Articles | Volume 10, issue 7
https://doi.org/10.5194/bg-10-4767-2013
© Author(s) 2013. This work is distributed under
the Creative Commons Attribution 3.0 License.
the Creative Commons Attribution 3.0 License.
https://doi.org/10.5194/bg-10-4767-2013
© Author(s) 2013. This work is distributed under
the Creative Commons Attribution 3.0 License.
the Creative Commons Attribution 3.0 License.
Research article
| 
15 Jul 2013
Research article |
| 15 Jul 2013
The role of benthic foraminifera in the benthic nitrogen cycle of the Peruvian oxygen minimum zone
N. Glock
Sonderforschungsbereich 754, Christian-Albrechts-University Kiel, Climate-Biogeochemistry Interactions in the Tropical Ocean, Kiel, Germany
GEOMAR, Helmholtz-Zentrum für Ozeanforschung Kiel, Wischhofstr. 1–3, 24148 Kiel, Germany
J. Schönfeld
GEOMAR, Helmholtz-Zentrum für Ozeanforschung Kiel, Wischhofstr. 1–3, 24148 Kiel, Germany
A. Eisenhauer
GEOMAR, Helmholtz-Zentrum für Ozeanforschung Kiel, Wischhofstr. 1–3, 24148 Kiel, Germany
C. Hensen
GEOMAR, Helmholtz-Zentrum für Ozeanforschung Kiel, Wischhofstr. 1–3, 24148 Kiel, Germany
J. Mallon
Sonderforschungsbereich 754, Christian-Albrechts-University Kiel, Climate-Biogeochemistry Interactions in the Tropical Ocean, Kiel, Germany
present address: Geoteam Schaffer, Gärtnerstrasse 38, 70329 Stuttgart, Gemany
S. Sommer
GEOMAR, Helmholtz-Zentrum für Ozeanforschung Kiel, Wischhofstr. 1–3, 24148 Kiel, Germany
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The biodiversity and abundance of benthic foraminifera tend to increase with distance within a transect from the Rainbow hydrothermal vent field. Miliolids dominate closer to the vents and may be better adapted to the potentially hydrothermal conditions than hyaline and agglutinated species. The reason for this remains unclear, but there are indications that elevated trace-metal concentrations in the porewater and intrusion of acidic hydrothermal fluids could have an influence on the foraminifera.
Babette A.A. Hoogakker, Catherine Davis, Yi Wang, Stephanie Kusch, Katrina Nilsson-Kerr, Dalton S. Hardisty, Allison Jacobel, Dharma Reyes Macaya, Nicolaas Glock, Sha Ni, Julio Sepúlveda, Abby Ren, Alexandra Auderset, Anya V. Hess, Katrin J. Meissner, Jorge Cardich, Robert Anderson, Christine Barras, Chandranath Basak, Harold J. Bradbury, Inda Brinkmann, Alexis Castillo, Madelyn Cook, Kassandra Costa, Constance Choquel, Paula Diz, Jonas Donnenfield, Felix J. Elling, Zeynep Erdem, Helena L. Filipsson, Sebastián Garrido, Julia Gottschalk, Anjaly Govindankutty Menon, Jeroen Groeneveld, Christian Hallmann, Ingrid Hendy, Rick Hennekam, Wanyi Lu, Jean Lynch-Stieglitz, Lélia Matos, Alfredo Martínez-García, Giulia Molina, Práxedes Muñoz, Simone Moretti, Jennifer Morford, Sophie Nuber, Svetlana Radionovskaya, Morgan Reed Raven, Christopher J. Somes, Anja S. Studer, Kazuyo Tachikawa, Raúl Tapia, Martin Tetard, Tyler Vollmer, Xingchen Wang, Shuzhuang Wu, Yan Zhang, Xin-Yuan Zheng, and Yuxin Zhou
Biogeosciences, 22, 863–957, https://doi.org/10.5194/bg-22-863-2025, https://doi.org/10.5194/bg-22-863-2025, 2025
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Joachim Schönfeld, Nicolaas Glock, Irina Polovodova Asteman, Alexandra-Sophie Roy, Marié Warren, Julia Weissenbach, and Julia Wukovits
J. Micropalaeontol., 42, 171–192, https://doi.org/10.5194/jm-42-171-2023, https://doi.org/10.5194/jm-42-171-2023, 2023
Short summary
Short summary
Benthic organisms show aggregated distributions due to the spatial heterogeneity of niches or food. We analysed the distribution of Globobulimina turgida in the Gullmar Fjord, Sweden, with a data–model approach. The population densities did not show any underlying spatial structure but a random log-normal distribution. A temporal data series from the same site depicted two cohorts of samples with high or low densities, which represent hypoxic or well-ventilated conditions in the fjord.
Nicolaas Glock
Biogeosciences, 20, 3423–3447, https://doi.org/10.5194/bg-20-3423-2023, https://doi.org/10.5194/bg-20-3423-2023, 2023
Short summary
Short summary
Ocean deoxygenation due to climate warming is an evolving threat for organisms that are not well adapted to O2 depletion, such as many pelagic fish species. Other better-adapted organisms, such as some benthic foraminifera species, might benefit from ocean deoxygenation. Benthic foraminifera are a group of marine protists and can have specific adaptations to O2 depletion such as the ability to respire nitrate instead of O2. This paper reviews the current state of knowledge about these organisms.
Gerd Krahmann, Damian L. Arévalo-Martínez, Andrew W. Dale, Marcus Dengler, Anja Engel, Nicolaas Glock, Patricia Grasse, Johannes Hahn, Helena Hauss, Mark Hopwood, Rainer Kiko, Alexandra Loginova, Carolin R. Löscher, Marie Maßmig, Alexandra-Sophie Roy, Renato Salvatteci, Stefan Sommer, Toste Tanhua, and Hela Mehrtens
Earth Syst. Sci. Data Discuss., https://doi.org/10.5194/essd-2020-308, https://doi.org/10.5194/essd-2020-308, 2021
Preprint withdrawn
Short summary
Short summary
The project "Climate-Biogeochemistry Interactions in the Tropical Ocean" (SFB 754) was a multidisciplinary research project active from 2008 to 2019 aimed at a better understanding of the coupling between the tropical climate and ocean circulation and the ocean's oxygen and nutrient balance. On 34 research cruises, mainly in the Southeast Tropical Pacific and the Northeast Tropical Atlantic, 1071 physical, chemical and biological data sets were collected.
Cited articles
Arrigo, K. R.: Marine microorganisms and global nutrient cycles, Nature, 437, 349–355, 2005.
Bernhard, J. M.: Postmortem vital staining in benthic foraminifera: duration and importance in population and distributional studies, J. Foramin. Res., 18, 143–146, 1988.
Bernhard, J. M.: Distinguishing live from dead foraminifera: methods review and proper applications, Micropaleontology, 46 (Suppl. 1), 38–46, 2000.
Bernhard, J. M. and Reimers, C. E.: Benthic foraminiferal population fluctuations related to anoxia – Santa Barbara Basin, Biogeochemistry, 15, 127–149, https://doi.org/10.1007/BF00003221, 1991.
Bernhard, J. M., Visscher, P. T., and Bowser, S. S.: Sub-millimiter life positions of bacteria, protists, and metazoans in laminated sediments of the Santa Barbara Basin, Limnol. Oceanogr., 48, 813–828, 2003.
Bernhard, J. M., Casciotti, K. L., McIlvin, M. R., Beaudoin, D. J., Visscher, P. T., and Edgcomb, V. P.: Potential importance of physiologically diverse benthic foraminifera in sedimentary nitrate storage and respiration, J. Geophys. Res., 117, G03002, https://doi.org/10.1029/2012JG001949, 2012a.
Bernhard, J. M., Edgcomb, V. P., Casciotti, K. L., McIlvin, M. R., and Beaudoin, D. J.: Denitrification likely catalyzed by endobionts in an allogromiid foraminifer, ISME J., 6, 951–960, 2012b.
Bohlen, L., Dale, A. W., Sommer, S., Mosch, T., Hensen, C., Noffke, A., Scholz, F., and Wallmann, K.: Benthic nitogen cycling traversing the Peruvian oxygen minimum zone, Geochim. Cosmochim. Ac., 75, 6094–6111, 2011.
Cabrita, M. T. and Brotas, V.: Seasonal variation in denitrification and dissolved nitrogen fluxes in intertidal sediments of the Tagus estuary, Portugal, Mar. Ecol.-Prog. Ser., 202, 51–65, 2000.
Caralp, M., Lamy, A., and Pujos, M.: Contribution a la connaissance de la distribution bathymétrique des Foraminifères dans le golfe de Gascone, Revista Española de Micropaleontología, 2, 55–84, 1970.
Corliss, B. H.: Microhabitats of benthic foraminifera within deep-sea sediments, Nature, 314, 435–438, 1985.
Den Dulk, M., Reichart, G. J., Memon, G. M., Roelofs, E. M. P., Zachariasse, W. J., and van der Zwaan, G. J.: Benthic foraminiferal response to variations in surface water productivity and oxygenation in the northern Arabian Sea, Mar. Micropaleontol., 35, 43–66, 1998.
Finlay, B. J., Span. A. S. W., and Harman, J. M. P.: Nitrate respiration in primitive eukaryotes, Nature, 303, 333–335, 1983.
Glock, N., Eisenhauer, A., Milker, Y., Liebetrau, V., Schönfeld, J., Mallon, J., Sommer, S., and Hensen, C.: Environmental influences on the pore-density in tests of Bolivina spissa, J. Foramin. Res., 41, 22–32, 2011.
Glock, N., Schönfeld, J. and Mallon, J.: The functionality of pores in benthic foraminifera and bottom water oxygenation. A Review, in ANOXIA: Evidence for eukaryote survival and paleontological strategies, Cellular Origin, Life in Extreme Habitats and Astrobiology 21, edited by: Altenbach, A. V., Bernhard, J. M., and Seckbach, J., Springer Science + Business Media, 2011, 537–552, https://doi.org/10.1007/978-94-007-1896-8_28, 2012.
Glud, R. N., Thamdrup, B., Stahl, H., Wenzhoefer, F., Glud, A., Nomaki, H., Oguri, K., Revsbech, N. P., and Kitazato, H.: Nitrogen cycling in a deep ocean margin sediment (Sagami Bay, Japan), Limnol. Oceanogr., 54, 723–734, 2009.
Gooday, A. J., Bernhard, J. M., Levin, L. A., and Suhr, S. B.: Foraminifera in the Arabian Sea Oxygen minimum zone and other oxygen deficient settings: taxonomic composition, diversity, and relation to metazoan faunas, Deep-Sea Res. Pt. II, 47, 25–54, 2000.
Graco, M., Farias, L., Molina, V., Gutierrez, D., and Nielsen, L. P.: Massive developments of microbial mats following phytoplankton blooms in a naturally eutrophic bay: Implications for nitrogen cycling, Limnol. Oceanogr., 46, 821–832, 2001.
Grasshoff, K., Ehrhardt, M., and Kremling, K.: Methods of Seawater Analysis, Wiley-VCH, Weinheim, 1999.
Gruber N.: The dynamics of the marine nitrogen cycle and its influence on atmospheric CO2 variations, in: Carbon Cycle and Climate, edited by: Follows, M. and Oguz, T., The Ocean, NATO ASI Series, Kluwer Academic, Dordrecht, 97–148, 2004.
Gruber, N. and Sarmiento, J. L.: Global patterns of marine nitrogen fixation and denitrification, Global Biogeochem. Cy., 11, 235–266, 1997.
Guitiérrez, D., Enríquez, E., Purca, S., Quipúzcoa, L., Marquina, R., Flores, G., and Graco, M.: Oxygenation episodes on the continental shelf of central Peru: Remote forcing and benthic ecosystem response, Prog. Oceanogr., 79, 177–189, 2008.
Haake, F. W.: Benthic foraminifera of surface samples and sediment cores off Senegal and Gambia, West Africa, in: Benthische Foraminiferen in Oberflächen-Sedimenten und Kernen des Ostatlantiks vor Senegal/Gambia (Westafrika), edited by: Haake, F. W., Meteor Forschungsergebnisse, Deutsche Forschungsgemeinschaft, Reihe C Geologie und Geophysik, Gebrüder Bornträger, Berlin, Stuttgart, C32, 1–29, https://doi.org/10.1594/PANGAEA.548467, 1980.
Heinz, P., Ruepp, D., and Hemleben, C.: Benthic foraminifera assembleges at Great Meteor Seamount, Mar. Biol., 144, 985–998, https://doi.org/10.1007/s00227-003-1257-7, 2004.
Høgslund, S., Revsbech, N. P. Cedhagen, T. Nielsen, L. P., and Gallardo, V. A.: Denitrification, nitrate turn over, and aerobic respiration by benthic foraminiferans in the oxygen minimum zone off Chile, J. Exp. Mar. Biol. Ecol., 359, 85–91, 2008.
Jannink, N. T., Zachariasse, W. J., and van der Zwaan, G. J.: Living (Rose bengal stained) Benthic foraminifera from the Pakistan continental margin (northern Arabian Sea), Deep-Sea Res. Pt. I, 45, 1483–1513, 1998.
Kamp, A., de Beer, D., Nitsch, J. L., Lavik, G., and Stief, P.: Diatoms respire nitrate to survive dark and anoxic conditions, P. Natl. Acad. Sci. USA, 108, 5649–5654, https://doi.org/10.1073/pnas.1015744108, 2011.
Kitazato, H.: Foraminiferal microhabitats in four marine environments around Japan, Mar. Micropaleontol., 24, 29–41, 1994.
Kuypers, M. M. M., Sliekers, A. O., Lavik, G., Schmid, M., Jørgensen, B. B., Kuenen, J. G., Sinninghe Damsteì, J. S., Strous, M., and Jetten, M. S. M.: Anaerobic ammonium oxidation by anammox bacteria in the Black Sea, Nature, 422, 608–611, https://doi.org/10.1038/nature01472, 2003.
Lam, P., Lavik, G., Jensen, M. M., van de Vossenberg, J., Schmid, M., Woebken, D., Gutiérrez, D., Amann, R., Jetten, M. S. M., and Kuypers, M. M. M.: Revising the nitrogen cycle in the Peruvian oxygen minimum zone, P. Natl. Acad. Sci. USA, 106, 4752–4757, 2009.
Lutze, G. F.: Relative abundance of live (Rose Bengal stained) benthic foraminifera in the small size fraction of surface sediment off NW Africa (Table 3), in: Depth distribution of benthic foraminifera on the continental margin off NW Africa, edited by: Lutze, G. F., Meteor Forschungsergebnisse, Deutsche Forschungsgemeinschaft, Reihe C Geologie und Geophysik, Gebrüder Bornträger, Berlin, Stuttgart, C32, 31–80 https://doi.org/10.1594/PANGAEA.536260, 1980.
Lutze, G. F. and Altenbach, A. V.: Technik und Signifikanz der Lebendfärbung benthischer Foraminiferen in Bengalrot, Geologisches Jahrbuch, Reihe A, 128, 251–265, 1991.
Mallon, J.: Benthic foraminifera of the Peruvian and Ecuadorian Continental Margin, dissertation, Geomar Helmholtz Centre for Ocean Research, Kiel, 2012.
Mallon, J., Glock, N., and Schönfeld, J.: The response of benthic foraminifera to low-oxygen conditions of the Peruvian oxygen minimum zone, in: ANOXIA: Evidence for eukaryote survival and paleontological strategies, Cellular Origin, Life in Extreme Habitats and Astrobiology 21, edited by: Altenbach, A. V., Bernhard, J. M., and Seckbach, J., Springer Science + Business Media, 305–321, https://doi.org/10.1007/978-94-007-1896-8_16, 2012.
Mosch, T., Sommer, S., Dengler, M., Noffke, A., Bohlen, L., Pfannkuche, O., Liebetrau, V., and Wallmann, K.: Factors influencing the distribution of epibenthic megafauna across the Peruvian oxygen minimum zone, Deep-Sea Res. Pt. I, 68, 123–135, https://doi.org/10.1016/j.dsr.2012.04.014, 2012.
Murray, J. W.: The niche of benthic foraminifera, critical thresholds and proxies, Mar. Micropaleontol., 41, 1–7, 2001.
Murray, J. W. and Bowser, S. S.: Mortality, protoplasm decay rate, and reliability of staining techniques to recognize `living' foraminifera: a review, J. Foramin. Res., 30, 66–77, 2000.
Noffke, A., Hensen, C., Sommer, S., Scholz, F., Bohlen, L., Mosch, T., Graco, M., and Wallmann, K.: Benthic iron and phosphorus fluxes across the Peruvian oxygen minimum zone, Limnol. Oceanogr., 57, 851–867, 2012.
Ohga, T. and Kitazato, H.: Seasonal changes in bathyal foraminiferal populations in response to the flux of organic matter (Sagami Bay, Japan), Terra Nova, 9, 33–37, 1997.
Otto, G. H.: Comparative tests of several methods of sampling heavy mineral concentrates, J. Sediment. Petrol., 3, 30–39, 1933.
Phleger, F. B. and Soutar, A.: Production of benthic foraminifera in three east Pacific oxygen minima, Micropaleontology, 19, 110–115, 1973.
Pierotti, D. and Rasmussen, R. A.: Nitrous oxide measurements in the eastern tropical Pacific Ocean, Tellus, 32, 56–72, 1980.
Piña-Ochoa, E., Høgslund, S., Geslin, E., Cedhagen, T., Revsbech, N. P., Nielsen, L. P., Schweizer, M., Jorissen, F., Rysgaard, S., and Risgaard-Petersen, N.: Widespread occurence of nitrate storage and denitrification among Foraminifera and Gromiida, P. Natl. Acad. Sci. USA, 107, 1148–1153, 2010a.
Piña-Ochoa, E., Koho, K. A., Geslin, E., and Risgaard-Petersen, N.: Survival and life strategy of the foraminiferan Globobulimina turgida through nitrate storage and denitrification, Mar. Ecol.-Prog. Ser., 417, 39–49, 2010b.
Prokopenko, M. G., Sigman, D. M., Berelson, W. M., Hammond, D. E., Barnett, B., Chong, L., and Townsend-Small, A.: Denitrification in anoxic sediments supported by biological nitrate transport, Geochim. Cosmochim. Ac., 75, 7180–7199, 2011.
Reimers, C. E., Ruttenberg, K. C., Canfield, D. E., Christiansen, M. B., and Martin, J. B.: Porewater pH and authigenic phases formed in the uppermost sediments of the Santa Barbara Basin, Geochim. Cosmochim. Ac., 60, 4037–4057, https://doi.org/10.1016/S0016-7037(96)00231-1, 1996.
Revsbech, N. P., Larsen, L. H., Gundersen, J., Dalsgaard, T., Ulloa, O., and Thamdrup, B.: Determination of ultra-low oxygen concentrations in oxygen minimum zones by the STOX sensor, Limnol. Oceanogr.-Meth., 7, 371–381, 2009.
Risgaard-Petersen, N., Langezaal, A. M., Ingvardsen, S., Schmid, M. C., Jetten, M. S., Op den Camp, H. J. M., Derksen, J. W. M., Piña-Ochoa, E., Eriksson, S. P., Nielsen, L. P., Revsbech, N. P., Cedhagen, T., and van der Zwaan, G. J.: Evidence for complete denitrification in a benthic foraminifer, Nature, 443, 93–96, 2006.
Ryabenko, E., Kock, A., Bange, H. W., Altabet, M. A., and Wallace, D. W. R.: Contrasting biogeochemistry of nitrogen in the Atlantic and Pacific Oxygen Minimum Zones, Biogeosciences, 9, 203–215, https://doi.org/10.5194/bg-9-203-2012, 2012.
Rysgaard, S., Fossing, H., and Jensen, M. M.: Organic matter degradation through oxygen respiration, denitrification, and manganese, iron, and sulfate reduction in marine sediments (the Kattegat and the Skagerrak), Ophelia, 55, 77–91, 2001.
Schmiedl, G., Mackensen, A., and Müller, P. J.: Recent benthic foraminifera from the eastern South Atlantic Ocean: Dependance on food supply and water masses, Mar. Micropaleontol., 32, 249–287, 1997.
Schönfeld, J.: History and development of methods in Recent benthic foraminiferal studies, J. Micropaleontol., 31, 53–72, 2012.
Schumacher, S., Jorissen, F. J., Dissard, D., Larkin, K. E., and Gooday, A. J.: Live (Rose Bengal stained) and dead benthic foraminifera from the oxygen minimum zone of the Pakistan continental margin (Arabian Sea), Mar. Micropaleontol., 62, 45–73, 2007.
Shoun, H. and Tanimoto, T.: Denitrification by the fungus Fusarium oxysporum and involvement of cytochrome P-450 in the respiratory nitrite reduction, J. Biol. Chem., 266, 11078–11082, 1991.
Sommer, S., Linke, P., Pfannkuche, O., Schleicher, T., Schneider v. Deimling, J., Reitz, A., Haeckel, M., Flögel, S., and Hensen, C.: Seabed methane emissions and the habitat of frenulate tubeworms on the Captain Arutyunov mud volcano (Gulf of Cadiz), Mar. Ecol.-Prog. Ser., 382, 69–86, https://doi.org/10.3354/meps07956, 2009.
Sommer S., McGinnis, D. F., Linke, P., Camilli, R., Mosch, T., and Pfannkuche, O.: Life at the edge – oscillating lower boundary of the Peruvian oxygen minimum zone, Eos Transactions AGU, 91, Ocean Science Meeting Supplementary, Abstract BO24C–08, 2010.
Tengberg, A., Hovdenes, J., Andersson, H. J. Brocandel, O., Diaz, R., Hebert, D., Arnerich, T., Huber, C., Körtzinger, A., Khripounoff, A., Rey, F., Rönning, C., Schimanski, J., Sommer, S., and Stangelmayer, A.: Evaluation of a lifetime-based optode to measure oxygen in aquatic systems, Limnol. Oceanogr.-Meth., 4, 7–17, 2006.
Thamdrup B. and Dalsgaard T.: Production of N2 through anaerobic ammonium oxidation coupled to nitrate reduction in marine sediments, Appl. Environ. Microb., 68, 1312–1318, 2002.
Timm, S.: Rezente Tiefsee-Benthosforaminiferen aus Oberflächensedimenten des Golfes von Guinea (Westafrika): Taxonomie, Verbreitung, Oekologie und Korngrössenfraktionen = Recent deep-sea benthic foraminifera from surface sediments of Gulf of Guinea (Westafrika): taxonomy, distributions, ecology and grain size fractions, dissertation, Geologisch-Paläontologisches Institut und Museum, Christian-Albrechts-Universität, 1992.
Uchio, T.: Ecology of living benthonic foraminifera from the San Diego, California, Area, Cushman Foundation for Foraminiferal Research, Special Publication, 5, 1–72, 1960.
Usuda, K., Toritsuka, N., Matsuo, Y., Kim, D. H., and Shoun, H.: Denitrification by the fungus Cylindrocarpon tonkinense: Anaerobic cell growth and two isozyme forms of cytochrome P-450nor, Appl. Environ. Microb., 61, 883–889, 1995.
Van de Graaf, A. A., Mulder, A., de Bruijn, P., Jetten, M. S. M., and Kuenen, J. G.: Anaerobic oxidation of ammonium is a biologically mediated process, Appl. Environ. Microb., 1, 1246–1251, 1995.
Walker, D. A., Linton, A. E., and Schafer, C. T.: Sudan Black B: A superior stain to Rose Bengal for distinguishing living from nonliving foraminifera, J. Foramin. Res., 4, 205–215, 1974.
Zumft, W. G.: Cell biology and molecular basis of denitrification, Microbiol. Mol. Biol. R., 61, 533–616, 1997.
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