Coupled physical/biogeochemical modeling including O2-dependent processes in the Eastern Boundary Upwelling Systems: Application in the Benguela

Biogeosciences

Volumn 10, Issue 6, 2013, Pages 3559-3591

  Gutknecht, E ^a   Dadou, I ^a   Le Vu, B ^a   Cambon, G ^a   Sudre, J ^a   Garcon V ^a   MacHu, E ^b   Rixen, T ^c   Kock, A ^d   Flohr, A ^c   Paulmier, A ^a,e   Lavik, G ^f  

^a CNES   (France)

^b Centre Bretagne   (France)

^c LEIBNIZ CENTRE FOR TROPICAL MARINE RESEARCH ZMT   (Germany)

^d GEOMAR HELMHOLTZ CENTRE FOR OCEAN RESEARCH KIEL   (Germany)

^e INSTITUTO DEL MAR DEL PERÚ   (Peru)

^f MAX PLANCK INSTITUTE FOR MARINE MICROBIOLOGY   (Germany)

Author keywords

[No Author keywords available]

Indexed keywords

BIOAVAILABILITY; CHLOROPHYLL A; CLIMATE CHANGE; EASTERN BOUNDARY CURRENT; GREENHOUSE GAS; NITRATE; NITROGEN CYCLE; OZONE; PARAMETERIZATION; SATELLITE DATA; SEASONAL VARIATION; SENSITIVITY ANALYSIS; VERTICAL DISTRIBUTION;

ANGOLA; BENGUELA; NAMIBIA;

EID: 84879120241     PISSN: 17264170     EISSN: 17264189     Source Type: Journal    
DOI: 10.5194/bg-10-3559-2013     Document Type: Article

References (121)

1. Aiken, J. and Bale, A. J.: An introduction to the Atlantic Meridional Transect (AMT) Programme, Prog. Oceanogr., 45, 251-256, doi:10.1016/S0079- 6611(00)00004-5, 2000. (Pubitemid 30329549)
2. Aiken, J., Woodward, M. S., Robinson, C., Rees, A., Serret, P., Bowie, A.,Woodd-Walker, R., Pilgrim, D., Holligan, P., Suggett, D., Hooker, S., Dempsey, C., Brown, J., Maritorena, S., Barlow, R., Lucas, M., and Tilstone, G.: Atlantic Meridional Transect, AMT 6 cruise report, 14 May 5 to 16 June 1998, Plymouth Marine Laboratory, Plymouth, UK, 147 pp., 1998.
3. Aiken, J., Rees, N., Hooker, S., Holligan, P., Bale, A., Robins, D., Moore, G., Harris, R., and Pilgrim, D.: The Atlantic Meridional Transect: overview and synthesis of data, Prog. Oceanogr., 45, 257-312, doi:10.1016/S0079-6611(00)00005-7, 2000. (Pubitemid 30329550)
4. Anderson, J. J., Okubo, A., Robbins, A. S., and Richards, F. A.: A model for nitrite and nitrate distributions in oceanic oxygen minimum zones, Deep-Sea Res., 29, 1113-1140, 1982. (Pubitemid 13489006)
5. Anonymous: Greenhouse gases hit modern day-highs, Nature, 456, 558-559, doi:10.1038/456558b, 2008.
6. Arrigo, R. A.: Marine microorganisms and global nutrient cycles, Nature, 437, 349-355, 2005.
7. Bakun, A., Field, D. B., Redondo-Rodriguez, A., and Scarla, J.: Greenhouse gas, upwelling-favorable winds, and the future of coastal ocean ,upwelling ecosystems, Glob. Change Biol., 16, 1213-1228, doi:10.1111/j.1365- 2486.2009.02094.x, 2010.
8. Banyte, D., Tanhua, T., Visbeck, M.,Wallace, D.W. R., Karstensen, J., Krahmann, G., Schneider, A., Stramma, L., and Dengler, M., Diapycnal diffusivity at the upper boundary of the tropical North Atlantic oxygen minimum zone, J. Geophys. Res., 117, C09016, doi:10.1029/2011JC007762, 2012.
9. Barlow, R. G., Aiken, J., Holligan, P. M., Cummings, D. G., Maritorena, S., and Hooker, S.: Phytoplankton pigment and absorption characteristics along meridional transects in the Atlantic Ocean, Deep Sea Res. Pt I, 49, 637-660, doi:10.1016/S0967-0637(01)00081-4, 2002. (Pubitemid 34227948)
10. Barlow, R. G., Aiken, J., Moore, G. F., Holligan, P. M., and Lavender, S.: Pigment adaptations in surface phytoplankton along the eastern boundary of the Atlantic Ocean, Mar. Ecol. Prog.-Ser., 281, 13-26, doi:10.3354/meps281013, 2004. (Pubitemid 39569269)
11. Barlow, R., Lamont, T., Mitchell-Innes, B., Lucas, M., and Thomalla, S.: Primary production in the Benguela ecosystem, 1999-2002, Afr. J. Mar. Sci., 31, 97-101, doi:10.2989/AJMS.2009.31.1.9.780, 2009.
12. Barnier, B., Siefridt, L., and Marchesiello, P.: Thermal forcing for a global ocean circulation model using a three-year climatology of ECMWF analyses, J. Marine Syst., 6, 363-380, doi:10.1016/0924-7963(94)00034-9, 1995. (Pubitemid 26438045)
13. Bianucci, L., Fennel, K., and Denman, K. L.: Role of sediment denitrification in water column oxygen dynamics: comparison of the North American East andWest Coasts, Biogeosciences, 9, 2673-2682, doi:10.5194/bg-9- 2673-2012, 2012.
14. Brown, P. C., Painting, S. J., and Cochrane, K. L.: Estimates of phytoplankton and bacterial biomass and production in the northern and southern Benguela ecosystems, S. Afr. J. Sci., 11, 537-564, 1991.
15. Caperon, J. and Meyer, J.: Nitrogen-limited growth of marine phytoplankton-I. Changes in population characteristics with steady-state growth rate, Deep Sea Res., 19, 601-618, doi:10.1016/0011-7471(72)90089-7, 1972a.
16. Caperon, J. and Meyer, J.: Nitrogen-limited growth of marine phytoplankton-II. Uptake kinetics and their role in nutrient limited growth of phytoplankton, Deep Sea Res., 19, 619-632, doi:10.1016/0011-7471(72)90090-3, 1972b.
17. Carlson, C. A. and Ducklow, H. W.: Dissolved organic carbon in the upper ocean of the central equatorial Pacific Ocean, 1992: Daily and finescale vertical variations, Deep Sea Res. Pt. II, 42, 639-656, doi:10.1016/0967- 0645(95)00023-J, 1995. (Pubitemid 26462815)
18. Carr, M. E.: Estimation of potential productivity in Eastern Boundary Currents using remote sensing, Deep Sea Res. Pt II, 49, 59-80, 2002.
19. Carr, M. and Kearns, E. J.: Production regimes in four Eastern Boundary Current systems, Deep Sea Res. Pt II, 50, 3199-3221, doi:10.1016/j.dsr2.2003.07. 015, 2003. (Pubitemid 37537794)
20. CARS: CSIRO Atlas of Regional Seas, http://www.marine.csiro.au/-dunn/ cars2006, 30 March 2011, 2006.
21. CARS: CSIRO Atlas of Regional Seas, http://www.marine.csiro.au/-dunn/ cars2009, 30 March 2011, 2009.
22. Casey, K. S. and Cornillon, P.: A comparison of satellite and in situ-based sea surface temperature climatologies, J. Climate, 12, 1848-1863, 1999. (Pubitemid 29534630)
23. Charria, G., Dadou, I., Cipollini, P., Dŕevillon, M., and Garçon, V.: Influence of Rossby waves on primary production from a coupled physical-biogeochemical model in the North Atlantic Ocean, Ocean Sci., 4, 199-213, doi:10.5194/os-4-199-2008, 2008.
24. Conkright, M. E. and O'Brien, T. D.: World Ocean Atlas 2001, Volume 6: Chlorophyll, Global Biogeochem. Cy., 8, 65-80, doi:199410.1029/93GB03318, 1994. (Pubitemid 24402983)
25. Conkright, M. E., O'Brien, T. D., Stephens, C., Locarnini, R. A., Garcia, H. E., Boyer, T. P., and Antonov, J. I.:World Ocean Atlas 2001, Volume 6: Chlorophyll, edited by:Levitus. S., NOAA Atlas NESDIS 52, US Government Printing Office, Washington, DC, 2002.
26. Cornejo, M., Faras, L., and Paulmier, A.: Temporal variability in N2O water content and its air-sea exchange in an upwelling area off central Chile (36 S), Mar. Chem., 101, 85-94, doi:10.1016/j.marchem.2006.01.004, 2006.
27. Da Silva, A. M., Young, C. C., and Levitus, S.: Atlas of Surface Marine Data 1994. Vol. 1: Algorithms and Procedures, NOAA Atlas NESDIS 6, Department of Commerce, Washington DC, USA 1994.
28. Dadou, I., Lamy, F., Rabouille, C., Ruiz-Pino, D., Andersen, V., Bianchi, M., and Garcon, V.: An integrated biological pump model from the euphotic zone to the sediment: a 1-D application in the Northeast tropical Atlantic, Deep Sea Res. Pt II, 48, 2345-2381, 2001. (Pubitemid 32507509)
29. Dadou, I., Evans, G., and Garcon, V.: Using JGOFS in situ and ocean color data to compare biogeochemical models and estimate their parameters in the subtropical North Atlantic Ocean, J. Mar. Res., 62, 565-594, 2004. (Pubitemid 39464404)
30. Debreu, L., Marchesiello, P., Penven, P., and Cambon, G.: Twoway nesting in split-explicit ocean models: algorithms, implementation and validation, Ocean Model., 49-50, 1-21, doi:10.1016/j.ocemod.2012.03.003, 2012.
31. Demarcq, H.: Trends in primary production, sea surface temperature and wind in upwelling systems (1998-2007), Prog. Oceanogr., doi:10.1016/j.pocean. 2009.07.022, 2009.
32. Deutsch, B., Forster, S., Wilhelm, M., Dippner, J. W., and Voss, M.: Denitrification in sediments as a major nitrogen sink in the Baltic Sea: an extrapolation using sediment characteristics, Biogeosciences, 7, 3259-3271, doi:10.5194/bg-7-3259-2010, 2010.
33. Dittmar, T. and Birkicht, M.: Regeneration of nutrients in the northern Benguela upwelling and the Angola-Benguela Front areas, S. Afr. J. Sci., 97, 239-246, 2001. (Pubitemid 32739428)
34. Duteil, O. and A. Oschlies, A.: Sensitivity of simulated extent and future evolution of marine suboxia to mixing intensity, Geophys. Res. Lett., 38, L06607, doi:10.1029/2011GL046877, 2011.
35. Eppley, R. W.: Temperature and Phytoplankton Growth in the Sea, Fish. B.-NOAA, 70, 1063-1085, 1972.
36. Eppley, R. W., Rogers, J. N. and McCarthy, J. J.: Half-Saturation Constants for Uptake of Nitrate and Ammonium by Marine Phytoplankton, Limnol. Oceanogr., 14, 912-920, 1969.
37. Evans, G. T. and Parslow, J. S.: A model of annual plankton cycles, Bio. Oceanogr., 3, 327-347, 1985.
38. Fasham, M. J. R., Ducklow, H.W. and McKelvie, S. M.: A nitrogenbased model of plankton dynamics in the oceanic mixed layer, J. Mar. Res., 48, 591-639, 1990.
39. Fasham, M. J. R., Boyd, P. W., and Savidge, G.: Modeling the Relative Contributions of Autotrophs and Heterotrophs to Carbon Flow at a Lagrangian JGOFS Station in the Northeast Atlantic: The Importance of DOC, Limnol. Oceanogr., 44, 80-94, 1999. (Pubitemid 29077770)
40. Fischer, T., Banyte, D., Brandt, P., Dengler, M., Krahmann, G., Tanhua, T., and Visbeck, M.: Diapycnal oxygen supply to the tropical North Atlantic oxygen minimum zone, Biogeosciences Discuss., 9, 14291-14325, doi:10.5194/bgd-9-14291-2012, 2012.
41. Freon, P., Barange, M., and Arstegui, J.: Eastern Boundary Upwelling Ecosystems: Integrative and comparative approaches, 2009.
42. Fussel, J., Lam, P., Lavik, G., Jensen, M. M., Holtappels, M., Gunter, M., and Kuypers, M. M. M.: Nitrite oxidation in the Namibian oxygen minimum zone, ISME J., 6, 1200-1209 doi:10.1038/ismej.2011.178, 2011.
43. Garcia, H. E. and Gordon, L. I.: Oxygen Solubility in Seawater: Better Fitting Equations, Limnol. Oceanogr., 37, 1307-1312, 1992.
44. Garreaud, R. D. and Falvey, M.: The coastal winds off western subtropical South America in future climate scenarios, Int. J. Climatol., doi::10.1002/joc.1716, 2008.
45. Gruber, N.: The Dynamics of the Marine Nitrogen Cycle and its Influence on Atmospheric CO2 Variations, in: Carbon Climate interactions, edited by: Oguz, T. and Follows, M., Kluwer, Dordrecht, 97-148, 2004.
46. Gutknecht, E., Dadou, I., Marchesiello, P., Cambon, G., Le Vu, B., Sudre, J., Garcon, V., Machu, E., Rixen, T., Kock, A., Flohr, A., Paulmier, A., and Lavik, G.: Nitrogen transfers off Walvis bay: a 3-D coupled physical/ biogeochemical modeling approach in the Namibian upwelling system, Biogeosciences, in press, 2013.
47. Hofmann, A. F., Peltzer, E. T., Walz, P. M., and Brewer, P. G., Hypoxia by degrees: Establishing definitions for a changing ocean, Deep-Sea Res. Pt. I, 58, 1212-1226, doi:10.1016/j.dsr.2011.09.004, 2011.
48. Holligan, P.: Atlantic Meridional Transect, AMT 17 Cruise Report: RRS Discovery, 15 October to 28 November 2005, 135 pp., Plymouth Marine Laboratory, Plymouth, UK, 2006.
49. Huggett, J., Verheye, H., Escribano, R., and Fairweather, T.: Copepod biomass, size composition and production in the Southern Benguela: Spatio-temporal patterns of variation, and comparison with other eastern boundary upwelling systems, Prog. Oceanogr., 83, 197-207, doi:10.1016/j.pocean. 2009.07.048, 2009.
50. Huret, M., Dadou, I., Dumas, F., Lazure, P., and Garcon, V.: Coupling physical and biogeochemical processes in the Rio de la Plata plume, Cont. Shelf Res., 25, 629-653, doi:10.1016/j.csr.2004.10.003, 2005. (Pubitemid 40240994)
51. Hurtt, G. C. and Armstrong, R. A.: A pelagic ecosystem model calibrated with BATS data, Deep Sea Res. Pt II, 43, 653-683, 1996.
52. Hutchings, L., van der Lingen, C. D., Shannon, L. J., Crawford, R. J. M., Verheye, H. M. S., Bartholomae, C. H., van der Plas, A. K., Louw, D., Kreiner, A., Ostrowski, M., Fidel, Q., Barlow, R. G., Lamont, T., Coetzee, J., Shillington, F., Veitch, J., Currie, J. C., and Monteiro, P. M. S.: The Benguela Current: An ecosystem of four components, Prog. Oceanogr., 83, 15-32, doi:10.1016/j.pocean.2009.07.046, 2009.
53. Jackett, D. R. and McDougall, T. J.: Minimal Adjustment of Hydrographic Profiles to Achieve Static Stability, J. Atmos. Ocean. Tech., 12, 381-389, 1995.
54. Jin, X. and Gruber, N.: Offsetting the radiative benefit of ocean iron fertilization by enhancing N2O emissions, Geophys. Res. Lett., 30, 2249, doi:10.1029/2003GL018458, 2003.
55. Kalvelage, T., Jensen, M. M., Contreras, S., Revsbech, N. P., Lam, P., Gunter, M., LaRoche, J., Lavik, G., and Kuypers, M. M. M.: Oxygen sensitivity of anammox and coupled N-Cycle processes in Oxygen Minimum Zones, PLoS ONE, 6, e29299, doi:10.1371/journal.pone.0029299, 2011.
56. Kartal, B., Kuypers, M. M. M., Lavik, G., Schalk, J., Op den Camp, H. J. M., Jetten, M. S. M., and Strous, M., Anammox bacteria disguised as denitrifiers: nitrate reduction to dinitrogen gas via nitrite and ammonium, Environ. Microb., 9, 635-642, doi:10.1111/j.1462-2920.2006.01183.x, 2007.
57. Keeling, R. F., Stephens, B. B., Najjar, R. G., Doney, S. C., Archer, D., and Heimann, M.: Seasonal variations in the atmospheric O2/N2 ratio in relation to the kinetics of airsea gas exchange, Global Biogeochem. Cy., 12, 141-163, doi:199810.1029/97GB02339, 1998.
58. Kirchman, D. L., Lancelot, C., Fasham, M., Legendre, L., Radach, G., and Scott, M.: Dissolved organic matter in biogeochemical models in the ocean, in: Towards a Model of Ocean Biogeochemical Processes, 209-225, Springer-Verlag, 1993.
59. Kone, V., Machu, E., Penven, P., Andersen, V., Garcon, V., Freon, P., and Demarcq, H.: Modeling the primary and secondary productions of the southern Benguela upwelling system: A comparative study through two biogeochemical models, Global Biogeochem. Cy., 19, GB4021, doi:10.1029/2004GB002427, 2005. (Pubitemid 43280684)
60. Kreiner, A. and Ayon, P.: Zooplankton dynamics from 1994 to 2006 in the upwelling systems off Peru and northern Namibia, Eastern boundary upwelling ecosystems, Las Palmas, Gran Canaria, Spain, 2-6 June 2008, P08OP05, 2008.
61. Kuypers, M. M. M., Lavik, G., Woebken, D., Schmid, M., Fuchs, B. M., Amann, R., Jorgensen, B. B., and Jetten, M. S. M.: Massive nitrogen loss from the Benguela upwelling system through anaerobic ammonium oxidation, P. Natl. Acad. Sci. USA, 102, 6478-6483, doi:10.1073/pnas.0502088102, 2005. (Pubitemid 40617468)
62. Lacroix, G. and Nival, P.: Influence of meteorological variability on primary production dynamics in the Ligurian Sea (NW Mediterranean Sea) with a 1D hydrodynamic/biological model, J. Marine Syst., 16, 23-50, doi:10.1016/S0924- 7963(97)00098-5, 1998. (Pubitemid 28407983)
63. Lam, P., Lavik, G., Jensen, M. M., van de Vossenberg, J., Schmid, M., Woebken, D., Gutierrez, D., Amann, R., Jetten, R. S. M., and Kuypers, M. M. M.: Revising the nitrogen cycle in the Peruvian oxygen minimum zone, 106, 4752-4757, P. Natl. Acad. Sci. USA, doi:10.1073/pnas.0812444106, 2009.
64. Large, W. G., McWilliams, J. C., and Doney, S. C.: Oceanic vertical mixing: A review and a model with a nonlocal boundary layer parameterization, Rev. Geophys., 32, 363-403, doi:199410.1029/94RG01872, 1994. (Pubitemid 24987221)
65. Lavik, G., Stuhrmann, T., Bruchert, V., Van der Plas, A., Mohrholz, V., Lam, P., Muszmann, M., Fuchs, B. M., Amann, R., Lass, U., and Kuypers, M. M. M.: Detoxification of sulphidic African shelf waters by blooming chemolithotrophs, Nature, 457, 581-584, doi:10.1038/nature07588, 2008.
66. Le Vu, B., Gutknecht, E. Machu, E., Sudre, J., Dadou, I., Veitch, J., and Garcon, V.: Physical and Biogeochemical processes maintaining the Oxygen Minimum Zone in the Benguela Upwelling System using an eddy resolving model, Biogeosciences, in preparation, 2013.
67. Liu, W. T., Tang, W., and Polito, P. S.: NASA scatterometer provides global ocean surface wind fields with more structures than numerical weather prediction, Geophys. Res. Lett., 25, 761-764, 1998. (Pubitemid 28424535)
68. Lueker, T. J., Walker, S. J., Vollmer, M. K., Keeling, R. F., Nevison, C. D., Weiss, R. F., and Garcia, H. E.: Coastal upwelling air-sea fluxes revealed in atmospheric observations of O2/N2, CO2 and N2O, Geophys. Res. Lett., 30, 1292, doi:10.1029/2002GL016615, 2003.
69. Marchesiello, P., McWilliams, J. C., and Shchepetkin, A.: Open boundary conditions for long-term integration of regional 5003(00)00013-5, 2001.
70. Marchesiello, P., McWilliams, J. C., and Shchepetkin, A.: Equilibrium structure and dynamics of the California Current System, J. Phys. Oceanogr., 33, 753-783, 2003. (Pubitemid 37196016)
71. McClain, C. R., Cleave, M. L., Feldman, G. C., Gregg, W. W., Hooker, S. B., and Kuring, N.: Science Quality SeaWiFS Data for Global Biosphere Research, NASA/Goddard Space Flight Center, Sea Technol., 39, 10-16, 1998.
72. Mohrholz, V., Bartholomae, C. H., van der Plas, A. K., and Lass, H. U.: The seasonal variability of the northern Benguela undercurrent and its relation to the oxygen budget on the shelf, Cont. Shelf Res., 28, 424-441, doi:10.1016/j.csr.2007.10.001, 2008.
73. Mohrholz, V., Heene, T., Tsanwani, M., Morris, T., and Muller, A.: 1-Working group "Hydrography", in Geochemistry and Ecology of the Namibian Upwelling System (GENUS Project) and St Helena Bay Monitoring Line (SHBML), cruise report, FRS Africana, 1 to 17 December 2009, 2009.
74. Monteiro, P. M. S. and van der Plas, A. K.: Low oxygen water (LOW) variability in the Benguela system: Key processes and forcing scales relevant to forecasting, in Benguela-Predicting a Large Marine Ecosystem, 14, 1-90, Elsevier, 2006.
75. Monteiro, P. M. S., van der Plas, A., Mohrholz, V., Mabille, E., Pascall, A., and Joubert, W.: Variability of natural hypoxia and methane in a coastal upwelling system: Oceanic physics or shelf biology, Geophys. Res. Lett., 33, L16614, doi:10.1029/2006GL026234, 2006. (Pubitemid 44945739)
76. Monteiro, P. M. S., van der Plas, A. K., Ḿelice, J., and Florenchie, P.: Interannual hypoxia variability in a coastal upwelling system: Ocean-shelf exchange, climate and ecosystem-state implications, Deep Sea Res. Pt I, 55, 435-450, doi:10.1016/j.dsr.2007.12.010, 2008. (Pubitemid 351479215)
77. Monteiro, P. M. S., Dewitte, B., Scranton, M. I., Paulmier, A., and van der Pla, A.: The role of open ocean boundary forcing on seasonal to decadal-scale variability and long-term change of natural shelf hypoxia, Environ. Res. Lett., 6, 025002, doi:10.1088/1748-9326/6/2/025002, 2011.
78. Morel, A. and Berthon, J. F.: Surface pigments, algal biomass pro-files, and potential production of the euphotic layer: Relationships reinvestigated in view of remote-sensing applications. Limnol. Oceanogr., 34, 1545-1562, 1989.
79. Nalewajko, C. and Garside, C.: Methodological Problems in the Simultaneous Assessment of Photosynthesis and Nutrient Uptake in Phytoplankton as Functions of Light Intensity and Cell Size, Limnol. Oceanogr., 28, 591-597, 1983.
80. Nevison, C., Butler, J. H., and Elkins, J. W.: Global distribution of N2O and the Delta N2O-AOU yield in the subsurface ocean, Global Biogeochem. Cy., 17, 1119, doi:10.1029/2003GB002068, 2003.
81. Olivieri, R. A. and Chavez, F. P.: A model of plankton dynamics for the coastal upwelling system of Monterey Bay, California, Deep-Sea Res. Pt II, 47, 1077-1106, 2000. (Pubitemid 30124491)
82. O'Reilly, J. E., Maritorena, S., Siegel, D., O 'Brien, M., Toole, D., Greg Mitchell, B., Kahru, M., Chavez, F., Strutton, P., Cota, G., Hooker, S., McClain, C., Carder, K., Muller-Karger, F., Harding, L., Magnuson, A., Phinney, D., Moore, G., Aiken, J., Arrigo, K., Letelier, R., and Culver, M.: Ocean color chlorophyll-a algorithms for SeaWiFS, OC2, and OC4: Version 4, edited by: O'Reilly, J. E., and 24 coauthors, in: SeaWiFS Postlaunch Calibration and Validation Analyses, Part 3. NASA Tech. Memo. 2000-206892, Vol. 11, edited by: Hooker, S. B. and Firestone, E. R., NASA Goddard Space Flight Center, Greenbelt, Maryland, 9-23, 2000.
83. Oschlies, A. and Garcon, V.: An eddy-permitting coupled physicalbiological model of the North Atlantic-1. Sensitivity to advection numerics and mixed layer physics, Global Biogeochem. Cy., 13, 135-160, doi:10.1029/98GB02811, 1999. (Pubitemid 29365970)
84. Paulmier, A., Ruiz-Pino, D., and Garcon, V.: The oxygen minimum zone (OMZ) off Chile as intense source of CO2 and N2O, Cont. Shelf Res., 28, 2746-2756, doi:10.1016/j.csr.2008.09.012, 2008.
85. Pena, M. A., Katsev, S., Oguz, T., and Gilbert, D.: Modeling dissolved oxygen dynamics and hypoxia, Biogeosciences, 7, 933-957, doi:10.5194/bg-7-933- 2010, 2010.
86. Penven, P., Roy, C., Brundrit, G. B., de Verdiere, A. C., Freon, P., Johnson, A. S., Lutjeharms, J. R. E., and Shillington, F. A.: A regional hydrodynamic model of upwelling in the Southern Benguela, S. Afr. J. Sci., 97, 472-475, 2001. (Pubitemid 34147301)
87. Penven, P., Echevin, V., Pasapera, J., Colas, F., and Tam, J.: Average circulation, seasonal cycle, and mesoscale dynamics of the Peru Current System: A modeling approach, J. Geophys. Res., 110, C10021, doi:10.1029/2005JC002945, 2005. (Pubitemid 41763349)
88. Penven, P., Debreu, L., Marchesiello, P., and McWilliams, J. C.: Evaluation and application of the ROMS 1-way embedding procedure to the central california upwelling system, Ocean Model., 12, 157-187, doi:10.1016/j.ocemod. 2005.05.002, 2006a. (Pubitemid 41799925)
89. Penven, P., Lutjeharms, J. R. E., and Florenchie, P.: Madagascar: A pacemaker for the Agulhas Current system, Geophys. Res. Lett., 33, L17609, doi:10.1029/2006GL026854, 2006b. (Pubitemid 44945651)
90. Penven, P., Marchesiello, P., Debreu, L., and Lefevre, J.: Software tools for pre-and post-processing of oceanic regional simulations, Environ. Modell. Softw., 23, 660-662, doi:10.1016/j.envsoft.2007.07.004, 2008.
91. Peterson, W. T., Painting, S. J., and Hutchings, L.: Diel variations in gut pigment content, diel vertical migration and estimates of grazing impact for copepods in the southern Benguela upwelling region in October 1987, J. Plankton Res., 12, 259-281, doi:10.1093/plankt/12.2.259, 1990.
92. Popova, E. E., Lozano, C. J., Srokosz, M. A., Fasham, M. J. R., Haley, P. J., and Robinson, A. R.: Coupled 3D physical and biological modelling of the mesoscale variability observed in North-East Atlantic in spring 1997: biological processes, Deep Sea Res. Pt I, 49, 1741-1768, doi:10.1016/S0967-0637(02)00091- 2, 2002. (Pubitemid 35399937)
93. Postel, L., Arndt, E. A., and Brenning, U.: Rostock zooplankton studies off West Africa, Helgolander Meeresunters., 49, 829-847, 1995. (Pubitemid 26448771)
94. Redfield, J., Ketchum, B. H., and Richards, F. A.: The influence of organisms on the composition of sea-water, in: The sea, Vol 2, edited by: Hill, M. N., Academic Press, NY, 26-77, 1963.
95. Richards, F. A.: Anoxic basins and fjords, in Chemical Oceanography, edited by: Riley, J. P. and Skirrow, G., 1, Academic Press, NY, 611-645, 1965.
96. Rouault, M., Illig, S., Bartholomae, C., Reason, C. J. C., and Bentamy, A.: Propagation and origin of warm anomalies in the Angola Benguela upwelling system in 2001, J. Marine Syst., 68, 477-488, 2007.
97. Shannon, L. V.: A plan comes together, in: Benguela: Predicting a Large Marine Ecosystem, vol. 14.
98. edited by: Shannon, V., Hempel, G., Malanotte-Rizzoli, P., Moloney, C., and Woods, J., Shchepetkin, A. F. and McWilliams, J. C.: A method for computing horizontal pressure-gradient force in an oceanic model with a nonaligned vertical coordinate, J. Geophys. Res., 108, 3090, doi:10.1029/2001JC001047, 2003.
99. Shchepetkin, A. F. and McWilliams, J. C.: The regional oceanic modeling system (ROMS): a split-explicit, free-surface, topography-following-coordinate oceanic model, Ocean Model., 9, 347-404, doi:10.1016/j.ocemod.2004.08.002, 2005. (Pubitemid 40306754)
100. Stramma, L., Johnson, G. C., Sprintall, J., and Mohrholz, V.: Expanding oxygen-minimum zones in the tropical oceans, Science, 320, 655-658, doi:10.1126/science.1153847, 2008. (Pubitemid 351928345)
101. Stramma, L., Johnson, G. C., Firing, E., and Schmidtko, S.: Eastern Pacific oxygen minimum zones: Supply paths and multidecadal changes, J. Geophys. Res., 115, C09011, doi:201010.1029/2009JC005976, 2010.
102. Strous, M., and Jetten, M. S. M.: Anaerobic oxidation of methane and ammonium, Annu. Rev. Microbiol., 58, 99-117, 2004. (Pubitemid 39551980)
103. Strous, M., Pelletier, E., Mangenot, S., Rattei, T., Lehner, A., Taylor, M. W., Horn, M., Daims, H., Bartol-Mavel, D., Wincker, P., Barbe V., Fonknechten, N., Vallenet, D., Segurens, B., Schenowitz-Truong, C., Medigue, C., Collingro, A., Snel, B., Dutilh, B. E., Op den Camp, H. J. M., van der Drift, C., Cirpus, I., van de Pas-Schoonen, K. T., Harhangi, H. R., van Niftrik, L., Schmid, M., Keltjens, J., van de Vossenberg, J., Kartal, B., Meier, H., Frishman, D., Huy-nen, M. A., Mewes, H.-W., Weissenbach, J., Jetten, M. S. M., Wagner, M., and Le Paslier, D.: Deciphering the evolution and metabolism of an anammox bacterium from a community genome, Nature, 440, 790-794, doi:10.1038/nature04647, 2006.
104. Suntharalingam, P., Sarmiento, J. L., and Toggweiler, J. R.: Global significance of nitrous-oxide production and transport from oceanic low-oxygen zones: A modeling study, Glob. Biogeochem. Cy., 14, 1353-1370, 2000. (Pubitemid 32281842)
105. Suntharalingam, P., Buitenhuis, E., Le Quere, C., Dentener, F., Nevison, C., and Butler, J.: Quantifying the Impact of Anthropogenic Nitrogen Deposition on Oceanic Nitrous Oxide, Geophys. Res. Lett., 39, L07605, doi:10.1029/ 2011GL050778, 2012.
106. Taylor, K. E.: Summarizing multiple aspects of model performance in a single diagram, J. Geophys. Res., 106, 7183-7192, 2001.
107. Tian, R.C., Vezina, A., Legendre, L., Ingram, R. G., Klein, B., Packard, T., Roy, S., Savenkoff, C., Silverberg, N., Therriault, J. C., and Tremblay, J.E.: Effects of pelagic food-web interactions and nutrient remineralization on the biogeochemical cycling of carbon: a modeling approach, Deep Sea Res. Pt II, 47, 637-662, doi:10.1016/S0967-0645(99)00121-6, 2000. (Pubitemid 30063408)
108. Tian, R. C., V́ezina, A. F., Starr, M., and Saucier, F.: Seasonal dynamics of coastal ecosystems and export production at high latitudes: A modeling study, Limnol. Oceanogr., 46, 1845-1859, doi:10.4319/lo.2001.46.8.1845, 2001. (Pubitemid 34009936)
109. Tilstone, G., Smyth, T., Poulton, A., and Hutson, R.: Measured and remotely sensed estimates of primary production in the Atlantic Ocean from 1998 to 2005, Deep Sea Res. Pt II, 56, 918-930, doi:10.1016/j.dsr2.2008.10.034, 2009.
110. Thamdrup, B., Jensen, M. M., Dalsgaard, T., Ulloa, O., Farias, L., and Escribano, R.: Anaerobic ammonium oxidation in the oxygen-deficient waters off northern Chile, Limnol. Oceanogr. 51, 2145-2156, 2006. (Pubitemid 44520911)
111. Toniazzo, T. and Woolnough, S.: Development of warm SST errors in the southern tropical Atlantic in CMIP5 decadal hindcasts, Clim. Dynam, 1-25, doi:10.1007/s00382-013-1691-2, 2013.
112. Tyrrell, T. and Lucas,M. I.: Geochemical evidence of denitrification in the Benguela upwelling system, Cont. Shelf Res., 22, 2497-2511, 2002. (Pubitemid 35371797)
113. Veitch, J., Penven, P., and Shillington, F.: The Benguela: A laboratory for comparative modeling studies, Prog. Oceanogr., 83, 296-302, doi:10.1016/j.pocean.2009.07.008, 2009.
114. Verheye, H. M. and Ekau, W.: Geochemistry and Ecology of the Namibian Upwelling System (GENUS Project) and St Helena Bay Monitoring Line (SHBML), cruise report, FRS Africana, 1 to 17 December 2009, 2009.
115. Walter, S., Bange, H. W., Breitenbach, U., and Wallace, D. W. R.: Nitrous oxide in the North Atlantic Ocean, Biogeosciences, 3, 607-619, doi:10.5194/bg-3-607-2006, 2006. (Pubitemid 44878466)
116. Wanninkhof, R.: Relationship Between Wind Speed and Gas Exchange Over the Ocean, J. Geophys. Res., 97, 7373-7382, 1992.
117. Ware, D. M.: Production characteristics of upwelling systems and the trophodynamic roleof hake, in: Benguela trophic functioning, edited by: Payne, A. I. L., Brink, K. H., Mann, K. H., and Hilborn, R., 501-513, 1992. (Pubitemid 24413539)
118. Weiss, R. F. and Price, B. A.: Nitrous oxide solubility in water and seawater, Mar. Chem., 8, 347-359, doi:10.1016/0304-4203(80)90024-9, 1980. (Pubitemid 10150957)
119. Yakushev, E. V., Pollehne, F., Jost, G., Kuznetso, I., Schneider, B., and Urnlauf, L.: Analysis of the water column oxic/anoxic interface in the Black and Baltic seas with a numerical model, Mar. Chem., 107, 388-410, doi:10.1016/j.marchem.2007.06.003, 2007. (Pubitemid 350089980)
120. Zabel, M., Ahke, A., Aspetsberger, F., Eberwein, A., Ferdelman, T., Franke, C., Heene, T., Heslop, D., Hessler, S., Holzwarth, U., Inthorn, M., Kahl, G., Kasten, S., Langreder, J., Mohrholz, V., Mulitza, S., Nordhausen, A., Ochsenhirt, W.-T., Paul, A., Rajes, B., Rathmann, S., Riedinger, N., Romero, O., Schafer, R., Schmidt, M., von Dobeneck, T., Wilhelm, O., Witte, U., and Wulbers, A.: Report and preliminary results of METEOR Cruise M57/2, Walvis Bay-Walvis Bay, 11 February-12 March 2003, cruise report, Berichte, 220 Fachbereich Geowissenschaften, Universitat Bremen 2003.
121. Zamora, L. M., Oschlies, A., Bange, H. W., Huebert, K. B., Craig, J. D., Kock, A., and L̈oscher, C. R.: Nitrous oxide dynamics in low oxygen regions of the Pacific: insights from the MEMENTO database, Biogeosciences, 9, 5007-5022, doi:10.5194/bg-9-5007-2012, 2012. supporting GENUS