References

Biogeosciences

1726-4189

Copernicus Publications

Göttingen, Germany

10.5194/bg-11-7125-2014

Drivers of trophic amplification of ocean productivity trends in a changing climate

Stock

C. A.

¹ Dunne

J. P.

¹ John

J. G.

https://orcid.org/0000-0003-2696-277X

NOAA Geophysical Fluid Dynamics Laboratory, 201 Forrestal Road, Princeton, NJ 08540-6649, USA

16 12 2014

11 24 7125 7135

2014

This work is licensed under the Creative Commons Attribution 3.0 Unported License. To view a copy of this licence, visit https://creativecommons.org/licenses/by/3.0/

This article is available from https://bg.copernicus.org/articles/11/7125/2014/bg-11-7125-2014.html

The full text article is available as a PDF file from https://bg.copernicus.org/articles/11/7125/2014/bg-11-7125-2014.pdf

Pronounced projected 21st century trends in regional oceanic net primary production (NPP) raise the prospect of significant redistributions of marine resources. Recent results further suggest that NPP changes may be amplified at higher trophic levels. Here, we elucidate the role of planktonic food web dynamics in driving projected changes in mesozooplankton production (MESOZP) found to be, on average, twice as large as projected changes in NPP by the latter half of the 21st century under a high emissions scenario in the Geophysical Fluid Dynamics Laboratory's ESM2M–COBALT (Carbon, Ocean Biogeochemistry and Lower Trophics) earth system model. Globally, MESOZP was projected to decline by 7.9% but regional MESOZP changes sometimes exceeded 50%. Changes in three planktonic food web properties – zooplankton growth efficiency (ZGE), the trophic level of mesozooplankton (MESOTL), and the fraction of NPP consumed by zooplankton (zooplankton–phytoplankton coupling, ZPC), explain the projected amplification. Zooplankton growth efficiencies (ZGE) changed with NPP, amplifying both NPP increases and decreases. Negative amplification (i.e., exacerbation) of projected subtropical NPP declines via this mechanism was particularly strong since consumers in the subtropics have limited surplus energy above basal metabolic costs. Increased mesozooplankton trophic level (MESOTL) resulted from projected declines in large phytoplankton production. This further amplified negative subtropical NPP declines but was secondary to ZGE and, at higher latitudes, was often offset by increased ZPC. Marked ZPC increases were projected for high-latitude regions experiencing shoaling of deep winter mixing or decreased winter sea ice – both tending to increase winter zooplankton biomass and enhance grazer control of spring blooms. Increased ZPC amplified projected NPP increases in the Arctic and damped projected NPP declines in the northwestern Atlantic and Southern Ocean. Improved understanding of the physical and biological interactions governing ZGE, MESOTL and ZPC is needed to further refine estimates of climate-driven productivity changes across trophic levels.

References 1

Anderson, J. L., Balaji, V., Broccoli, A. J., Cooke, W. F., Delworth, T. L., Dixon, K. W., Donner, L. J., Dunne, K. A., Freidenreich, S. M., Garner, S. T., Gudgel, R. G., Gordon, C. T., Held, I. M., Hemler, R. S., Horowitz, L. W., Klein, S. A., Knutson, T. R., Kushner, P. J., Langenhost, A. R., Lau, N. C., Liang, Z., Malyshev, S. L., Milly, P. C. D., Nath, M. J., Ploshay, J. J., Ramaswamy, V., Schwarzkopf, M. D., Shevliakova, E., Sirutis, J. J., Soden, B. J., Stern, W. F., Thompson, L. A., Wilson, R. J., Wittenberg, A. T., Wyman, B. L., and Dev, G. G. A. M.: The new GFDL global atmosphere and land model AM2-LM2: Evaluation with prescribed SST simulations, J. Climate, 17, 4641–4673, 2004.

Armstrong, R. A., Lee, C., Hedges, J. I., Honjo, S., and Wakeham, S. G.: A new, mechanistic model for organic carbon fluxes in the ocean based on the quantitative association of POC with ballast minerals, Deep-Sea Res.-Pt. II, 49, 219–236, 2002.

Aumont, O., Maier-Reimer, E., Blain, S., and Monfray, P.: An ecosystem model of the global ocean including Fe, Si, P co-limitation, Global Biogeochem. Cy., 17, 1060, <a href="http://dx.doi.org/10.1029/2001GB001745">https://doi.org/10.1029/2001GB001745</a>, 2003.

Baines, S. B. and Pace, M. L.: The production of dissolved organic matter by phytoplankton and its importance to bacteria: Patterns across marine and freshwater systems, Limnol. Oceanogr., 36, 1078–1090, 1991.

Barange, M., Merino, G., Blanchard, J. L., Scholtens, J., Harle, J., Allison, E. H., Allen, J. I., Holt, J., and Jennings, S.: Impacts of climate change on marine ecosystem production in societies dependent on fisheries, Nature Climate Change, 4, 211–216, <a href="http://dx.doi.org/10.1038/Nclimate2119">https://doi.org/10.1038/Nclimate2119</a>, 2014.

Bopp, L., Monfray, P., Aumont, O., Dufresne, J. L., Le Treut, H., Madec, G., Terray, L., and Orr, J. C.: Potential impact of climate change on marine export production, Global Biogeochem. Cy., 15, 81–99, 2001.

Bopp, L., Resplandy, L., Orr, J. C., Doney, S. C., Dunne, J. P., Gehlen, M., Halloran, P., Heinze, C., Ilyina, T., Séférian, R., Tjiputra, J., and Vichi, M.: Multiple stressors of ocean ecosystems in the 21st century: projections with CMIP5 models, Biogeosciences, 10, 6225–6245, <a href="http://dx.doi.org/10.5194/bg-10-6225-2013">https://doi.org/10.5194/bg-10-6225-2013</a>, 2013.

Burd, A. B. and Jackson, G. A.: Particle aggregation, Annual Review of Marine Science, 1, 65–90, <a href="http://dx.doi.org/10.1146/annurev.marine.010908.163904">https://doi.org/10.1146/annurev.marine.010908.163904</a>, 2009.

Calbet, A. and Landry, M. R.: Phytoplankton growth, microzooplankton grazing, and carbon cycling in marine systems, Limnol. Oceanogr., 49, 51–57, 2004.

Carlotti, F., Giske, J., and Werner, F. E.: Modeling zooplankton dynamics, in: Zooplankton methodology manual, edited by: Harris, R. P., Wiebe, P. H., Lenz, J., Skjoldal, H. R., and Huntley, M. E., Elsevier, New York, 571–667, 2000.

Chai, F., Dugdale, R. C., Peng, T. H., Wilkerson, F. P., and Barber, R. T.: One-dimensional ecosystem model of the equatorial pacific upwelling system. Part I: Model development and silicon and nitrogen cycle, Deep-Sea Res.-Pt. II, 49, 2713–2745, 2002.

Chust, G., Allen, J. I., Bopp, L., Schrum, C., Holt, J., Tsiaras, K., Zavatarelli, M., Chifflet, M., Cannaby, H., Dadou, I., Daewel, U., Wakelin, S. L., Machu, E., Pushpadas, D., Butenschon, M., Artioli, Y., Petihakis, G., Smith, C., Garcon, V., Goubanova, K., Le Vu, B., Fach, B. A., Salihoglu, B., Clementi, E., and Irigoien, X.: Biomass changes and trophic amplification of plankton in a warmer ocean, Glob. Change Biol., 20, 2124–2139, <a href="http://dx.doi.org/10.1111/gcb.12562">https://doi.org/10.1111/gcb.12562</a>, 2014.

Dahms, H. U.: Dormancy in the copepoda – an overview, Hydrobiologia, 306, 199–211, <a href="http://dx.doi.org/10.1007/Bf00017691">https://doi.org/10.1007/Bf00017691</a>, 1995.

del Giorgio, P. A. and Cole, J. J.: Bacterial energetics and growth efficiency, in: Microbial ecology of the oceans, edited by: Kirchman, D. L., John Wiley and Sons, Inc., New York, 289–325, 2000.

Delworth, T. L., Broccoli, A. J., Rosati, A., Stouffer, R. J., Balaji, V., Beesley, J. A., Cooke, W. F., Dixon, K. W., Dunne, J., Dunne, K. A., Durachta, J. W., Findell, K. L., Ginoux, P., Gnanadesikan, A., Gordon, C. T., Griffies, S. M., Gudgel, R., Harrison, M. J., Held, I. M., Hemler, R. S., Horowitz, L. W., Klein, S. A., Knutson, T. R., Kushner, P. J., Langenhorst, A. R., Lee, H., Lin, S., Lu, J., Malyshev, S. L., Milly, P. C. D., Ramaswamy, V., Russell, J., Schwarzkopf, M. D., Shevliakova, E., Sirutis, J. J., Spelman, M. J., Stern, W. F., Winton, M., Wittenberg, A. T., Wyman, B., Zeng, F., and Zhang, R.: GFDL's CM2 global coupled climate models, Part I: Formulation and simulation characteristics, J. Climate, 19, 643–674, 2006.

Doney, S. C.: Oceanography – plankton in a warmer world, Nature, 444, 695–696, 2006.

Doney, S. C., Glover, D. M., and Najjar, R. J.: A new coupled, one-dimensional biological-physical model for the upper ocean: Applications to the JGOFS bermuda atlantic time-series study (BATS) site, Deep-Sea Res.-Pt. II, 43, 591–624, 1996.

Dunne, J. P., Armstrong, R. A., Gnanadesikan, A., and Sarmiento, J. L.: Empirical and mechanistic models for the particle export ratio, Global Biogeochem. Cy., 19, Gb4026, <a href="http://dx.doi.org/10.1029/2004gb002390">https://doi.org/10.1029/2004gb002390</a>, 2005.

Dunne, J. P., John, J. G., Adcroft, A. J., Griffies, S. M., Hallberg, R. W., Shevliakova, E., Stouffer, R. J., Cooke, W., Dunne, K. A., Harrison, M. J., Krasting, J. P., Malyshev, S. L., Milly, P. C. D., Phillipps, P. J., Sentman, L. T., Samuels, B. L., Spelman, M. J., Winton, M., Wittenberg, A. T., and Zadeh, N.: GFDL's ESM2 global coupled climate-carbon earth system models, Part I: Physical formulation and baseline simulation characteristics, J. Climate, 25, 6646–6665, <a href="http://dx.doi.org/10.1175/Jcli-D-11-00560.1">https://doi.org/10.1175/Jcli-D-11-00560.1</a>, 2012.

Dunne, J. P., John, J. G., Shevliakova, E., Stouffer, R. J., Krasting, J. P., Malyshev, S. L., Milly, P. C. D., Sentman, L. T., Adcroft, A. J., Cooke, W., Dunne, K. A., Griffies, S. M., Hallberg, R. W., Harrison, M. J., Levy, H., Wittenberg, A. T., Phillips, P. J., and Zadeh, N.: GFDL's ESM2 global coupled climate-carbon earth system models, Part II: Carbon system formulation and baseline simulation characteristics, J. Climate, 26, 2247–2267, <a href="http://dx.doi.org/10.1175/Jcli-D-12-00150.1">https://doi.org/10.1175/Jcli-D-12-00150.1</a>, 2013.

Edwards, K. F., Thomas, M. K., Klausmeier, C. A., and Litchman, E.: Allometric scaling and taxonomic variation in nutrient utilization traits and maximum growth rate of phytoplankton, Limnol. Oceanogr., 57, 554–566, <a href="http://dx.doi.org/10.4319/lo.2012.57.2.0554">https://doi.org/10.4319/lo.2012.57.2.0554</a>, 2012.

Eppley, R. W.: Temperature and phytoplankton growth in the sea, Fisheries Bulletin, 70, 1063–1085, 1972.

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.

Flato, G., Marotzke, J., Abiodun, B., Braconnot, P., Chou, S. C., Collins, W., Cox, P., Driouech, F., Emori, S., Eyring, V., Forest, C., Gleckler, P., Guilyardi, E., Jakob, C., Kattsov, V., Reason, C., and Rummukainen, M.: Evaluation of climate models, in: Climate change 2013: The physical science basis, Contributions of working group I to the fifth assessment report of the intergovernmental panel on climate change, edited by: Stocker, T. F., Qin, D., Plattner, G.-K., Tignor, M., Allen, S. K., Boschung, J., Nauels, A., Xia, Y., Bex, V., and Midgley, P. M., Cambridge University Press, Cambridge, UK, 741–866, 2013.

Flynn, K. J.: Incorporating plankton respiration in models of aquatic ecosystem function, in: Respiration in aquatic ecosystems, edited by: del Giorgio, P. A. and Williams, P. J. B., Oxford University Press, Oxford, 248–266, 2005.

Friedland, K. D., Stock, C., Drinkwater, K. F., Link, J. S., Leaf, R. T., Shank, B. V., Rose, J. M., Pilskaln, C. H., and Fogarty, M. J.: Pathways between primary production and fisheries yields of large marine ecosystems, Plos One, 7, e28945, <a href="http://dx.doi.org/10.1371/journal.pone.0028945">https://doi.org/10.1371/journal.pone.0028945</a>, 2012.

Fuchs, H. L. and Franks, P. J. S.: Plankton community properties determined by nutrients and size-selective feeding, Mar. Ecol.-Prog. Ser., 413, 1–15, <a href="http://dx.doi.org/10.3354/meps08716">https://doi.org/10.3354/meps08716</a>, 2010.

Fuhrman, J.: Impact of viruses on bacterial processes, in: Microbial ecology of the oceans, 1st ed., edited by: Kirchman, D. L., Wiley series in ecology and microbiology, John Wiley and Sons, New York, 327–350, 2000.

Geider, R. J.: Respiration: Taxation without representation, in: Primary productivity and biogeochemical cycles in the sea, edited by: Falkowski, P. G. and Woodhead, A. D., Plenum Press, New York, 333–360, 1992.

Geider, R., MacIntyre, H. L., and Kana, T. M.: Dynamic model of phytoplankton growth and acclimation: Responses of the balanced growth rate and chlorophyll a: carbon ratio to light, nutrient limitation and temperature, Mar. Ecol.-Prog. Ser., 148, 187–200, 1997.

Griffies, S. M.: Elements of MOM4p1, NOAA/GFDL, Princeton, NJ, 377, 444 pp., 2009.

Griffies, S. M., Gnanadesikan, A., Dixon, K. W., Dunne, J. P., Gerdes, R., Harrison, M. J., Rosati, A., Russell, J. L., Samuels, B. L., Spelman, M. J., Winton, M., and Zhang, R.: Formulation of an ocean model for global climate simulations, Ocean Science, 1, 45–79, 2005.

Guilyardi, E., Wittenberg, A., Fedorov, A., Collins, M., Wang, C. Z., Capotondi, A., van Oldenborgh, G. J., and Stockdale, T.: Understanding El Nino in ocean-atmosphere general circulation models progress and challenges, B. Am. Meteorol. Soc., 90, 325–340, <a href="http://dx.doi.org/10.1175/2008bams2387.1">https://doi.org/10.1175/2008bams2387.1</a>, 2009.

Hansen, B. W., Bjornsen, P. K., and Hansen, P. J.: The size ratio between planktonic predators and their prey, Limnol. Oceanogr., 39, 395–402, 1994.

Hansen, P. J., Bjornsen, P. K., and Hansen, B. W.: Zooplankton grazing and growth: Scaling within the 2-2000-μm body size range, Limnol. Oceanogr., 42, 687–704, 1997.

Hegerl, G. C., Zwiers, F. W., Braconnot, P., Gillett, N. P., Luo, Y., Marengo Orsini, J. A., Nicholls, N., Penner, J. E., and Stott, P. A.: Understanding and attributing climate change, in: Climate change 2007: The physical science basis, Contribution of working group I to the fourth assessment report of the intergovernmental panel on climate edited by: Solomon, S., Qin, D., Manning, M., Chen, Z., Marquis, M., Averyt, K. B., Tignor, M., and Miller, H. L., Cambridge University Press, Cambridge UK and New York, NY, USA, 663–745, 2007.

Jackson, G. A.: A model of the formation of marine algal flocs by physical coagulation processes, Deep-Sea Res.-Pt. I, 37, 1197–1211, 1990.

Johnson, C. L., Leising, A. W., Runge, J. A., Head, E. J. H., Pepin, P., Plourde, S., and Durbin, E. G.: Characteristics of calanus finmarchicus dormancy patterns in the northwest atlantic, ICES J. Mar. Sci., 65, 339–350, <a href="http://dx.doi.org/10.1093/icesjms/fsm171">https://doi.org/10.1093/icesjms/fsm171</a>, 2008.

Jumars, P. A., Penry, D. L., Baross, J. A., Perry, M. J., and Frost, B. W.: Closing the microbial loop: Dissolved carbon pathway to heterotrophic bacteria from incomplete ingestion, digestion, and absorption in animals, Deep-Sea Res., 36, 483–495, 1989.

Kearney, K. A., Stock, C., and Sarmiento, J. L.: Amplification and attenuation of increased primary production in a marine food web, Mar. Ecol.-Prog. Ser., 491, 1–14, <a href="http://dx.doi.org/10.3354/Meps10484">https://doi.org/10.3354/Meps10484</a>, 2013.

Kishi, M. J., Kashiwai, M., Ware, D. M., Megrey, B. A., Eslinger, D. L., Werner, F. E., Noguchi-Aita, M., Azumaya, T., Fuji, M., Hashimoto, S., Huang, D., Iizumi, H., Ishida, Y., Kang, S., Kantakov, G. A., Kim, H.-c., Komatsu, K., Navrotsky, V. V., Smith, S. L., Tadokoro, K., Tsuda, A., Yamamura, O., Yamanaka, Y., Yakouchi, K., Yoshi, N., Zhang, J., Zuenko, Y. I., and Zvalinsky, V. I.: Nemuro-a lower trophic level model for the north pacific marine ecosystem, Ecol. Model., 202, 12–25, 2007.

Klaas, C. and Archer, D. E.: Association of sinking organic matter with various types of mineral ballast in the deep sea: Implications for the rain ratio, Global Biogeochem. Cy., 16, 1116, <a href="http://dx.doi.org/10.1029/2001GB001765">https://doi.org/10.1029/2001GB001765</a>, 2002.

Landry, M. R., Al-Mutairi, H., Selph, K. E., Christensen, S., and Nunnery, S.: Seasonal patterns of mesozooplankton abundance and biomass at station aloha, Deep-Sea Res.-Pt. II, 48, 2037–2061, 2001.

Lin, S. J.: A "vertically lagrangian" finite-volume dynamical core for global models, Mon. Weather Rev., 132, 2293–2307, <a href="http://dx.doi.org/10.1175/1520-0493(2004)132<2293:Avlfdc>2.0.Co;2">https://doi.org/10.1175/1520-0493(2004)132<2293:Avlfdc>2.0.Co;2</a>, 2004.

Mauchline, J.: The biology of calanoid copepods, Advances in marine biology, Academic Press, San Diego, 710 pp., 1998.

Meehl, G. A., Stocker, T. F., Collins, W. D., Friedlingstein, P., Gaye, A. T., Gregory, J. M., Kitoh, A., Knutti, R., Murphy, J. M., Noda, A., Raper, S. P. B., Watterson, I. G., Weaver, A. J., and Zhao, Z.-C.: Global climate projections, in: Climate change 2007: The physical science basis. Contribution of working group I to the fourth assessment report of the intergovernmental panel on climate change, edited by: Solomon, S., Qin, D., Manning, M., Chen, Z., Marquis, M., Averyt, K. B., Tignor, M., and Miller, H. L., Cambridge University Press, Cambridge, United Kingdom and New York, NY, USA., 589–662, 2007.

Merino, G., Barange, M., Blanchard, J. L., Harle, J., Holmes, R., Allen, I., Allison, E. H., Badjeck, M. C., Dulvy, N. K., Holt, J., Jennings, S., Mullon, C., and Rodwell, L. D.: Can marine fisheries and aquaculture meet fish demand from a growing human population in a changing climate?, Global Environ. Chang., 22, 795–806, 2012.

Munk, W. H. and Riley, G. A.: Absorption of nutrients by aquatic plants, J. Mar. Res., 11, 215–240, 1952.

Nagata, T.: Production mechanisms of dissolved organic matter, in: Microbial ecology of the oceans, edited by: Kirchman, D. L., John Wiley and Sons, Inc., New York, 121–152, 2000.

O'Brien, T. D.: Copepod: A global plankton database, Technical Memorandum, 19, 136 pp., 2005.

Pauly, D., Christensen, V., and Walters, C.: Ecopath, Ecosim, and Ecospace as tools for evaluating ecosystem impact of fisheris, ICES J. Mar. Sci., 57, 697–706, 2000.

Record, N. R., Pershing, A. J., and Maps, F.: Emergent copepod communities in an adaptive trait-structured model, Ecol. Model., 260, 11–24, <a href="http://dx.doi.org/10.1016/j.ecolmodel.2013.03.018">https://doi.org/10.1016/j.ecolmodel.2013.03.018</a>, 2013.

Reichler, T. and Kim, J.: How well do coupled models simulate today's climate?, B. Am. Meteorol. Soc., 89, 303–311, <a href="http://dx.doi.org/10.1175/BAMS-89-3-303">https://doi.org/10.1175/BAMS-89-3-303</a>, 2008.

Riahi, K., Rao, S., Krey, V., Cho, C. H., Chirkov, V., Fischer, G., Kindermann, G., Nakicenovic, N., and Rafaj, P.: RCP 8.5 – a scenario of comparatively high greenhouse gas emissions, Clim. Change, 109, 33–57, <a href="http://dx.doi.org/10.1007/s10584-011-0149-y">https://doi.org/10.1007/s10584-011-0149-y</a>, 2011.

Roman, M. R., Adolf, H. A., Landry, M. R., Madin, L. P., Steinberg, D. K., and Zhang, X.: Estimates of oceanic mesozooplankton production: A comparison using the Bermuda and Hawaii time-series data, Deep-Sea Res.-Pt. II, 49, 175–192, 2002.

Rose, K. A., Allen, J. I., Artioli, Y., Barange, M., Blackford, J., Carlotti, F., Cropp, R., Daewel, U., Edwards, K., Flynn, K., Hill, S., Hille Ris Lambers, R., Huse, G., Mackinson, S., Megrey, B. A., Moll, A., Rivkin, R., Salihoglu, B., Schrum, C., Shannon, L., Shin, Y., Smith, S. L., Smith, C., Solidoro, C., St John, M., and Zhou, M.: End-to-end models for the analysis of marine ecosystems: Challenges, issues, and next steps, Marine and Coastal Fisheries: Dynamics, Management and Ecosystem Science, 2, 115–130, 2010.

Ruggio, R., Vichi, M., Paparella, F., and Masina, S.: Climatic trends of the equatorial undercurrent: A backup mechanism for sustaining the equatorial pacific production, J. Marine Syst., 121, 11–23, <a href="http://dx.doi.org/10.1016/j.jmarsys.2013.04.001">https://doi.org/10.1016/j.jmarsys.2013.04.001</a>, 2013.

Rykaczewski, R. R. and Dunne, J. P.: Enhanced nutrient supply to the California Current ecosystem with global warming and increased stratification in an earth system model, Geophys. Res. Lett., 37, L21606, <a href="http://dx.doi.org/10.1029/2010GL045019">https://doi.org/10.1029/2010GL045019</a>, 2010.

Ryther, J. H.: Photosynthesis and fish production in the sea, Science, 166, 72–76, 1969.

Sieburth, J. M., Smetacek, V., and Lenz, J.: Pelagic ecosystem structure: Heterotrophic compartments of the plankton and their relationship to plankton size fractions, Limnol. Oceanogr., 23, 1256–1263, 1978.

Steinacher, M., Joos, F., Frölicher, T. L., Bopp, L., Cadule, P., Cocco, V., Doney, S. C., Gehlen, M., Lindsay, K., Moore, J. K., Schneider, B., and Segschneider, J.: Projected 21st century decrease in marine productivity: a multi-model analysis, Biogeosciences, 7, 979–1005, <a href="http://dx.doi.org/10.5194/bg-7-979-2010">https://doi.org/10.5194/bg-7-979-2010</a>, 2010.

Stock, C. A. and Dunne, J. P.: Controls on the ratio of mesozooplankton production to primary production in marine ecosystems, Deep-Sea Res.-Pt. I, 57, 95–112, 2010.

Stock, C. A., Powell, T. M., and Levin, S. A.: Bottom-up and top-down forcing in a simple size-structured plankton dynamics model, J. Marine Syst., 74, 134–152, 2008.

Stock, C. A., Alexander, M. A., Bond, N. A., Brander, K. M., Cheung, W. W. L., Curchitser, E. N., Delworth, T. L., Dunne, J. P., Griffies, S. M., Haltuch, M. A., Hare, J. A., Hollowed, A. B., Lehodey, P., Levin, S. A., Link, J. S., Rose, K. A., Rykaczewski, R. R., Sarmiento, J. L., Stouffer, R. J., Schwing, F. B., Vecchi, G. A., and Werner, F. E.: On the use of IPCC-class models to assess the impact of climate on living marine resources, Prog. Oceanogr., 88, 1–27, 2011.

Stock, C. A., Dunne, J. P., and John, J. G.: Global-scale carbon and energy flows through the marine planktonic food web: An analysis with a coupled physical-biological model, Prog. Oceanogr., 120, 1–28, <a href="http://dx.doi.org/10.1016/j.pocean.2013.07.001">https://doi.org/10.1016/j.pocean.2013.07.001</a>, 2014.

Straile, D.: Gross growth efficiencies of protozoan and metazoan zooplankton and their dependence on food concentration, predator-prey weight ratio, and taxonomic group, Limnol. Oceanogr., 42, 1375–1385, 1997.

Sumaila, U. R., Cheung, W. W. L., Lam, V. W. Y., Pauly, D., and Herrick, S.: Climate change impacts on the biophysics and economics of world fisheries, Nature Climate Change, 1, 449–456, <a href="http://dx.doi.org/10.1038/Nclimate1301">https://doi.org/10.1038/Nclimate1301</a>, 2011.

Suttle, C. A.: The significance of viruses to mortality in aquatic microbial communities, Microb. Ecol., 28, 237–243, 1994.

Winton, M., Adcroft, A., Griffies, S. M., Hallberg, R. W., Horowitz, L. W., and Stouffer, R. J.: Influence of ocean and atmosphere components on simulated climate sensitivities, J. Climate, 26, 231–245, <a href="http://dx.doi.org/10.1175/Jcli-D-12-00121.1">https://doi.org/10.1175/Jcli-D-12-00121.1</a>, 2013.