Biogeosciences www.biogeosciences.net 1726-4170 1726-4189 7 1 2010 10.5194/bg-7-371-2010 http://www.biogeosciences.net/7/371/2010/ http://www.biogeosciences.net/7/371/2010/bg-7-371-2010.html http://www.biogeosciences.net/7/371/2010/bg-7-371-2010.pdf 371 386 2010-01-29 Interannual variability of alongshore spring bloom dynamics in a coastal sea caused by the differential influence of hydrodynamics and light climate G. Brandt brandt@bpt-info.de K. W. Wirtz GKSS-Research Centre, Institute for Coastal Research, Max-Planck-Str. 1, 21502 Geesthacht, Germany currently at: Bundesamt für Seeschifffahrt und Hydrographie, Bernhard-Nocht-Str. 78, 20359 Hamburg, Germany Timing and spatial distribution of phytoplankton blooms in coastal oceans are highly variable. The interactions of various biological and physical factors leading to the observed variability are complex and remain poorly understood. We present an example for distinct differences in the spatio-temporal chlorophyll <i>a</i> (CHL-<i>a</i>) distribution on an interannual scale, integrating high-frequency data from an autonomous measuring device (FerryBox), which operated on an alongshore route in the coastal German Bight (North Sea). While in one year the distribution of CHL-<i>a</i> was spatially homogeneous (2004), a bloom only developed in one part of the transect in the following spring period (2005). We use a one-dimensional Lagrangian particle tracking model, which operates along the mean current direction, combined with a NPZ-model to identify the mechanisms controlling the observed interannual bloom variability on the alongshore transect. Our results clearly indicate that in 2004 the local light climate determined the spatial and temporal dynamics of the spring bloom. In contrast, the import of a water mass with elevated CHL-<i>a</i> concentrations from the adjacent Southern Bight triggered the spring bloom in 2005. The inflow event did, however, not last long enough to spread the bloom into the eastern part of the study area, where high turbidity prevented local phytoplankton growth. The model identifies two interacting mechanisms, light climate and hydrodynamics, that controlled the alongshore dynamics. Especially the occurrence of a pronounced spring bloom despite unfavourable light conditions in 2005 underlines the need to carefully consider hydrodynamics to understand the dynamics of the plankton community in coastal environments. Ainsworth, C.: Oceanography: FerryBoxes Begin to Make Waves, Science, 322, 1627, 2008. Beddig, S., Brockmann, U., Dannecker, W., Körner, D., Pohlmann, T., Puls, W., Radach, G., Rebers, A., Rick, H J., Schatzmann, M., et al.: Nitrogen fluxes in the German Bight, Mar. Pollut. Bull., 34, 382–394, 1997. Behrenfeld, M J. and Falkowski, P G.: Photosynthetic rates derived from satellite-based chlorophyll concentration, Limnol. Oceanogr., 42, 1–20, 1997. Behrenfeld, M J., Boss, E., Siegel, D A., and Shea, D M.: Carbon-based ocean productivity and phytoplankton physiology from space, Global Biogeochem. Cy., 19, doi:10.1029/2004GB002299, 2005. Brandt, G., Wehrmann, A., and Wirtz, K.: Rapid invasion of \emphCrassostrea gigas into the German Wadden Sea dominated by larval supply, J. Sea Res., 59, 279–296, 2008. Cadée, G C. and Hegeman, J.: Phytoplankton in the Marsdiep at the end of the 20th century; 30 years monitoring biomass, primary production, and Phaeocystis blooms, J. Sea Res., 48, 97–110, 2002. Cloern, J E.: An empirical model of the phytoplankton chlorophyll: Carbon ratio – the conversion factor between productivity and growth rate, Limnol. Oceanogr., 40, 1313–1321, 1995. Cloern, J E.: Phytoplankton bloom dynamics in coastal ecosystems: A review with some general lessons from sustained investigation of San Francisco Bay, California, Rev. Geophys., 34, 127–168, 1996. Conover, R J.: Assimilation of organic matter by zooplankton, Limnol. Oceanogr., 11, 338–345, 1966. Davies-Colley, R J. and Smith, D G.: Turbidity, suspended sediment, and water clarity: A review, J. Am. Water. Resour. As., 37, 1085–1101, 2001. Devlin, M J., Barry, J., Mills, D K., Gowen, R J., Foden, J., Sivyer, D., and Tett, P.: Relationships between suspended particulate material, light attenuation and Secchi depth in UK marine waters, Estuar., Coast. and Shelf Sci., 79, 429–439, 2008. Ebenhöh, W., Baretta-Bekker, J G., and Baretta, J W.: The primary production model in the ecosystem model ERSEM II, J. Sea Res., 38, 173–192, 1997. Edwards, A M. and Brindley, J.: Oscillatory behaviour in a three-component plankton population model, Dynam. Stabil. Syst., 11, 347–370, 1996. Elser, J., Sterner, R., Galford, A., Chrzanowski, T., Findlay, D., Mills, K., Paterson, M., Stainton, M., and Schindler, D.: Pelagic C: N: P stoichiometry in a eutrophied lake: responses to a whole-lake food-web manipulation, Ecosystems, 3, 293–307, 2000. Faure, V., Pinazo, C., Torréton, J P., and Douillet, P.: Relevance of various formulations of phytoplankton chlorophyll \empha: carbon ratio in a 3D marine ecosystem model, C. R. Biologies, 329, 813–822, 2006. Furnas, M J.: In situ growth rates of marine phytoplankton: approaches to measurement, community and species growth rates, J. Plankton Res., 12, 1117–1151, 1990. Geider, R J.: Light and temperature depenence of the carbon to chlorophyll \empha ratio in microalgae and cyanobacteria: implications for physiology and growth of phytoplankton, New Phytol., 106, 1–34, 1987. Geider, R J., MacIntyre, H L., and Kana, T M.: A dynamic regulatory model of phytoplanktonic acclimation to light, nutrients, and temperature, Limnol. Oceanogr., 43, 679–694, 1998. Gentleman, W., Leising, A., Frost, B., Strom, S., and Murray, J.: Functional responses for zooplankton feeding on multiple resources: a review of assumptions and biological dynamics, Deep-Sea Res. Pt II, 50, 2847–2875, 2003. Gismervik, I.: Stoichiometry of some marine planktonic crustaceans, J. Plankton Res., 19, 279–285, 1997. Greve, W., Reiners, F., Nast, J., and Hoffmann, S.: Helgoland Roads meso- and macrozooplankton time-series 1974 to 2004: lessons from 30 years of single spot, high frequency sampling at the only off-shore island of the North Sea, Helgol. Mar. Res., 58, 274–288, 2004. Halsband-Lenk, C., Nival, S., Carlotti, F., and Hirche, H.: Seasonal cycles of egg production of two planktonic copepods, \emphCentropages typicus and \emphTemora stylifera, in the north-western Mediterranean Sea, J. Plankton Res., 23, 597–609, 2001. Hansen, P J., Bjornsen, P K., and Hansen, B W.: Zooplankton grazing and growth: Scaling within the 2&minus;2000 \unit\mum body size range, Limnol. Oceanogr., 42, 687–704, 1997. Hecky, R E. and Kilham, P.: Nutrient limitation of phytoplankton in freshwater and marine environments: A review of recent evidence on the effects of enrichment., Limnol. Oceanogr., 33, 796–822, 1988. Hirst, A G. and Bunker, A J.: Growth of marine planktonic copepods: global rates and patterns in relation to chlorophyll \empha, temperature, and body weight, Limnol. Oceanogr., 48, 1988–2010, 2003. Iriarte, A. and Purdie, D A.: Factors controlling the timing of major spring bloom events in an UK south coast estuary, Estuar., Coast. and Shelf Sci., 61, 679–690, 2004. Irigoien, X., Flynn, K J., and Harris, R P.: Phytoplankton blooms: a "loophole" in microzooplankton grazing impact?, J. Plankton Res., 27, 313–321, 2005. Joint, I. and Pomroy, A.: Phytoplankton biomass and production in the southern North Sea, Mar. Ecol. Prog. Ser., 99, 169–169, 1993. Klausmeier, C., Litchman, E., and Levin, S.: Phytoplankton growth and stoichiometry under multiple nutrient limitation, Limnol. Oceanogr., 49, 1463–1470, 2004. Kuipers, B R. and van Noort, G J.: Towards a natural Wadden Sea?, J. Sea Res., 60, 44–53, 2008. Lacroix, G., Ruddick, K., Park, Y., Gypens, N., and Lancelot, C.: Validation of the 3D biogeochemical model MIRO&CO with field nutrient and phytoplankton data and MERIS-derived surface chlorophyll \empha images, J. Marine Sys., 64, 66–88, 2007. Levin, S.: The problem of pattern and scale in ecology: the Robert H. MacArthur award lecture, Ecology, 73, 1943–1967, 1992. Llewellyn, C A., Fishwick, J R., and Blackford, J.: Phytoplankton community assemblage in the English Channel: a comparison using chlorophyll a derived from HPLC-CHEMTAX and carbon derived from microscopy cell counts, J. Plankton Res., 27, 103–119, 2005. Loebl, M. and Beusekom, J. E. E V.: Seasonality of microzooplankton grazing in the northern Wadden Sea, J. Sea Res., 59, 203–216, 2008. Loebl, M., Colijn, F., van Beusekom, J. E E., Baretta-Bekker, J G., Lancelot, C., Philippart, C. J M., Rousseau, V., and Wiltshire, K H.: Recent patterns in potential phytoplankton limitation along the Northwest European continental coast, J. Sea Res., 61, 34–43, submitted, 2010. Lucas, L., Koseff, J., Cloern, J., Monismith, S., and Thompson, J.: Processes governing phytoplankton blooms in estuaries. I: The local production-loss balance, Mar. Ecol. Prog. Ser., 187, 1–15, 1999a. Lucas, L., Koseff, J., Monismith, S., Cloern, J., and Thompson, J.: Processes governing phytoplankton blooms in estuaries 2: The role of horizontal transport, Mar. Ecol. Prog. Ser., 187, 17–30, 1999b. Lucas, L., Koseff, J., Monismith, S., and Thompson, J.: Shallow water processes govern system-wide phytoplankton bloom dynamics: A modeling study, J. Marine Sys., 75, 70–86, 2009. Macedo, M F., Duarte, P., Mendes, P., and Ferreira, J G.: Annual variation of environmental variables, phytoplankton species composition and chotosynthetic carameters in a coastal lagoon, J. Plankton Res., 23, 719–732, 2001. Martin, A.: Phytoplankton patchiness: the role of lateral stirring and mixing, Prog. Oceanogr., 57, 125–174, 2003. May, C L., Koseff, J R., Lucas, L V., Cloern, J E., and Schoellhamer, D H.: Effects of spatial and temporal variability of turbidity on phytoplankton blooms, Mar. Ecol. Prog. Ser., 254, 111–128, 2003. Muylaert, K., Gonzales, R., Franck, M., Lionard, M., Van~der Zee, C., Cattrijsse, A., Sabbe, K., Chou, L., and Vyverman, W.: Spatial variation in phytoplankton dynamics in the Belgian coastal zone of the North Sea studied by microscopy, HPLC-CHEMTAX and underway fluorescence recordings, J. Sea Res., 55, 253–265, 2006. Nielsen, T G.: Contribution of zooplankton grazing to the decline of a Ceratium bloom, Limnol. Oceanogr., 36, 1091–1106, 1991. Pahlow, M.: Linking chlorophyll-nutrient dynamics to the Redfield N:C ratio with a model of optimal phytoplankton growth, Mar. Ecol. Prog. Ser., 287, 33–43, 2005. Peperzak, L., Colijn, F., Gieskes, W. W C., and Peeters, J. C H.: Development of the diatom-Phaeocystis spring bloom in the Dutch coastal zone of the North Sea: the silicon depletion versus the daily irradiance threshold hypothesis, J. Plankton Res., 20, 517–537, 1998. Petersen, W., Petschatnikov, M., Schroeder, F., and Colijn, F.: FerryBox systems for monitoring coastal waters, in: Building the European Capacity in Operational Oceanography: Proc. Third International Conference on EuroGOOS, Elsevier Oceanography Series Publication series, edited by: Dahlien, H., Flemming, N C., Knittis, K., and Petersson, S E., 19, 325–333, 2003. Petersen, W., Wehde, H., Krasemann, H., Colijn, F., and Schroeder, F.: FerryBox and MERIS – Assessment of coastal and shelf sea ecosystems by combining in situ and remotely sensed data, Estuar. Coast. Shelf S., 77, 296–307, 2008. Platt, T., Fuentes-Yaco, C., and Frank, K T.: Marine ecology: spring algal bloom and larval fish survival, Nature, 423, 398–399, 2003. Prins, T., Smaal, A., Pouwer, A., and Dankers, N.: Filtration and resuspension of particulate matter and phytoplankton on an intertidal mussel bed in the Oosterschelde estuary (SW Netherlands), Mar. Ecol. Prog. Ser., 142, 121–134, 1996. Radach, G.: Ecosystem functioning in the German Bight under continental nutrient inputs by rivers, Estuaries Coasts, 15, 477–496, 1992. Ragueneau, O., Quéguiner, B., and Tréguer, P.: Contrast in biological responses to tidally-induced vertical mixing for two macrotidal ecosystems of western Europe, Estuar. Coast. Shelf S., 42, 645–665, 1996. Raven, J A. and Geider, R J.: Temperature and algal growth, New. Phytol., 110, 441–461, 1988. Renz, J., Mengedoht, D., and Hirche, H.: Reproduction, growth and secondary production of \emphPseudocalanus elongatus Boeck (Copepoda, Calanoida) in the southern North Sea, J. Plankton Res., 30, 511–528, 2008. Siegismund, F. and Schrum, C.: Decadal changes in the wind forcing over the North Sea, Climate Res., 18, 39–45, 2001. Skogen, M D. and Moll, A.: Importance of ocean circulation in ecological modeling: An example from the North Sea, J. Marine Sys., 57, 289–300, 2005. Skogen, M D., Söiland, H., and Svendson, E.: Effects of changing nutrient loads to the North Sea, J. Marine Sys., 46, 23–38, 2004. Smetacek, V. and Cloern, J E.: Oceans: On Phytoplankton Trends, Science, 319, 1346, 2008. Soetaert, A. F. H K. and Middelburg, J J.: Present nitrogen and carbon dynamics in the Scheldt estuary using a novel 1-D model, Biogeosciences, 5, 981–1006, 2008. Sommer, U.: Biologische Meereskunde, Springer, 1998. Sommer, U. and Lengfellner, K.: Climate change and the timing, magnitude, and composition of the phytoplankton spring bloom, Glob. Change Biolo., 14, 1199–1208, 2008. Staneva, J., Stanev, E V., Wolff, J O., Badewien, T., Reuter, R., Flemming, B., Bartholomä, and Bolding, K.: Hydrodynamics and sediment dynamics in the German Bight. A focus on observations and numerical modelling in the East Frisian Sea, Cont. Shelf. Res., 29, 302–319, 2009. Stelfox-Widdicombe, C E., Archer, S D., Burkill, P H., and Stefels, J.: Microzooplankton grazing in Phaeocystis and diatom-dominated waters in the southern North Sea in spring, J. Sea Res., 51, 37–51, 2004. Stips, A., Bolding, K., Pohlmann, T., and Burchard, H.: Simulating the temporal and spatial dynamics of the North Sea using the new model GETM (General Estuarine Transport Model), Ocean Dyn., 54, 266–283, 2004. Taylor, A H., Geider, R J., and Gilbert, F. J H.: Seasonal and latitudinal dependencies of phytoplankton carbon-to-chlorophyll \empha ratios: results of a modelling study, Mar. Ecol. Prog. Ser., 152, 51–66, 1997. Tett, P., Heaney, S I., and Droop, M R.: The redfield ratio and phytoplankton growth rate., J. Mar. Biol. Assoc. U.K., 65, 487–504, 1985. Thomas, A., Townsend, D., and Weatherbee, R.: Satellite-measured phytoplankton variability in the Gulf of Maine, Cont. Shelf. Res., 23, 971–989, 2003. Tian, T., Merico, A., Su, J., Staneva, J., Wiltshire, K., and Wirtz, K.: Importance of resuspended sediment dynamics for the phytoplankton spring bloom in a coastal marine ecosystem, J. Sea Res., 62, 214–228, 2009. Tilzer, M M.: The importance of fractional light absorption by photosynthetic pigments for phytoplankton productivity in Lake Constance., Limnol. Oceanogr., 28, 833–846, 1983. Townsend, D., Cammen, L., Holligan, P., Campbell, D., and Pettigrew, N.: Causes and consequences of variability in the timing of spring phytoplankton blooms, Deep-Sea Res. Pt I, 41, 747–765, 1994. van der Zee, C. and Chou, L.: Seasonal cycling of phosphorus in the Southern Bight of the North Sea, Biogeosciences, 2, 27–42, 2005. Verity, P G.: Measurement and simulation of prey uptake by marine planktonic ciliates fed plastidic and aplastidic nanoplankton., Limnol. Oceanogr., 36, 729–750, 1991. Weston, K., Greenwood, N., Fernand, L., Pearce, D J., and Sivyer, D B.: Environmental controls on phytoplankton community composition in the Thames plume, UK, J. Sea Res., 60, 262–270, 2008. Wiltshire, K H. and Manly, B. F J.: The warming trend at Helgoland Roads, North Sea: phytoplankton response, Helgol. Mar. Res., 58, 269–273, 2004. Wiltshire, K H., Malzahn, A M., Wirtz, K W., Greve, W., Janisch, S., Mangelsdorf, P., Manly, B. F J., and Boersma, M.: Resilience of North Sea phytoplankton spring bloom dynamics: An analysis of long-term data at Helgoland Roads, Limnol. Oceanogr., 53, 1294–1302, 2008. Wirtz, K W. and Eckhardt, B.: Effective variables in ecosystem models with an application to phytoplankton succession, Ecol. Modell., 92, 33–53, 1996. Woods, J., Perilli, A., and Barkmann, W.: Stability and predictability of a virtual plankton ecosystem created with an individual-based model, Prog. Oceanogr., 2005.