Articles | Volume 23, issue 17
https://doi.org/10.5194/bg-23-6317-2026
© Author(s) 2026. This work is distributed under
the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
https://doi.org/10.5194/bg-23-6317-2026
© Author(s) 2026. This work is distributed under
the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
A boost on the final stretch: intense river metabolism and wetland discharge increase aquatic CO2 dynamics in the Danube Delta
Marie-Sophie Maier
Department of Surface Waters, Research and Management, Eawag, 6047 Kastanienbaum, Switzerland
Department of Environmental Systems Science, ETH Zurich, 8092 Zurich, Switzerland
Bernhard Wehrli
CORRESPONDING AUTHOR
Department of Surface Waters, Research and Management, Eawag, 6047 Kastanienbaum, Switzerland
Department of Environmental Systems Science, ETH Zurich, 8092 Zurich, Switzerland
Cristian R. Teodoru
National Research – Development Institute for Marine Geology and Geoecology (GeoEcoMar), Dimitrie Onciu Street 23–25, 024053 Bucharest, Romania
Cited articles
Abril, G. and Borges, A. V.: Ideas and perspectives: Carbon leaks from flooded land: do we need to replumb the inland water active pipe?, Biogeosciences, 16, 769–784, https://doi.org/10.5194/bg-16-769-2019, 2019.
Aho, K. S., Hosen, J. D., Logozzo, L. A., McGillis, W. R., and Raymond, P. A.: Highest rates of gross primary productivity maintained despite CO2 depletion in a temperate river network, Limnology and Oceanography Letters, 6, 200–206, https://doi.org/10.1002/lol2.10195, 2021.
Battin, T. J., Lauerwald, R., Bernhardt, E. S., Bertuzzo, E., Gener, L. G., Hall, R. O., Jr., Hotchkiss, E. R., Maavara, T., Pavelsky, T. M., Ran, L., Raymond, P., Rosentreter, J. A., and Regnier, P.: River ecosystem metabolism and carbon biogeochemistry in a changing world, Nature, 613, 449–459, https://doi.org/10.1038/s41586-022-05500-8, 2023.
Bernhardt, E. S., Heffernan, J. B., Grimm, N. B., Stanley, E. H., Harvey, J. W., Arroita, M., Appling, A. P., Cohen, M. J., McDowell, W. H., Hall, R. O., Read, J. S., Roberts, B. J., Stets, E. G., and Yackulic, C. B.: The metabolic regimes of flowing waters, Limnol. Oceanogr., 63, https://doi.org/10.1002/lno.10726, 2017.
Bondar, C.: Referitor la alimentarea si tranzitul apelor Dunarii prin interiorul deltei, Analele ICPDD, III/2, 259–261, 1994.
Borges, A. V., Darchambeau, F., Lambert, T., Morana, C., Allen, G. H., Tambwe, E., Toengaho Sembaito, A., Mambo, T., Nlandu Wabakhangazi, J., Descy, J.-P., Teodoru, C. R., and Bouillon, S.: Variations in dissolved greenhouse gases (CO2, CH4, N2O) in the Congo River network overwhelmingly driven by fluvial-wetland connectivity, Biogeosciences, 16, 3801–3834, https://doi.org/10.5194/bg-16-3801-2019, 2019.
Cai, W. J. and Wang, Y.: The chemistry, fluxes, and sources of carbon dioxide in the estuarine waters of the Satilla and Altamaha Rivers, Georgia, Limnol. Oceanogr., 43, 657–668, 1998.
Canning, A., Wehrli, B., and Körtzinger, A.: Methane in the Danube Delta: the importance of spatial patterns and diel cycles for atmospheric emission estimates, Biogeosciences, 18, 3961–3979, https://doi.org/10.5194/bg-18-3961-2021, 2021.
Cole, J. J., Prairie, Y. T., Caraco, N. F., McDowell, W. H., Tranvik, L. J., Striegl, R. G., Duarte, C. M., Kortelainen, P., Downing, J. A., Middelburg, J. J., and Melack, J.: Plumbing the Global Carbon Cycle: Integrating Inland Waters into the Terrestrial Carbon Budget, Ecosystems, 10, 172–185, https://doi.org/10.1007/s10021-006-9013-8, 2007.
Coops, H., Buijse, L. L., Buijse, A. D., Constantinescu, A., Covaliov, S., Hanganu, J., Ibelings, B. W., Menting, G., Navodaru, I., Oosterberg, W., Staras, M., and Török, L.: Trophic gradients in a large-river Delta: ecological structure determined by connectivity gradients in the Danube Delta (Romania), River Res. Appl., 24, 698–709, https://doi.org/10.1002/rra.1136, 2008.
Csagoly, P., Magnin, G., and Hulea, O.: Lower Danube Green Corridor, in: The Wetland Book., edited by: Finlayson, C. M., Milton, G., Prentice, R., and Davidson, N., Springer, Dodrecht, https://doi.org/10.1007/978-94-007-4001-3_251, 2018.
Dalmagro, H. J., Lathuilliere, M. J., Hawthorne, I., Morais, D. D., Pinto, O. B., Couto, E. G., and Johnson, M. S.: Carbon biogeochemistry of a flooded Pantanal forest over three annual flood cycles, Biogeochemistry, 139, 1–18, https://doi.org/10.1007/s10533-018-0450-1, 2018.
Degens, E., Kempe, S., and Ittekkot, V.: Monitoring carbon in world rivers, Environment: Science and Policy for Sustainable Development, 26, 29–33, https://doi.org/10.1080/00139157.1984.9932533, 1984.
DelVecchia, A. G., Rhea, S., Aho, K. S., Stanley, E. H., Hotchkiss, E. R., Carter, A., and Bernhardt, E. S.: Variability and drivers of CO2, CH4, and N2O concentrations in streams across the United States, Limnol. Oceanogr., 68, 394–408, https://doi.org/10.1002/lno.12281, 2023.
Diamond, J. S., Truong, A. N., Abril, G., Bertuzzo, E., Chanudet, V., Lamouroux, R., and Moatar, F.: Inorganic carbon dynamics and their relation to autotrophic community regime shift over three decades in a large, alkaline river, Limnol. Oceanogr., 70, 1122–1136, https://doi.org/10.1002/lno.70016, 2025.
Dodds, W. K., Veach, A. M., Ruffing, C. M., Larson, D. M., Fischer, J. L., and Costigan, K. H.: Abiotic controls and temporal variability of river metabolism: multiyear analyses of Mississippi and Chattahoochee River data, Freshw. Sci., 32, 1073–1087, https://doi.org/10.1899/13-018.1, 2013.
Durisch-Kaiser, E., Doberer, A., Reutimann, J., Pavel, A., Balan, S., Radan, S., and Wehrli, B.: Organic matter governs N and P balance in Danube Delta lakes, Aquat. Sci., 73, 21–33, https://doi.org/10.1007/s00027-010-0156-5, 2010.
Dybdahl, A. K., Holmboe, C. M. H., Baattrup-Pedersen, A., Riis, T., and Pastor, A.: Temperature dependence of biofilm metabolism in a lowland stream, Freshw. Sci., 43, 277–287, https://doi.org/10.1086/731873, 2024.
Gatescu, P.: The Danube Delta: Geographical Characteristics and Ecological Recovery, GeoJournal, 29, 57–67, https://link.springer.com/article/10.1007/BF00806866 (last access: 10 September 2026), 1993.
Gomez-Baggethun, E., Tudor, M., Doroftei, M., Covaliov, S., Nastase, A., Onara, D. F., Mierla, M., Marinov, M., Dorosencu, A. C., Lupu, G., Teodorof, L., Tudor, I. M., Kohler, B., Museth, J., Aronsen, E., Johnsen, S. I., Ibram, O., Marin, E., Craciun, A., and Cioaca, E.: Changes in ecosystem services from wetland loss and restoration: An ecosystem assessment of the Danube Delta (1960–2010), Ecosyst. Serv., 39, 17, https://doi.org/10.1016/j.ecoser.2019.100965, 2019.
Gomez-Gener, L., Rocher-Ros, G., Battin, T., Cohen, M. J., Dalmagro, H. J., Dinsmore, K. J., Drake, T. W., Duvert, C., Enrich-Prast, A., Horgby, A., Johnson, M. S., Kirk, L., Machado-Silva, F., Marzolf, N. S., McDowell, M. J., McDowell, W. H., Miettinen, H., Ojala, A. K., Peter, H., Pumpanen, J., Ran, L. S., Riveros-Iregui, D. A., Santos, I. R., Six, J., Stanley, E. H., Wallin, M. B., White, S. A., and Sponseller, R. A.: Global carbon dioxide efflux from rivers enhanced by high nocturnal emissions, Nat. Geosci., 14, 289–294, https://doi.org/10.1038/s41561-021-00722-3, 2021.
Haque, M. M., Begum, M. S., Nayna, O. K., Tareq, S. M., and Park, J.-H.: Seasonal shifts in diurnal variations of CO2 and O2 in the lower Ganges River, Limnology and Oceanography Letters, 7, 191–201, https://doi.org/10.1002/lol2.10246, 2022.
Hastie, A., Lauerwald, R., Ciais, P., and Regnier, P.: Aquatic carbon fluxes dampen the overall variation of net ecosystem productivity in the Amazon basin: An analysis of the interannual variability in the boundless carbon cycle, Glob. Change Biol., 25, 2094–2111, https://doi.org/10.1111/gcb.14620, 2019.
Hemes, K. S., Chamberlain, S. D., Eichelmann, E., Knox, S. H., and Baldocchi, D. D.: A Biogeochemical Compromise: The High Methane Cost of Sequestering Carbon in Restored Wetlands, Geophys. Res. Lett., 45, 6081–6091, https://doi.org/10.1029/2018gl077747, 2018.
Hotchkiss, E. R., Hall Jr., R. O., Sponseller, R. A., Butman, D., Klaminder, J., Laudon, H., Rosvall, M., and Karlsson, J.: Sources of and processes controlling CO2 emissions change with the size of streams and rivers, Nat. Geosci., 8, 696–699, https://doi.org/10.1038/ngeo2507, 2015.
Huertas, I. E., Flecha, S., Figuerola, J., Costas, E., and Morris, E. P.: Effect of hydroperiod on CO2 fluxes at the air-water interface in the Mediterranean coastal wetlands of Donana, J. Geophys. Res.-Biogeo., 122, 1615–1631, https://doi.org/10.1002/2017JG003793, 2017.
Jankowski, K., Schindler, D. E., and Lisi, P. J.: Temperature sensitivity of community respiration rates in streams is associated with watershed geomorphic features, Ecology, 95, 2707–2714, https://doi.org/10.1890/14-0608.1, 2014.
Kathilankal, J. C., O'Halloran, T. L., Schmidt, A., Hanson, C. V., and Law, B. E.: Development of a semi-parametric PAR (Photosynthetically Active Radiation) partitioning model for the United States, version 1.0, Geosci. Model Dev., 7, 2477–2484, https://doi.org/10.5194/gmd-7-2477-2014, 2014.
Li, J. J., Yuan, J. J., Ciais, P., Kang, H. J., Freeman, C., Huang, Y. Y., Dong, Y. H., Liu, D. Y., Li, Y., and Ding, W. X.: Two decades of improved wetland carbon sequestration in northern mid-to-high latitudes are offset by tropical and southern declines, Nature Ecology and Evolution, 9, https://doi.org/10.1038/s41559-025-02809-1, 2025.
Lu, W. Z., Xiao, J. F., Liu, F., Zhang, Y., Liu, C. A., and Lin, G. H.: Contrasting ecosystem CO2 fluxes of inland and coastal wetlands: a meta-analysis of eddy covariance data, Glob. Change Biol., 23, 1180–1198, https://doi.org/10.1111/gcb.13424, 2017.
Maier, M.-S., Teodoru, C. R., and Wehrli, B.: Spatio-temporal variations in lateral and atmospheric carbon fluxes from the Danube Delta, Biogeosciences, 18, 1417–1437, https://doi.org/10.5194/bg-18-1417-2021, 2021.
Maier, M.-S., Canning, A. R., Brennwald, M. S., Teodoru, C. R., and Wehrli, B.: Spatial Mapping of Dissolved Gases in the Danube Delta Reveals Intense Plant-Mediated Gas Transfer, Frontiers in Environmental Science, 10, https://doi.org/10.3389/fenvs.2022.838126, 2022.
Marzolf, N. S., Small, G. E., Oviedo-Vargas, D., Ganong, C. N., Duff, J. H., Ramirez, A., Pringle, C. M., Genereux, D. P., and Ardon, M.: Partitioning inorganic carbon fluxes from paired O2–CO2 gas measurements in a Neotropical headwater stream, Costa Rica, Biogeochemistry, 160, 259–273, https://doi.org/10.1007/s10533-022-00954-4, 2022.
Mulholland, P. J., Fellows, C. S., Tank, J. L., Grimm, N. B., Webster, J. R., Hamilton, S. K., Martí, E., Ashkenas, L., Bowden, W. B., Dodds, W. K., McDowell, W. H., Paul, M. J., and Peterson, B. J.: Inter-biome comparison of factors controlling stream metabolism, Freshwater Biol., 46, 1503–1517, https://doi.org/10.1046/j.1365-2427.2001.00773.x, 2001.
Nguyen, A. T., Abril, G., Diamond, J. S., Lamouroux, R., Martinet, C., and Moatar, F.: Multidecadal trends in CO2 evasion and aquatic metabolism in a large temperate river, Biogeosciences, 22, 4923–4951, https://doi.org/10.5194/bg-22-4923-2025, 2025.
Oosterberg, W., Staras, M., Bodgdan, L., Buijse, A. D., Constantinescu, A., Coops, H., Hanganu, J., Ibelings, B. W., Menting, G. A. M., and Navodaru, I.: Ecological gradients in the Danube Delta lakes: present state and man-induced changes, RIZA Rijkswaterstaat, NL, ISBN 90.369.5309x, 2000.
Oprea, A., Sîrbu, C., Doroftei, M., and Covaliov, S.: New Contributions to Vegetation Knowledge of the Danube Delta, Romania (I), Journal of Plant Development, 31, 159–181, https://doi.org/10.47743/jpd.2024.31.1.958, 2024.
Perkins, D. M., Yvon-Durocher, G., Demars, B. O. L., Reiss, J., Pichler, D. E., Friberg, N., Trimmer, M., and Woodward, G.: Consistent temperature dependence of respiration across ecosystems contrasting in thermal history, Glob. Change Biol., 18, 1300–1311, https://doi.org/10.1111/j.1365-2486.2011.02597.x, 2012.
Rabaey, J. S., Holgerson, M. A., Richardson, D. C., Andersen, M. R., Bansal, S., Bortolotti, L. E., Cotner, J. B., Hornbach, D. J., Martinsen, K. T., Moody, E. K., and Schloegel, O. F.: Freshwater Biogeochemical Hotspots: High Primary Production and Ecosystem Respiration in Shallow Waterbodies, Geophys. Res. Lett., 51, https://doi.org/10.1029/2023gl106689, 2024.
Raymond, P. A., Hartmann, J., Lauerwald, R., Sobek, S., McDonald, C., Hoover, M., Butman, D., Striegl, R., Mayorga, E., Humborg, C., Kortelainen, P., Durr, H., Meybeck, M., Ciais, P., and Guth, P.: Global carbon dioxide emissions from inland waters, Nature, 503, 355–359, https://doi.org/10.1038/nature12760, 2013.
Reiman, J. H. and Xu, Y. J.: Dissolved carbon export and CO2 outgassing from the lower Mississippi River – Implications of future river carbon fluxes, J. Hydrol., 578, https://doi.org/10.1016/j.jhydrol.2019.124093, 2019.
Richardson, J. L., Desai, A. R., Thom, J., Lindgren, K., Laudon, H., Peichl, M., Nilsson, M., Campeau, A., Järveoja, J., Hawman, P., Mishra, D. R., Smith, D., D'Acunha, B., Knox, S. H., Ng, D., Johnson, M. S., Blackstock, J., Malone, S. L., Oberbauer, S. F., Detto, M., Wickland, K. P., Forbrich, I., Weston, N., Hung, J. K. Y., Edgar, C., Euskirchen, E. S., Bret-Harte, S., Dobkowski, J., Kling, G., Kane, E. S., Badiou, P., Bogard, M., Bohrer, G., O'Halloran, T., Ritson, J., Arias-Ortiz, A., Baldocchi, D., Oikawa, P., Shahan, J., and Matsumura, M.: On the Relationship Between Aquatic CO2 Concentration and Ecosystem Fluxes in Some of the World's Key Wetland Types, Wetlands, 44, https://doi.org/10.1007/s13157-023-01751-x, 2024.
Rocher-Ros, G., Gomez-Gener, L., Jativa, C., Lannergård, E. E., Laudon, H., Lupon, A., Martí, E., Peñarroya, X., Sponseller, R. A., and Bernal, S.: Emerging patterns of CO2 : O2 dynamics in rivers and their link to ecosystem carbon processing, Limnology and Oceanography Letters, https://doi.org/10.1002/lol2.70057, 2025.
Rode, M., Wade, A. J., Cohen, M. J., Hensley, R. T., Bowes, M. J., Kirchner, J. W., Arhonditsis, G. B., Jordan, P., Kronvang, B., Halliday, S. J., Skeffington, R. A., Rozemeijer, J. C., Aubert, A. H., Rinke, K., and Jomaa, S.: Sensors in the Stream: The High-Frequency Wave of the Present, Environ. Sci. Technol., 50, 10297–10307, https://doi.org/10.1021/acs.est.6b02155, 2016.
Romanova, Y., Shakirzanova, Z., Ovcharuk, V., Todorova, O., Medvedieva, I., and Ivanchenko, A.: Temporal variation of water discharges in the lower course of the Danube River across the area from Reni to Izmail under the influence of natural and anthropogenic factors, Energetika, 65, 144–160, https://doi.org/10.6001/energetika.v65i2-3.4108, 2019.
Rosentreter, J. A., Laruelle, G. G., Bange, H. W., Bianchi, T. S., Busecke, J. J. M., Cai, W.-J., Eyre, B. D., Forbrich, I., Kwon, E. Y., Maavara, T., Moosdorf, N., Najjar, R. G., Sarma, V. V. S. S., Van Dam, B., and Regnier, P.: Coastal vegetation and estuaries are collectively a greenhouse gas sink, Nat. Clim. Change, 13, 579–587, https://doi.org/10.1038/s41558-023-01682-9, 2023.
Sabater, S., Timoner, X., Borrego, C., and Acuña, V.: Stream Biofilm Responses to Flow Intermittency: From Cells to Ecosystems, Frontiers in Environmental Science, 4, https://doi.org/10.3389/fenvs.2016.00014, 2016.
Shangguan, Q. P., Degrandpre, M. D., Hall, R. O., Jr., and Payn, R. A.: Freshwater carbonate buffering revisited, Limnology and Oceanography Letters, 10, 619–635, https://doi.org/10.1002/lol2.70047, 2025.
Solano, V., Duvert, C., Birkel, C., Maher, D. T., Garcia, E. A., and Hutley, L. B.: Stream respiration exceeds CO2 evasion in a low-energy, oligotrophic tropical stream, Limnol. Oceanogr., 68, 1132–1146, https://doi.org/10.1002/lno.12334, 2023.
Tschikof, M., Gericke, A., Venohr, M., Weigelhofer, G., Bondar-Kunze, E., Kaden, U. S., and Hein, T.: The potential of large floodplains to remove nitrate in river basins – The Danube case, Sci Total Environ., 843, https://doi.org/10.1016/j.scitotenv.2022.156879, 2022.
Vachon, D., Sadro, S., Bogard, M. J., Lapierre, J. F., Baulch, H. M., Rusak, J. A., Denfeld, B. A., Laas, A., Klaus, M., Karlsson, J., Weyhenmeyer, G. A., and Giorgio, P. A.: Paired O2–CO2 measurements provide emergent insights into aquatic ecosystem function, Limnology and Oceanography Letters, 5, 287–294, https://doi.org/10.1002/lol2.10135, 2020.
van der Knaap, J., Harpenslager, S. F., Aben, R. C. H., Weideveld, S. T. J., van Giersbergen, Q., van Dijk, G., Wintjen, P., Buzacott, A. J. V., Fritz, C., Kruijt, B., and Kosten, S.: Disproportionately High Contribution of Ditches to Landscape Greenhouse Gas Emissions in Drained Peatlands, Ecosystems, 28, https://doi.org/10.1007/s10021-025-01005-3, 2025.
van Heuven, S., Pierrot, D., Rae, J. W. B., Lewis, E., and Wallace, D. W. R.: MATLAB Program Developed for CO2 System Calculations, Carbon Dioxide Information Analysis Center, Oak Ridge National Laboratory, U. S. Department of Energy, Oak Ridge, Tennessee, https://doi.org/10.3334/CDIAC/otg.CO2SYS_MATLAB_v1.1, 2011.
Vorobyev, S. N., Kolesnichenko, L. G., Kolesnichenko, Y., Prokushkin, A. S., Lugovaya-Dolmatova, A., Karlsson, J., and Pokrovsky, O. S.: Floodplain carbon dioxide emissions strongly exceed those of the main river stem: A case study of the Ob River, western Siberia, J. Hydrol., 638, https://doi.org/10.1016/j.jhydrol.2024.131468, 2024.
Wehrli, B.: bernhardwehrli/CO2-O2-stat: CO2 dynamics in the Danube Delta (Versions O2_CO2_statistics), Zenodo [code], https://doi.org/10.5281/zenodo.21456491, 2026.
Wehrli, B., Maier, M.-S., and Teodoru, C. R.: CO2-O2 timeseries Danube Delta, Research Collection ETH Zurich [data set], https://doi.org/10.3929/ethz-c-000786936, 2025.
Weiss, R. F.: Carbon dioxide in water and seawater: the solubility of a non-ideal gas, Mar. Chem., 2, 203–215, https://doi.org/10.1016/0304-4203(74)90015-2, 1974.
Xue, H., Ding, H., Han, X. K., Lang, Y. C., Wang, T. J., Li, P., Qiao, M. R., Liu, D. D., Liu, Z. H., and Liu, C. Q.: Ditches as key players in carbon emissions in managed Phragmites-dominated wetland, J. Hydrol., 647, https://doi.org/10.1016/j.jhydrol.2024.132355, 2025.
Zou, J. Y., Ziegler, A. D., Chen, D. L., McNicol, G., Ciais, P., Jiang, X., Zheng, C. M., Wu, J., Wu, J., Lin, Z. Y., He, X. Y., Brown, L. E., Holden, J., Zhang, Z. T., Ramchunder, S. J., Chen, A. P., and Zeng, Z. Z.: Rewetting global wetlands effectively reduces major greenhouse gas emissions, Nat. Geosci., 15, 627–632, https://doi.org/10.1038/s41561-022-00989-0, 2022.
Zuijdgeest, A., Baumgartner, S., and Wehrli, B.: Hysteresis effects in organic matter turnover in a tropical floodplain during a flood cycle, Biogeochemistry, 131, 49–63, https://doi.org/10.1007/s10533-016-0263-z, 2016.
Zurbrügg, R., Wamulume, J., Kamanga, R., Wehrli, B., and Senn, D. B.: River-floodplain exchange and its effects on the fluvial oxygen regime in a large tropical river system (Kafue Flats, Zambia), J. Geophys. Res.-Biogeo., 117, https://doi.org/10.1029/2011jg001853, 2012.
Short summary
Monitoring stations in the Danube Delta revealed two orders of magnitude difference in the intensity of oxygen and carbon dioxide cycles. Biological processes were driving more intense day-night cycles in delta channels compared to river reaches and were mainly controlled by changes in temperature and cloud cover. Wetland discharge added dissolved CO2 from anaerobic processes and caused higher emission rates in delta channels and downstream river reaches.
Monitoring stations in the Danube Delta revealed two orders of magnitude difference in the...
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