Biogeosciences www.biogeosciences.net 1726-4170 1726-4189 3 4 2006 10.5194/bg-3-635-2006 http://www.biogeosciences.net/3/635/2006/ http://www.biogeosciences.net/3/635/2006/bg-3-635-2006.html http://www.biogeosciences.net/3/635/2006/bg-3-635-2006.pdf 635 650 2006-12-18 Seasonal dynamics of Pseudocalanus minutus elongatus and Acartia spp. in the southern Baltic Sea (Gdańsk Deep) – numerical simulations L. Dzierzbicka-Głowacka dzierzb@iopan.gda.pl L. Bielecka S. Mudrak Institute of Oceanology, Polish Academy of Sciences Powstańcóy 55, 81-712 Sopot, Poland Institute of Oceanography, University of Gdańsk Al. Marsz. J. Piłsudskiego 46, 81-378 Gdynia, Poland A population dynamics model for copepods is presented, describing the seasonal dynamics of Pseudocalanus minutus elongatus and Acartia spp. in the southern Baltic Sea (Gdańsk Deep). The copepod model was coupled with a one-dimensional physical and biological upper layer model for nutrients (total inorganic nitrogen, phosphate), phytoplankton, microzooplankton, and an early juvenile of herring as a predator. In this model, mesozooplankton (herbivorous copepods) has been introduced as an animal having definite patterns of growth in successive stages, reproduction and mortality. The populations are represented by 6 cohorts in different developmental stages, thus assuming that recruitment of the next generation occurs after a fixed period of adult life. The copepod model links trophic processes and population dynamics, and simulates individual growth within cohorts and the changes in biomass between cohorts. The simulations of annual cycles of copepods contain one complete generation of Pseudocalanus and two generations of Acartia in the whole column water, and indicate the importance of growth in the older stages of 6 cohorts of each species, to arrive at a total population biomass. The peaks of copepods' biomass are larger at the turn of June and July for Pseudocalanus and smaller in July for Acartia, lagging that of phytoplankton by ca. two mouths, due to the growth of cohorts in successive stages and egg production by females. The numerical results show that the investigated species could not be the main factor limiting the spring phytoplankton bloom in the Gdańsk Deep, because the initial development was slow for Acartia and faster for Pseudocalanus, but the main development formed after the bloom, in both cases. The phytoplankton bloom is very important in the diet of the adults of the copepods, but it is not particularly important for the youngest part of new generation (early nauplii). However, the simulated microzooplankton biomass was enough high to conclude, in our opinion, that, in this case, it was a major cause in limiting phytoplankton bloom. The model presented here is a next step in understanding how the population dynamics of a dominant species in the southern Baltic Sea interact with the environment. Ambler, J. W.: Seasonal factors affecting egg production and viability of eggs of \it Acartia tonsa Dana, from East Lagoon, Galveston, Texas, Estuar. cstl. Shelf Sci., 20, 743–760, 1985. Bautista, B., Harris, R. P., Rodriguez, V., and Guerrero, F.: Temporal variability in copepod fecundity during two different spring bloom periods in coastal waters off Plymouth (SW England), J. Plankton Res., 16, 1367–1377, 1994. Beckman, B. C. and Peterson, W. T.: Egg production of \it Acartia tonsa Dana in Long Island Sound, J. Plankton Res., 8, 917–925, 1986. Berggreen, U., Hansen, B., and Kiørboe, T.: Food size spectra, ingestion and growth of the copepod \it Acartia tonsa during development: implications for determination of copepod production, Marine Biology, 99, 341–352, 1988. Billen, G., Lancelot, C., and Maybeck, M.: N, P and Si retention along the aquatic continuum form land to ocean, in: Ocean Margin Processes in Global Change, edited by: Mantoura, R. F. C., Martin, J. M., and Wollast, R., Phys., Chem., Earth Sci. Res., Report 9, 19–44, Wiley and Sons, New York, 1991. Carlotti, F. and Sciandra, A.: Population dynamics model of \it Euterpina acutifrons (Copepoda: Harpacticoida) couplong individual growth and larval development, Mar. Ecol. Prog. Ser., 56, 225–242, 1989. Carlotti, F. and Nival, P.: Model of copepod growth and development: moulting and mortality in relation to physiological processes during an individual moult cycle, Mar. Ecol. Prog. Ser., 84, 219–233, 1992. Carlotti, F. and Radach, G.: Seasonal dynamics of phytoplankton and \it Calanus finmarchicus in the North Sea as revealed by a~coupled one-dimensional model, Limnol. Oceanogr., 41(3), 522–539, 1996. Carlotti, F. and Wolf, K. U.: A Lagrangian ensemble model of \it Calanus finmarchicus coupled with a~1-D ecosystem model, Fish. Oceanogr., 7(3/4), 191–204, 1998. Checkley, D. M.: Food limitation of egg production by a marine planktonic copepod in the sea off southern California, Limnol. Oceanogr., 25, 991–998, 1980. Christoffersen, K. and Jespersen, A. M.: Gut evacuation rates and ingestion rates of \it Eudiaptomus graciloides measured by means of the gut fluorescence method, J. Plankton Res., 8, 973–983, 1986. Ciszewski, P. and Witek, Z.: Production of older stages of copepods \it Acartia bifilosa Giesb. and \it Pseudocalanus elongatus Boeck in Gda\'nsk Bay, Pol. arch. Hydrobiol., 24, 449–459, 1977. Corkett, C. J. and McLaren, I. A.: The biology of \it Pseudocalanus, Ad. Mar. Biol., 15, 1–231, 1978. Dagg, M. J. and Wyman, K. D.: Natural ingestion rates of the copepods \it Neocalanus plumchrus and \it N. cristatus, calculated from gut contents, Mar. Ecol. Prog. Ser., 13, 37–46, 1983. Dam, H. G. and Peterson, W. T.: The effect of temperature on the gut clearance rate constant of planktonic copepods, J. exp. mar. Biol. Ecol., 123, 1–14, 1988. Durbin, E. G., Durbin, A. G., Smayda, T. J., and Verity, P. G.: Food limitation of production by adult \it Acartia tonsa in Narragansett Bay, Rhode Island, Limnol. Oceanogr., 28, 1199–1213, 1983. Durbin, E. G. and Durbin, A. G.: Effects of temperature and food abundance on grazing and short-term weight change in the marine copepod \it Acartia hudsonica, Limnol. Oceanogr., 37, 361–378, 1992. Dzierzbicka-G\lowacka, L.: Numerical analysis of the influence of the grazing on the two-dimensional distribution function of the phytoplankton concentration in a stratified sea, Oceanologia, 36(2), 155–173, 1994. Dzierzbicka-G\lowacka, L.: Growth and development of copepodite stages of \it Pseudocalanus~spp, J. Plankton Res., 26, 49–60, 2004a. Dzierzbicka-G\lowacka, L.: The dependence of body weight in copepodite stages of \it Pseudocalanus~spp on variations of ambient temperature and food concentration, Oceanologia, 46, 45–63, 2004b. Dzierzbicka-G\lowacka, L.: A numerical investigation of phytoplankton and \it Pseudocalanus elongatus dynamics in the spring bloom time in the Gdansk Gulf, J. Mar. Sys., 53, 19–36, 2005a. Dzierzbicka-G\lowacka, L.: Modelling the seasonal dynamics of marine plankton in southern Baltic Sea. Part 1. A Coupled Ecosystem Model, Oceanologia, 47(4), 591–619, 2005b. Dzierzbicka-G/lowacka, L.: Equivalence of rates of growth and egg production of \it Pseudocalanus, Ocean. Hydrobiol. Stud., 34(4), 19–32, 2005c. Dzierzbicka-G\lowacka, L.: Encounter rate in zooplankton, Pol. J. Environ. Stud., 15(2), 243–257, 2006. Fennel, W.: Modeling of copepods with links to circulation model, J. Plankton Res., 23, 1217–1232, 2001. Fennel, W. and Neumann, T.: Variability of copepods as seen in a coupled physical biological model of the Baltic Sea, ICEM Marine Science Symposia, 219, 208–219, 2003. Fey, D. P.: Differences in temperature conditions and somatic growth rate of larval and early juvenile spring-spawned herring from the Vistula Lagoon, Baltic Sea manifested in the otolith to fish size relationship, J. Fish Biol., 58, 1257–1273, 2001. Fransz, H., Moammerts, J., and Radach, G.: Ecological modelling of the North Sea, Nether. J. Sea Res. 28, 67–140, 1991. Frost, B. W.: A taxonomy of the marine calanoid copepod genus \it Pseudocalanus, Can. J. Zool., 67, 525–551, 1989. Hay, S. J.: Egg production and secondary production of common North Sea copepods: field estimates ith regional and seasonal comparisons, ICES J. Mar. Sci., 52, 315–327, 1995. Hernroth, L.: Recommendations on methods for marine biological studies in the Baltic Sea. Mesozooplankton biomass assessment, The Baltic Marine Biologists, 10, 1–32, 1985. Kiørboe, T., Møhlenberg, F. and Nicolajsen, H.: Ingestion rate and gut clearance in the planktonic copepod \it Centropages hamatus (lilljeborg) in relation to food concentration and temperature, Ophelia, 21, 181–194, 1982. Kiørboe, T., Møhlenberg, F., and Hamburger, K.: Bioenergetics of the planktonic copepod \it Acartia tonsa: relation between feeding, egg production and respiration, and composition of specific dynamic action, Mar. Ecol. Prog. Ser., 26, 85–97, 1985. Kiørboe, T. and Johansen, K.: Studies of a larval herring (\it Clupea harengus L.) patch in the Buchan area. IV Zooplankton distribution and productivity in relation to hydrographic features, Dana, 6, 37–51, 1986. Klein Breteler, W. C. M., Gonzales, S. R., and Schogt, N.: Development of Pseudocalanus elongatus (Copepoda, Calanoida) cultured at different temperature and food conditions, Mar. Ecol. Progr. Ser., 119, 99–110, 1995. Kleppel, G. S., Holliday, D. V., and Pieper, R. E.: Trophic interactions between copepods and micoplankton: a question about the role of diatoms, Limnol. Oceanogr., 36, 172–178, 1991. Kleppel, G. S.: Environmental regulation of feeding and egg production by \it Acartia tonsa off southern California, Mar. Biol., 112, 57–65, 1992. Koski, M., Breteler, W. K. and Schogt, N.: Effect of food quality on rate growth and development of the pelagic copepod \it Pseudocalanus elongatus (Copepoda, Calanoida), Mar. Ecol. Prog. Ser., 170, 169–187, 1998. Kremer, J. N. and Nixon, S. W.: A coastal marine ecosystem. Simulation and analysis, Ecological studies 24, Springer-Verlag, Heidelberg, 1978. Landry, M. R.: The development of marine calanoids with comment on the isochronal rule, Limnol. Oceanogr., 28, 614–624, 1983. Last, J. M.: The food of four species of pleuronectiform larvae in the eastern English Channel and southern North Sea, Mar. Biol., 45, 359–368, 1978a. Last, J. M.: The food of three species of gadoid larvae in the eastern English Channel and southern North Sea, Mar. Biol., 45, 377–386, 1978b. Last, J. M.: The food of twenty species of fish larvae in the west-central North Sea, Fisheries Research Technical report, Lowestoft, 60, pp 44, 1980. Maritime Branch Materials Institute of Meteorology and Water Management, Environmental conditions in the Polish zone of the southern Baltic Sea during 1999, 2000. Margo\'nski, P.: The abundance, growth rate and mortality of the early life stages of herring (\it Clupea harengus) and smelt (\it Osmerus eperlanus) in the Vistula Lagoon (southern Baltic Sea) during 1998–1999, International Council for the Exploration of the Sea ICES CM 2000/N:21, 2000. McLaren, I. A.: Generation lengths of some temperate marine copepods: estimation, production and implications, J. Fish Res. Bd. Can., 345, 1330–1342, 1978. McLaren, I. A., Corkett, C. J., and Zillioux, E. J.: Temperature adaptations of copepod eggs from the arctic to the tropics, Biol. Bull., 137, 486–493, 1969. McLaren, I. A. and Leonard, A.: Assessing the equivalence of growth and egg production of copepods, ICES J. Mar. Sci., 52, 397–408, 1995. McLaren, I. A., Sévigny, J. M., and Corkett, C. J.: Temperature-dependent development in \it Pseudocalanus species, Can. J. Zool., 67, 559–564, 1989. Miller, T. J., Crowder, L. B., Rice, J. A., and Marshall, E. A.: Larval size and recruitment mechanisms in fishes: toward a~conceptual framework, Can. J. Fish. Aquat. Sci., 45, 1657–1670, 1988. Miller, C. A. and Landry, M. R.: Ingestion-independent rates of ammonium excretion by the copepod \it Calanus pacificus, Mar. Biol., 78, 263–270, 1984. Moll, A. and Stegert, C.: Modelling \it Pseudocalanus elongatus stage-structured population dynamics embedded in a water column ecosystem model for the northern North Sea, J. Marine Sys., 64, 35–46, 2007. Mudrak, S.: Short- and long-term variability of zooplankton in coastal Baltic water: using the Gulf of Gda\'nsk as an example, PhD thesis, Gda\'nk Univ., Gdynia, (in Polish), 323 pp., 2004. Mudrak, S., Bielecka, L., and /Zmijewska, M. I.: Diel vertical migrations of copepoda from Gdansk Deep (Baltic sea), Abstracts, 39th European Marine Biology Symposium, Genoa, 2004. Paffenhöfer, G. A.: Grazing and ingestion rates of nauplii, copepodids and adults of the marine planktonic copepod \it Calanus helgolandicus, Mar. Biol., 11, 286–298, 1971. Paffenhöfer, G.-A. and Harris, R. P.: Feeding, growth and reproduction of the marine planktonic copepod \it Pseudocalanus elongatus Boeck, J. Mar. Biol. Ass. UK, 56, 327–344, 1976. Peterson, W. T.: Rates of egg production by the copepod \it Calanus marshallae in the laboratoy and in the sea off Oregon, USA, Mar. Ecol. Prog. Ser., 47, 229–237, 1988. Piontkovskii, S. A. and Petipa, T. S.: Quantitative description of the behavior of copepod \it Acartia clausi during feeding on algae, Soviet. J. Mar. Biol., 2, 40–46, 1976. Radach, G., Carlotti, F., and Spangenberg, A.: Variability of zooplankton population dynamics caused by annual weather variability, Fish. Oceanogr., 7(3/4), 272–281, 1998. Renk, H.: Primary production in the southern Baltic, Sea Fisheries Institute, Gdynia, Poland, 2000. Renk, H., Ochocki, S., and Pytel, H.: Short-term fluctuations of primary production and chlorophyll-a concentration in the Gda\'nsk Deep, Pol. Ecol. Stud., 9(3), 341–359, 1983. Riley, G. A.: Factors controlling phytoplankton populations on Georges Bank, J. Mar. Res., 6, 54–73, 1946. Riley, G. A.: A mathematical model of regional variations in plankton, Limnol. Oceanogr., 10, 202–215, 1965. Roman, M. R.: Utilization of detritus by the copepod \it Acartia tonsa, Limnol. Oceanogr., 29, 949–959, 1984. Rothschild, B. J.: Biodynamics of the sea: the ecology of high dimensionality systems, in: Toward a theory of biological – physical interactions in the world ocean, edited by: Rothschild, B. J., Kluwer, Boston, 527–548, 1988. Runge, J. A.: Egg production of the marine planktonic copepod, \it Calanus pacificus Brodsky: laboratory observations, J. Exp. Mar. Biol. Ecol., 74, 53–66, 1984. Runge, J. A.: Relationship of egg production of \it Calanus pacificus to seasonal changes in phytoplankton availability in Puget Sound, Washington, Limnol. Oceanogr., 30, 382–396, 1985. Sciandra, A.: Study and modelling of development of \it Euterpina acutifrons (Copepoda, Harpacticoida), J. Plankton Res., 8, 1149–1162, 1986. Schmidt, K., Kähler, P., and Bodungen, B.: Copepod egg production rates in the Pomeranian Bay (southern Baltic Sea) as a function of phytoplankton abundance and taxonomic composition, Mar. Ecol. Prog. Ser., 174, 183–195, 1998. Sekiguchi, H., McLaren, I. A., and Corkett, C. J.: Relationship between growth rate and egg production in the copepod \it Acartia clausi Hudsonica, Mar. Biol., 58, 133–138, 1980. Steele, J. H.: The Structure of Marine Ecosystems, Harvard University Press, Cambridge, 1974. Steele, J. and Henderson, E. W.: Simulation of vertical structure in a planktonic ecosystem, Scott. Fish. Res. Rep., 5, 1–27, 1976. Steele, J. H. and Mullin, M. M.: Zooplankton dynamics, in: The sea Vol. 6, edited by: Goldberg, E. D., McCave, I. N., O'Brien, J. J., and Steele, J. H., Interscience Publ. New York, London, Sydney, Toronto, 857–887, 1977. Stoecker, D. K. and Egloff, D. A.: Predation by \it Acartia tonsa Dana on planktonic ciliates and rotifers, J. Exp. Mar. Biol. Ecol., 110, 53–68, 1987. Thompson, B. M.: The biology of \it Pseudocalanus elongatus (Boeck), PhD thesis, University of East Anglia, Norwich, 1976. Thompson, B. M.: Growth and development of \it Pseudocalanus elongatus and \it Calanus~sp in the laboratory, J. Mar. Biol. Ass. UK., 62, 359–372, 1982. Thompson, A. M., Durbin, E. G., and Durbin, A. G.: Seasonal changes in maximum ingestion rate of \it Acartia tonsa in Narragansett Bay, Rhode Island, USA, Mar. Ecol. Prog. Ser., 108, 91–105, 1994. Turner, J. T. and Tester, P. A.: Zooplankton feeding ecology: nonselective grazing by the copepods \it Acartia tonsa Dana, \it Centropages velificatus De Oliveira, and \it Eucalanus pileatus giesbrecht in the plume of the Mississippi River, J. Exp. Mar. Biol. Ecol., 126, 21–43, 1989. Witek, Z.: Structure and function of marine ecosystem in the Gda/nsk Basin on the basis of studies performed in 1987, Scientific Committee on Oceanic Research, 63, 1–123, 1993. Witek, Z.: Biological production and its utilization within a~marine ecosystem in the western Gdansk basin, Sea Fischeries Institute, Gdynia, Poland, 1995. Wlodarczyk, E., Durbin, A. G., and Durbin, E. G.: Effect of temperature on lower feeding thresholds, gut evacuation rate, and diel feeding behavior in the copepod \it Acartia hudsonica, Mar. Ecol. Prog. Ser., 85, 93–106, 1992. Wroblewski, J. S.: Formulation of growth and mortality of larvae of northern anchovy in a turbulent feeding environment, Mar. Ecol. Prog. Ser., 20, 13–22, 1984. Wroblewski, J. S. and Richman, J. G.: The non-linear response of plankton to wind mixing events – implications for survival of larval northern anchovy, J. Plankton Res., 9, 103–123, 1987. Verity, P. G. and Smayda, T. J.: Nutritional value of \it Phaeocystis pouchetii (Prymnesiophyceae) and other phytoplankton for \it Acartia spp (Copepoda): ingestion, egg production, and growth of nauplii, Mar. Biol., 100, 161–171, 1989. Vidal, J.: Physioecology of Zooplancton. I. Effects of phytoplankton concentration, temperature, and body size on the growth rate of \it Calanus pacificus and \it Pseudocalanus~sp., Mar. Biol., 56, 111–134, 1980a. Vidal, J.: Physioecology of Zooplancton. II. Effects of phytoplankton concentration, temperature, and body size on the development and molting rates of \it Calanus pacificus and \it Pseudocalanus~sp., Mar. Biol., 56, 135–146, 1980b. Vinogradov, M. E. and Shushkina, E. A.: Functioning of pelagic plankton communities in the ocean, Nauka Moskwa (in Russian), 1987. Za\llachowski, W., Szypu/la, J., Krzykawski, S., and Krzykawska, I.: Feeding of some commercial fishes in the southern region of the Baltic Sea in 1971 and 1972, Pol. Arch. Hydrobiol., 22, 429–448, 1975.