N₂ fixation in the Mediterranean Sea related to the composition of the diazotrophic community and impact of dust under present and future environmental conditions

<p><span class="inline-formula">N₂</span> fixation rates were measured in the 0–1000 <span class="inline-formula">m</span> layer at 13 stations located in the open western and central Mediterranean Sea (MS) during the PEACETIME cruise (late spring 2017). While the spatial variability in <span class="inline-formula">N₂</span> fixation was not related to Fe, P nor N stocks, the surface composition of the diazotrophic community indicated a strong longitudinal gradient increasing eastward for the relative abundance of non-cyanobacterial diazotrophs (NCDs) (mainly <span class="inline-formula"><i>γ</i></span>-Proteobacteria) and conversely decreasing eastward for photo-heterotrophic group A (UCYN-A) (mainly UCYN-A1 and UCYN-A3), as did <span class="inline-formula">N₂</span> fixation rates. UCYN-A4 and UCYN-A3 were identified for the first time in the MS. The westernmost station influenced by Atlantic waters and characterized by highest stocks of N and P displayed a patchy distribution of diazotrophic activity with an exceptionally high rate in the euphotic layer of 72.1 <span class="inline-formula"><math xmlns="http://www.w3.org/1998/Math/MathML" id="M8" display="inline" overflow="scroll" dspmath="mathml"><mrow class="unit"><mi mathvariant="normal">nmol</mi><mspace linebreak="nobreak" width="0.125em"/><mi mathvariant="normal">N</mi><mspace width="0.125em" linebreak="nobreak"/><msup><mi mathvariant="normal">L</mi><mrow><mo>-</mo><mn mathvariant="normal">1</mn></mrow></msup><mspace width="0.125em" linebreak="nobreak"/><msup><mi mathvariant="normal">d</mi><mrow><mo>-</mo><mn mathvariant="normal">1</mn></mrow></msup></mrow></math><span><svg:svg xmlns:svg="http://www.w3.org/2000/svg" width="71pt" height="13pt" class="svg-formula" dspmath="mathimg" md5hash="329d8ea18d72e83e28e52b406b72b63f"><svg:image xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="bg-19-415-2022-ie00001.svg" width="71pt" height="13pt" src="bg-19-415-2022-ie00001.png"/></svg:svg></span></span>, which could support up to 19 % of primary production. At this station at 1 <span class="inline-formula">% PAR</span> (photosynthetically available radiation) depth, UCYN-A4 represented up to 94 % of the diazotrophic community. These in situ observations of greater relative abundance of UCYN-A at stations with higher nutrient concentrations and dominance of NCDs at more oligotrophic stations suggest that nutrient conditions – even in the nanomolar range – may determine the composition of diazotrophic communities and in turn <span class="inline-formula">N₂</span> fixation rates. The impact of Saharan dust deposition on <span class="inline-formula">N₂</span> fixation and diazotrophic communities was also investigated, under present and future projected conditions of temperature and pH during short-term (3–4 <span class="inline-formula">d</span>) experiments at three stations. New nutrients from simulated dust deposition triggered a significant stimulation of <span class="inline-formula">N₂</span> fixation (from 41 % to 565 %). The strongest increase in <span class="inline-formula">N₂</span> fixation was observed at the stations dominated by NCDs and did not lead on this short timescale to changes in the diazotrophic community composition. Under projected future conditions, <span class="inline-formula">N₂</span> fixation was either increased or unchanged; in that later case this was probably due to a too-low nutrient bioavailability or an increased grazing pressure. The future warming and acidification likely benefited NCDs (<i>Pseudomonas</i>) and<span id="page416"/> UCYN-A2, while disadvantaged UCYN-A3 without knowing which effect (alone or in combination) is the driver, especially since we do not know the temperature optima of these species not yet cultivated as well as the effect of acidification.</p>