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Abstract Biological dinitrogen (N2) fixation supplies nitrogen to the oceans, supporting primary productivity, and is carried out by some bacteria and archaea referred to as diazotrophs. Cyanobacteria are conventionally considered to be the major contributors to marine N2 fixation, but non-cyanobacterial diazotrophs (NCDs) have been shown to be distributed throughout ocean ecosystems. However, the biogeochemical significance of marine NCDs has not been demonstrated. This review synthesizes multiple datasets, drawing from cultivation-independent molecular techniques and data from extensive oceanic expeditions, to provide a comprehensive view into the diversity, biogeography, ecophysiology, and activity of marine NCDs. A NCD nifH gene catalog was compiled containing sequences from both PCR-based and PCR-free methods, identifying taxa for future studies. NCD abundances from a novel database of NCD nifH-based abundances were colocalized with environmental data, unveiling distinct distributions and environmental drivers of individual taxa. Mechanisms that NCDs may use to fuel and regulate N2 fixation in response to oxygen and fixed nitrogen availability are discussed, based on a metabolic analysis of recently available Tara Oceans expedition data. The integration of multiple datasets provides a new perspective that enhances understanding of the biology, ecology, and biogeography of marine NCDs and provides tools and directions for future research. 

Abstract. Marine diazotrophs convert dinitrogen (N2) gas intobioavailable nitrogen (N), supporting life in the global ocean. In 2012, thefirst version of the global oceanic diazotroph database (version 1) waspublished. Here, we present an updated version of the database (version 2),significantly increasing the number of in situ diazotrophic measurements from13 565 to 55 286. Data points for N2 fixation rates, diazotrophic cellabundance, and nifH gene copy abundance have increased by 184 %, 86 %, and809 %, respectively. Version 2 includes two new data sheets for the nifH genecopy abundance of non-cyanobacterial diazotrophs and cell-specific N2fixation rates. The measurements of N2 fixation rates approximatelyfollow a log-normal distribution in both version 1 and version 2. However,version 2 considerably extends both the left and right tails of thedistribution. Consequently, when estimating global oceanic N2 fixationrates using the geometric means of different ocean basins, version 1 andversion 2 yield similar rates (43–57 versus 45–63 Tg N yr−1; rangesbased on one geometric standard error). In contrast, when using arithmeticmeans, version 2 suggests a significantly higher rate of 223±30 Tg N yr−1 (mean ± standard error; same hereafter) compared to version 1(74±7 Tg N yr−1). Specifically, substantial rate increases areestimated for the South Pacific Ocean (88±23 versus 20±2 Tg N yr−1), primarily driven by measurements in the southwestern subtropics,and for the North Atlantic Ocean (40±9 versus 10±2 Tg N yr−1). Moreover, version 2 estimates the N2 fixation rate in theIndian Ocean to be 35±14 Tg N yr−1, which could not be estimatedusing version 1 due to limited data availability. Furthermore, a comparisonof N2 fixation rates obtained through different measurement methods atthe same months, locations, and depths reveals that the conventional15N2 bubble method yields lower rates in 69 % cases compared tothe new 15N2 dissolution method. This updated version of thedatabase can facilitate future studies in marine ecology andbiogeochemistry. The database is stored at the Figshare repository(https://doi.org/10.6084/m9.figshare.21677687; Shao etal., 2022).