MC, Bereberich M, Ray N, Oczkowski A, Benavides M, Church M, Loeks B, Newell S, Olofsson M, Oppong JC, Roley S, Carmella V, Wilson S, Chowdhury S, Groffman P, Scott T, Marcarelli A, Supplementary Information for: Global Importance of Nitrogen Fixation Across Inland and Coastal Waters (v1.1.1). [Preprint] (2025). PubMed PMC

Fulweiler RW, Brown SM, Nixon SW, Jenkins BD, Evidence and a conceptual model for the co-occurrence of nitrogen fixation and denitrification in heterotrophic marine sediments. Mar Ecol Prog Ser 482, 57–68 (2013).

Laruelle GG, Dürr HH, Lauerwald R, Hartmann J, Slomp CP, Goossens N, Regnier PAG, Global multi-scale segmentation of continental and coastal waters from the watersheds to the continental margins. Hydrol Earth Syst Sci 17, 2029–2051 (2013).

Dürr HH, Laruelle GG, van Kempen CM, Slomp CP, Meybeck M, Middelkoop H, Worldwide typology of nearshore coastal systems: defining the estuarine filter of river inputs to the oceans. Estuaries and coasts 34, 441–458 (2011).

Richardson DC, Holgerson MA, Farragher MJ, Hoffman KK, King KBS, Alfonso MB, Andersen MR, Cheruveil KS, Coleman KA, Farruggia MJ, A functional definition to distinguish ponds from lakes and wetlands. Sci Rep 12, 10472 (2022). PubMed PMC

Cael BB, Seekell DA, The size-distribution of Earth’s lakes. Sci Rep 6, 29633 (2016). PubMed PMC

Andreadis KM, Schumann GJ, Pavelsky T, A simple global river bankfull width and depth database. Water Resour Res 49, 7164–7168 (2013).

Agawin NSR, Ferriol P, Cryer C, Alcon E, Busquets A, Sintes E, Vidal C, Moyà G, Significant nitrogen fixation activity associated with the phyllosphere of Mediterranean seagrass Posidonia oceanica: first report. Mar Ecol Prog Ser 551, 53–62 (2016).

Rosentreter JA, V Borges A, Deemer BR, Holgerson MA, Liu S, Song C, Melack J, Raymond PA, Duarte CM, Allen GH, Olefeldt D, Poulter B, Battin TI, Eyre BD, Half of global methane emissions come from highly variable aquatic ecosystem sources. Nat Geosci 14, 225–230 (2021).

Alder J, Putting the coast in the Sea Around Us Project (2003). http://seaaroundus.org/newsletter/Issue15.pdf.

Zhang Z, Fluet-Chouinard E, Jensen K, McDonald K, Hugelius G, Gumbricht T, Carroll M, Prigent C, Bartsch A, Poulter B, Development of the global dataset of wetland area and dynamics for methane modeling (WAD2M). Earth Syst Sci Data 13, 2001–2023 (2021).

Saunois M, Stavert AR, Poulter B, Bousquet P, Canadell JG, Jackson RB, Raymond PA, Dlugokencky EJ, Houweling S, Patra PK, The global methane budget 2000–2017. Earth System Science Data Discussions 2019, 1–136 (2019).

Pi X, Luo Q, Feng L, Xu Y, Tang J, Liang X, Ma E, Cheng R, Fensholt R, Brandt M, Mapping global lake dynamics reveals the emerging roles of small lakes. Nat Commun 13, 5777 (2022). PubMed PMC

Giri C, Ochieng E, Tieszen LL, Zhu Z, Singh A, Loveland T, Masek J, Duke N, Status and distribution of mangrove forests of the world using earth observation satellite data. Global ecology and biogeography 20, 154–159 (2011).

Bunting P, Rosenqvist A, Hilarides L, Lucas RM, Thomas N, Tadono T, Worthington TA, Spalding M, Murray NJ, Rebelo L-M, Global mangrove extent change 1996–2020: Global mangrove watch version 3.0. Remote Sens (Basel) 14, 3657 (2022).

Mcowen CJ, V Weatherdon L, Van Bochove J-W, Sullivan E, Blyth S, Zockler C, Stanwell-Smith D, Kingston N, Martin CS, Spalding M, A global map of saltmarshes. Biodivers Data J (2017). PubMed PMC

NEP-WCMC, F. Short, Global distribution of seagrasses (version 7.1). Seventh update to the data layer used in Green and Short (2003), UN Environment World Conservation Monitoring Centre. (2021).

McKenzie LJ, Nordlund LM, Jones BL, Cullen-Unsworth LC, Roelfsema C, Unsworth RKF, The global distribution of seagrass meadows. Environmental Research Letters 15, 074041 (2020).

Murray NJ, Phinn SR, DeWitt M, Ferrari R, Johnston R, Lyons MB, Clinton N, Thau D, Fuller RA, The global distribution and trajectory of tidal flats. Nature 565, 222–225 (2019). PubMed

Galloway JN, Bleeker A, Erisman JW, The human creation and use of reactive nitrogen: a global and regional perspective. Annu Rev Environ Resour 46, 255–288 (2021).

Tian H, Bian Z, Shi H, Qin X, Pan N, Lu C, Pan S, Tubiello FN, Chang J, Conchedda G, History of anthropogenic Nitrogen inputs (HaNi) to the terrestrial biosphere: A 5-arcmin resolution annual dataset from 1860 to 2019. Earth System Science Data Discussions 2022, 1–32 (2022).

Malagó A, Bouraoui F, Global anthropogenic and natural nutrient fluxes: from local to planetary assessments. Environmental Research Letters 16, 054074 (2021).

Galloway JN, Dentener FJ, Capone DG, Boyer EW, Howarth RW, Seitzinger SP, Asner GP, Cleveland CC, Green PA, Holland EA, Nitrogen cycles: past, present, and future. Biogeochemistry 70, 153–226 (2004).

Yu T, Zhuang Q, Modeling biological nitrogen fixation in global natural terrestrial ecosystems. Biogeosciences 17, 3643–3657 (2020).

Davies‐Barnard T, Friedlingstein P, The global distribution of biological nitrogen fixation in terrestrial natural ecosystems. Global Biogeochem Cycles 34, e2019GB006387 (2020).

Gruber N, Galloway JN, An Earth-system perspective of the global nitrogen cycle. Nature 451, 293 (2008). PubMed

Meyerholt J, Zaehle S, Smith MJ, Variability of projected terrestrial biosphere responses to elevated levels of atmospheric CO 2 due to uncertainty in biological nitrogen fixation. Biogeosciences 13, 1491–1518 (2016).

Wang Y, Houlton BZ, Nitrogen constraints on terrestrial carbon uptake: Implications for the global carbon‐climate feedback. Geophys Res Lett 36 (2009).

Luo Y-W, Doney SC, Anderson LA, Benavides M, Berman-Frank I, Bode A, Bonnet S, Boström KH, Böttjer D, Capone DG, Database of diazotrophs in global ocean: abundance, biomass and nitrogen fixation rates. Earth Syst Sci Data 4, 47–73 (2012).

Tang W, Li Z, Cassar N, Machine Learning Estimates of Global Marine Nitrogen Fixation. 717–730 (2019).

Dutkiewicz S, Ward BA, Monteiro F, Follows MJ, Interconnection of nitrogen fixers and iron in the Pacific Ocean: Theory and numerical simulations. Global Biogeochem Cycles 26 (2012).

Bianchi D, Dunne JP, Sarmiento JL, Galbraith ED, Data‐based estimates of suboxia, denitrification, and N2O production in the ocean and their sensitivities to dissolved O2. Global Biogeochem Cycles 26 (2012).

Gruber N, Sarmiento JL, Global patterns of marine nitrogen fixation and denitrification. Global Biogeochem Cycles 11, 235–266 (1997).

Brandes JA, Devol AH, A global marine‐fixed nitrogen isotopic budget: Implications for Holocene nitrogen cycling. Global Biogeochem Cycles 16, 61–67 (2002).

Deutsch C, Sarmiento JL, Sigman DM, Gruber N, Dunne JP, Spatial coupling of nitrogen inputs and losses in the ocean. Nature 445, 163–7 (2007). PubMed

Landolfi A, Koeve W, Dietze H, Kähler P, Oschlies A, A new perspective on environmental controls of marine nitrogen fixation. Geophys Res Lett 42, 4482–4489 (2015).

Jickells TD, Buitenhuis E, Altieri K, Baker AR, Capone D, Duce RA, Dentener F, Fennel K, Kanakidou M, LaRoche J, A reevaluation of the magnitude and impacts of anthropogenic atmospheric nitrogen inputs on the ocean. Global Biogeochem Cycles 31, 289–305 (2017).

Großkopf T, Mohr W, Baustian T, Schunck H, Gill D, Kuypers MMM, Lavik G, Schmitz R. a., Wallace DWR, LaRoche J, Doubling of marine dinitrogen-fixation rates based on direct measurements. Nature 488, 361–364 (2012). PubMed

Somes CJ, Oschlies A, On the influence of “non‐Redfield” dissolved organic nutrient dynamics on the spatial distribution of N2 fixation and the size of the marine fixed nitrogen inventory. Global Biogeochem Cycles 29, 973–993 (2015).

Global importance of nitrogen fixation across inland and coastal waters