Estuaries and Coasts

, Volume 37, Issue 4, pp 911–924 | Cite as

Vertical Carbon Flux of Biogenic Matter in a Coastal Area of the Aegean Sea: The Importance of Appendicularians

  • S. Zervoudaki
  • C. Frangoulis
  • C. Svensen
  • E. D. Christou
  • E. Tragou
  • E. G. Arashkevich
  • T. N. Ratkova
  • Ι. Varkitzi
  • E. Krasakopoulou
  • K. Pagou
Article
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Abstract

This work focuses on the direct measurement of the vertical flux of appendicularian houses in order to assess their importance as a component of vertical carbon flux in coastal areas. For this purpose, arrays of cylindrical sediment traps were deployed for 5 to 8 days at two depths in a coastal area of the northern Aegean Sea (inner Thermaikos Gulf) during spring. The data support the contention that resuspension was minimal. Fecal pellet (FP) production and grazing experiments with the dominant copepods (Acartia clausi) were conducted to provide additional information on the potential FP contribution to the total carbon flux. The magnitude of the vertical flux of particulate organic carbon (POC) ranged between 310 and 724 mg C m−2 day−1. The proportion of phytoplankton carbon in the POC vertical flux was up to 45 %. The contribution of zooplankton FPs to the total carbon never exceeded 5 %. On the contrary, appendicularian houses were an essential component of the biogenic carbon flux contributing up to 55.3 % of the total vertical carbon flux. Consequently, both phytoplankton and appendicularian houses contributed equally to the biogenic carbon flux exceeding 80 % of the total sinking POC. Taking into account the sinking speed of the particles and the environment in the area, all this carbon probably reaches the seafloor, thus indicating a strong pelagic–benthic coupling.

Keywords

Vertical flux Appendicularians Coastal area Northern Aegean Sea 
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Notes

Acknowledgments

This study was supported by the European Commission (Research Directorate General–Environment Program–Marine Ecosystems) through the FATE project “Transfer and Fate of Harmful Algal Bloom (HAB) Toxins in European Marine Waters” (contract number EVK3-2001-00055, contract holder E. Granéli) as part of the EC EUROHAB cluster. We gratefully acknowledge G. Papas, C. Charalambous, and C. Wexels Riser for their help in the field, Y. N. Krestenitis (Aristotle University of Thessaloniki) and A. Pavlidou (HCMR) for providing temperature, salinity, and nutrient data, as well as Prof. V. Galanopoulos and Th. Kanelopoulos for their help in the SEM observations. Finally the two anonymous reviewers for their valuable comments on the manuscript.

References

  1. Acuña, J.L., D. Deibel, and C.C. Morris. 1996. Particle capture mechanism of the pelagic tunicate Oikopleura vanhoeffeni. Limnology and Oceanography 41(8): 1800–1814.CrossRefGoogle Scholar
  2. Agawin, N.S.R., and C.M. Duarte. 2002. Evidence of direct particle trapping by a tropical seagrass meadow. Estuaries 25: 1205–1209.CrossRefGoogle Scholar
  3. Alldredge, A.L. 1972. Abandoned larvacean houses: A unique food source in the pelagic environment. Science 177: 885–887.CrossRefGoogle Scholar
  4. Alldredge, A.L. 2005. The contribution of discarded appendicularian houses to the flux of particulate organic carbon from oceanic surface waters. In Response of marine ecosystems to global change: Ecological impact of appendicularians, ed. G. Gorsky, M.J. Youngbluth, and D. Deibel, 309–326. Paris: GBScientific Publisher.Google Scholar
  5. Alldredge, A.L., and M. Silver. 1988. Characteristics, dynamics and significance of marine snow. Progress in Oceanography 20: 41–82.CrossRefGoogle Scholar
  6. Balopoulos, E.T.H., and N.C.H. Friligos. 1994. Water circulation and eutrophication in the northwestern Aegean Sea. Thermaikos Gulf Toxicological and Environmental Chemistry 41(3): 155–167.CrossRefGoogle Scholar
  7. Båmstedt, U., D.J. Gifford, X. Irigoien, A. Atkinson, and M. Roman. 2000. Feeding. In ICES zooplankton methodology manual, ed. R. Harris, P. Wiebe, J. Lenz, H.R. Skjoldal, and M. Huntley, 297–399. London: Academic.CrossRefGoogle Scholar
  8. Barron, C., J.J. Middelburg, and C.M. Duarte. 2006. Phytoplankton trapped within seagrass (Posidonia oceanica) sediments are a nitrogen source: an in situ isotope labeling experiment. Limnology and Oceanography 51: 1648–1653.Google Scholar
  9. Bauerfeind, E., C. Garrity, M. Krumbholz, R.O. Ramseier, and M. Voß. 1997. Seasonal variability of sediment trap collections in the northeast water polynya. Part 2. Biochemical and microscopic composition of sedimenting matter. Journal of Marine Systems 10: 371–389.CrossRefGoogle Scholar
  10. Bedo, A.W., J.L. Acuna, D. Robins, et al. 1993. Grazing in the micron and the submicron particle-size range—The case of Oikopleura dioica (Appendicularia). Bulletin of Marine Science 53(1): 2–14.Google Scholar
  11. Bochdansky, A.B., and G.J. Herndl. 1992. Ecology of amorphous aggregations (marine snow) in the Northern Adriatic Sea. V. Role of fecal pellets in marine snow. Marine Ecology Progress Series 89(2–3): 297–303.CrossRefGoogle Scholar
  12. Bodungen, B.V., K.V. Bröckel, V. Smetacek, and B. Zeitzschel. 1981. Growth and sedimentation of the phytoplankton spring bloom in the Bornholm Sea (Baltic Sea). Kieler Meeresforsch Sonderh 5: 49–60.Google Scholar
  13. Buesseler, K.O. 1991. Do upper-ocean sediment traps provide an accurate record of particle flux? Nature 353: 420–423.CrossRefGoogle Scholar
  14. Buesseler, K.O., A.N. Antia, M. Chen, S.W. Fowler, W.D. Gardner, O. Gustafsson, K. Harada, A.F. Michaels, M. Rutgers van der Loeff, M. Sarin, D.K. Steinberg, and T. Trull. 2007. An assessment of the use of sediment traps for estimating upper ocean particle fluxes. Journal of Marine Research 65: 345–416.CrossRefGoogle Scholar
  15. Calbet, A., S. Garrido, E. Saiz, M. Alcaraz, and C.M. Duarte. 2001. Annual zooplankton succession in coastal NW Mediterranean waters: The importance of the smaller size fractions. Journal of Plankton Research 23(3): 319–331.CrossRefGoogle Scholar
  16. Christou, E.D., and G.C. Verriopoulos. 1993. Length, weight and condition factor of Acartia clausi (Copepoda) in the Eastern Mediterranean. Journal of Marine Biological Association UK 73(2): 343–353.CrossRefGoogle Scholar
  17. Christou, E.D., K. Pagou, S., Christianidis and Papathanassiou, E. 1995. Temporal and spatial variability of plankton communities in a shallow embayment of the eastern Mediterranean. In Biology and ecology of shallow coastal waters, ed. A. Eleftheriou, 3–10. Denmark: Olsen & OlsenGoogle Scholar
  18. Coppola, L., M. Roy-Barman, P. Wassmann, S. Mulsow, and C. Jeandel. 2002. Calibration of sediment traps and particulate organic carbon export using Th-234 in the Barents Sea. Marine Chemistry 80: 11–26.CrossRefGoogle Scholar
  19. Cutter, G.A. and J. Radford-Knoery. 1991. Determination of carbon, nitrogen, sulfur, and inorganic sulfur species in marine particles. In: Marine Particles: Analysis and Characterization, eds. D. Hurd and D. Spencer pp. 57–63. Washington: American Geophysical Union.Google Scholar
  20. Dauby, P., A.J. Bale, N. Bloomer, C. Canon, R.D. Ling, A. Norro, J.E. Robertson, A. Simon, J.M. Théate, A.J. Watson, and M. Frankignoulle. 1995. Particle fluxes over a Mediterranean seagrass bed: A one year case study. Marine Ecology Progress Series 126: 233–246.CrossRefGoogle Scholar
  21. Deibel, D., and C.V.L. Powell. 1987. Ultrastructure of the pharyngeal filter of the appendicularian Oikopleura vanhoeffeni—Implications for particle-size selection and fluid-mechanics. Marine Ecology 35(3): 243–250.CrossRefGoogle Scholar
  22. Elder, D.L., and S.W. Fowler. 1977. Polychlorinated biphenyls: Penetration into the deep ocean by zooplankton fecal pellet transport. Science 197: 459–461.CrossRefGoogle Scholar
  23. Fernández de Puelles, M.L., J. Valencia, and L. Vicente. 2004. Zooplankton variability and climatic anomalies from 1994 to 2001 in the Balearic Sea (Western Mediterranean). ICES Journal of Marine Science 61(4): 492–500.CrossRefGoogle Scholar
  24. Fowler, S.W., and G.A. Knauer. 1986. Role of large particles in the transport of elements and organic-compounds through the oceanic water column. Progress in Oceanography 16(3): 147–194.CrossRefGoogle Scholar
  25. Frangoulis, C., E.D. Christou, and J.H. Hecq. 2005. Comparison of marine copepod outfluxes: Nature, rate, fate and role in the carbon, and nitrogen cycles. Advances in Marine Biology 47: 251–307.Google Scholar
  26. Frangoulis, C., S. Psarra, V. Zervakis, T. Meador, P. Mara, A. Gogou, S. Zervoudaki, A. Giannakourou, P. Pitta, A. Lagaria, E. Krasakopoulou, and I. Siokou-Frangou. 2010. Connecting export fluxes to plankton food-web efficiency in the Black Sea waters inflowing into the Mediterranean Sea. Journal of Plankton Research 32(8): 1203–1216.CrossRefGoogle Scholar
  27. Frangoulis, C., N. Skliris, G. Lepoint, K. ElKalay, A. Goffart, J.K. Pinnegar, and J.H. Hecq. 2011. Importance of copepod carcasses versus faecal pellets in the upper water column of an oligotrophic area. Estuaries Coastal and Shelf Science 92: 456–463.CrossRefGoogle Scholar
  28. Friligos, N., J.C. Kondylakis, R. Psyllidou-Giouranovits, and E. Georgakopoulou-Gregoriadou. 1997. Eutrophication and phytoplankton abundance in the Thermaikos Gulf. Greece Fresenius Environmental Bulletin 6(1–2): 27–31.Google Scholar
  29. Frost, B.W. 1972. Effect of size and concentration of food particles on the feeding behaviour of the marine planktonic copepod Calanus pacificus. Limnology and Oceanography 17: 805–815.CrossRefGoogle Scholar
  30. Gacia, E., and C.M. Duarte. 2001. Sediment retention by a Mediterranean Posidonia oceanica meadow: The balance between deposition and resuspension. Estuarine, Coastal and Shelf Science 52: 505–514.CrossRefGoogle Scholar
  31. Ganoulis, J.G. 1991. Water quality assessment and protection measures of a semi-enclosed coastal area: the Bay of Thermaikos (NE Mediterranean Sea). Marine Pollution Bulletin 23: 83–87.CrossRefGoogle Scholar
  32. Gardner, W.D., M.J. Richardson, C.A. Carlson, D. Hansell, and A.V. Mishonov. 2003. Determining true particulate organic carbon: Bottles, pumps and methodologies. Deep Sea Research II Topical Studies Oceanography 50(3–4): 655–674.CrossRefGoogle Scholar
  33. González, H.E., and V. Smetacek. 1994. The possible role of the cyclopoid copepod Oithona in retarding vertical flux of zooplankton faecal material. Marine Ecology Progress Series 113: 233–246.Google Scholar
  34. González, H.E., S. González, and G.J. Brummer. 1994. Short-term sedimentation pattern of zooplankton, faeces and microplankton at a permanent station in the Bjørnafjorden (Norway) during April–May 1992. Marine Ecology Progress Series 105: 31–45.CrossRefGoogle Scholar
  35. Gotsis-Skretas, O., and N. Friligos. 1990. Eutrophication and phytoplankton ecology in the Therrnaikos Gulf. Thalassographica 13: 1–12.Google Scholar
  36. Goutx, M., A. Momzikoff, L. Striby, et al. 2000. High-frequency fluxes of labile compounds in the central Ligurian Sea, Northwestern Mediterranean. Deep Sea Research 47: 533–556.CrossRefGoogle Scholar
  37. Hansen, J.L.S., T. Kiørboe, and A.L. Alldredge. 1996. Marine snow derived from abandoned larvacean houses: Sinking rates, particle content and mechanisms of aggregate formation. Marine Ecology Progress Series 141: 202–215.CrossRefGoogle Scholar
  38. Hasselmann, K., D.B. Ross, P. Müller, and W. Sell. 1976. A parametric wave prediction model. Journal of Physical Oceanography 6(2): 200–228.CrossRefGoogle Scholar
  39. HCMR. 2007. Monitoring the quality of the marine environment in the Gulf of Thessaloniki from 1984–2006. Hellenic Centre for Marine Research, Final Technical Report, Athens, 249 pp (in Greek).Google Scholar
  40. Holm-Hansen, O., C.J. Lorenzen, R.N. Hormes, and J.D.H. Strickland. 1965. Fluorometric determination of chlorophyll. Journal de Conseil pour l'Exploration de la Mer 30: 3–15.CrossRefGoogle Scholar
  41. Hyder, P., J.H. Simpson, S. Christopoulos, and Y. Krestenitis. 2002. The seasonal cycles of stratification and circulation in the Thermaikos Gulf region of freshwater influence (ROFI). North-west Aegean Continental Shelf Research 22(17): 2573–2597.CrossRefGoogle Scholar
  42. Jagger, R.A., W.J. Kimmererer, and G.P. Jenkins. 1988. Food of the cladoceran Podon intermedius in a marine embayment. Marine Ecology Progress Series 43: 245–250.CrossRefGoogle Scholar
  43. Karageorgis, A.P., M.S. Skourtos, V. Kapsimalis, A.D. Kontogianni, N.Th. Skoulikidis, K. Pagou, N.P. Nikolaidis, P. Drakopoulou, B. Zanou, H. Karamanos, Z. Levkov, and Ch. Anagnostou. 2005. An integrated approach to watershed management within the DPSIR framework: Axios River catchment and Thermaikos Gulf. Regional Environmental Change 5: 138–160.Google Scholar
  44. Kiørboe, T., J. Hansen, A.L. Alldredge, G. Jackson, U. Passow, H. Dam, D. Drapeau, A. Waite, and C. Garcia. 1996. Sedimentation of phytoplankton during a spring diatom bloom: Rates and mechanisms. Journal of Marine Research 54: 1123–1148.CrossRefGoogle Scholar
  45. Komar, P.D., A.P. Morse, L.F. Small, and S.W. Fowler. 1981. An analysis of sinking rates of natural copepod and euphausiid fecal pellets. Limnology and Oceanography 26: 172–180.CrossRefGoogle Scholar
  46. Kombiadou, K., and Y.N. Krestenitis. 2011. Simulating the fate of mechanically eroded masses in the Thermaikos Gulf. Continental Shelf Research 31: 817–831.CrossRefGoogle Scholar
  47. Kontoyiannis, H., Kourafalou, V.H., and Papadopoulos, V. 2003. Seasonal characteristics of the hydrology and circulation in the northwest Aegean Sea (eastern Mediterranean): Observations and modeling. Journal of Geophysical Research 108(C9). doi: 10.1029/2001JC001132.
  48. Koroleff, F. 1970. Revised version of “Direct determination of ammonia in natural waters as indophenol blue”. ICES information on techniques and methods for sea water analysis. Interlaboratory Report 3: 19–22.Google Scholar
  49. Kourafalou, V.H., Y.G. Savvidis, Y.N. Krestenitis, and Ch.G. Koutitas. 2004. Modelling studies on the processes that influence matter transfer on the Gulf of Thermaikos (NW Aegean Sea). Continental Shelf Research 24: 203–222.Google Scholar
  50. Lane, P.V.Z., S.L. Smith, J.L. Urban, and P.E. Biscaye. 1994. Carbon flux and recycling associated with zooplanktonic fecal pellets on the shelf of the Middle Atlantic Bight. Deep Seas Research II 41: 437–457.CrossRefGoogle Scholar
  51. Legendre, L., and F. Rassoulzadegan. 1996. Food-web mediated export of biogenic carbon in oceans: Hydrodynamic control. Marine Ecology Progress Series 145: 179–193.CrossRefGoogle Scholar
  52. Licandro, P., and F. Ibanez. 2000. Changes of zooplankton communities in the Gulf of Tigullio (Ligurian Sea, Western Mediterranean) from 1985 to 1995. Influence of hydroclimatic factors. Journal of Plankton Research 22: 2225–2253.CrossRefGoogle Scholar
  53. Lykousis, V., and G. Chronis. 1989. Mechanisms of sediment transport and deposition—Sediment sequences and accumulation during the holocene on the Thermaikos plateau, the continental-slope, and basin (Sporadhes Basin), northwestern Aegean Sea, Greece. Marine Geology 87(1): 15–26.CrossRefGoogle Scholar
  54. Lykousis, V., and M. Collins. 2005. Impact of Natural and Trawling Events on Resuspension, dispersion and fate of POLlutants (INTERPOL). Continental Shelf Research 25(19–20): 2309–2314.CrossRefGoogle Scholar
  55. Maar, M., T.G. Nielsen, K. Richardson, U. Christaki, O.S. Hansen, S. Zervoudaki, and E.D. Christou. 2002. Spatial and temporal variability of food web structure during the spring bloom in the Skagerrak. Marine Ecology Progress Series 239: 11–29.CrossRefGoogle Scholar
  56. Maar, M., T.G. Nielsen, S. Gooding, K. Tönnesson, P. Tiselius, S. Zervoudaki, E.D. Christou, A. Sell, and K. Richardson. 2004. Trophodynamic function of copepods, appendicularians and protozooplankton in the late summer zooplankton community in the Skagerrak. Marine Biology 144: 917–933.CrossRefGoogle Scholar
  57. Marty, J.C., E. Nicolas, J.C. Miquel, and S.W. Fowler. 1994. Particulate fluxes of organic compounds and their relationship to zooplankton faecal pellets in NW Mediterranean Sea. Marine Chemistry 46: 387–405.CrossRefGoogle Scholar
  58. Menden-Deuer, S., and E.J. Lessard. 2000. Carbon to volume relationships for dinoflagellates, diatoms, and other protist plankton. Limnology and Oceanography 45: 569–579.CrossRefGoogle Scholar
  59. Milos, C., and A. Malej. 2005. Gelatinous zooplankton assemblages in temperate coastal waters—Seasonal variations (Gulf of Trieste, Adriatic Sea). Annales Series History Natural 15: 11–20.Google Scholar
  60. Mochioka, N., and M. Iwamizu. 1996. Diet of anguilloid larvae: Leptocephali feed selectively on larvacean houses and faecal pellets. Marine Biology 125: 447–452.Google Scholar
  61. Moncheva, S., O. Gotsis-Skretas, K. Pagou, and A. Krastev. 2001. Phytoplankton blooms in Black Sea and Mediterranean ecosystems subjected to anthropogenic eutrophication: Similarities and differences. Estuarine, Coastal and Shelf Science 53: 281–295.CrossRefGoogle Scholar
  62. Moran, S.B., M.A. Charette, S.M. Pike, and C.A. Wicklund. 1999. Differences in seawater particulate organic carbon concentration in samples collected using small- and large-volume methods: The importance of DOC adsorption to the filter blank. Marine Chemistry 67(1–2): 33–42.CrossRefGoogle Scholar
  63. Mullin, N.J.B., and J.P. Riley. 1955. The colorimetric determination of silicate with special reference to sea and natural waters. Analytica Chimica Acta 12: 162–176.CrossRefGoogle Scholar
  64. Murphy, J., and J.P. Riley. 1962. A modified solution method for determination of phosphate in natural waters. Analytica Chimica Acta 27: 31–36.CrossRefGoogle Scholar
  65. Pagou, K. 2005. Eutrophication in Hellenic coastal areas. In State of the Hellenic marine environment, ed. E. Papathanassiou and A. Zenetos, 311–317. Athens: HCMR Publications. ISBN 960-86651-8-3.Google Scholar
  66. Pagou, K., Assimakopoulou, G., Krasakopoulou, E., and Pavlidou, A. 2000. Nutrient input, fluxes and cycling in relation to the biological production in Mediterranean ecosystems influenced by river discharges: Thermaikos Gulf. In Dynamics of matter transfer and biogeochemical cycles: Their modeling in coastal systems of the Mediterranean Sea, ed. Ch. Anagnostou, 50–58. Commission of the European Union: Final Scientific Report Metro-Med Project, EU MAS3-CT96-0049.Google Scholar
  67. Pagou, K., Assimakopoulou, G., Krasakopoulou, E., and Pavlidou, A. 2004. WP4: Transport pathways and cycling of the resuspended key elements and pollutants and released nutrients to the near bottom waters. Impact to the coastal-shelf ecosystem and associated biochemical processes. In EU—Energy, Environment and Sustainable Development—INTERPOL Project: Impact of natural and trawling events on resuspension, dispersion and fate of pollutants, ed. V. Lykousis (coord.). Final Scientific Report (1 January 2001–31 December 2003).Google Scholar
  68. Peinert, R., B. von Bodungen, and V. Smetacek. 1989. Food web structure and loss rate. In Productivity of the ocean: Present and past, ed. W.H. Berger, V.S. Smetacek, and G. Wefer, 35–48. New York: Wiley.Google Scholar
  69. Petersen, G.H., and M.A. Curtis. 1980. Differences in energy flow through major components of the subarctic, temperate and tropical marine shelf ecosystems. Dana 1: 53–64.Google Scholar
  70. Poulsen, L.K. and T. Kiørboe. 2005. Coprophagy and coprorhexy in the copepods Acartia tonsa and Temora longicornis: clearance rates and feeding behaviour. Marine Ecology Progress Series 299: 217–227.Google Scholar
  71. Puskaric, S., S.W. Fowler, and J.-C. Miquel. 1992. Temporal changes in particulate flux in the Northern Adriatic Sea. Estuarine, Coastal and Shelf Science 35: 267–287.CrossRefGoogle Scholar
  72. Putt, M., and D.K. Stoecker. 1989. An experimentally determined carbon:volume ratio for marine oligotrichous ciliates from estuarine and coastal waters. Limnology and Oceanography 34: 1097–1103.CrossRefGoogle Scholar
  73. Ratkova, T.N., P. Wassmann, P.G. Verity, and I.J. Andreassen. 1999. Abundance and biomass of pico-, nano- and microplankton along a transect on Nordvestbanken, north Norwegian shelf, in 1994. Sarsia 84: 213–225.Google Scholar
  74. Reigstad, M., C. Wexels Riser and C. Svensen. 2005. Fate of copepod faecal pellets and the role of Oithona. Marine Ecology Progress Series 304: 265–270.Google Scholar
  75. Ribera d’Alcalà, M., F. Conversano, F. Corato, P. Licandro, O. Mangoni, D. Marino, M.G. Mazzocchi, M. Modigh, M. Montresor, M. Nardella, V. Saggiomo, D. Sarno, and A. Zingone. 2004. Seasonal patterns in plankton communities in a pluriannual time series at a coastal Mediterranean site (Gulf of Naples): An attempt to discern recurrences and trends. Scientia Marina 68(1): 65–83.Google Scholar
  76. Rousseau, V., S. Mathot, and C. Lancelot. 1990. Calculating carbon biomass of Phaeocystis sp. from microscopic observations. Marine Biology 107: 305–314.CrossRefGoogle Scholar
  77. Sato, R., Y. Tanaka, and T. Ishimaru. 2001. House production by Oikopleura dioica (Tunicata, Appendicularia) under laboratory conditions. Journal of Plankton Research 23: 415–423.CrossRefGoogle Scholar
  78. Sato, R., Y. Ishibashi, Y. Tanaka, T. Ishimaru, and M.J. Dagg. 2008. Productivity and grazing impact of Oikopleura dioica (Tunicata, Appendicularia) in Tokyo Bay. Journal of Plankton Research 30(3): 299–309.CrossRefGoogle Scholar
  79. Silver, M.W., S.L. Coale, C.H. Pilskaln, and D.R. Steinberg. 1998. Giant aggregates: Importance as microbial centers and agents of material flux in the mesopelagic zone. Limnology and Oceanography 43: 498–507.CrossRefGoogle Scholar
  80. Simboura, N., and S. Reizopoulou. 2008. An intercalibration of classification metrics of benthic macroinvertebrates in coastal and transitional ecosystems of the Eastern Mediterranean ecoregion (Greece). Marine Pollution Bulletin 56: 116.CrossRefGoogle Scholar
  81. Siokou-Frangou, I. 1996. Zooplankton annual cycle in a Mediterranean coastal area. Journal of Plankton Research 18: 203–223.CrossRefGoogle Scholar
  82. Siokou-Frangou, I., and E. Papathanassiou. 1991. Differentiation of zooplankton populations in a polluted area. Marine Ecology Progress Series 76(1): 41–51.CrossRefGoogle Scholar
  83. Smayda, T.J. 1978. From phytoplankters to biovolume. In Phytoplankton manual, ed. A. Sournia, 273–279. Paris: UNESCO Publications.Google Scholar
  84. Smetacek, V. 1984. The supply of food to the benthos. In Flows of energy and materials in marine ecosystems: Theory and practice, ed. M.J.R. Fasham, 517–547. New York: Plenum.CrossRefGoogle Scholar
  85. Souissi, S., O. Daly Yahia-Kéfi, and M.N. Daly Yahia. 2000. Spatial characterisation of nutrient dynamics in the Bay of Tunis (south-western Mediterranean) using multivariate analyses: Consequences for phyto- and zooplankton distribution. Journal of Plankton Research 22: 2039–2059.CrossRefGoogle Scholar
  86. Steinberg, D.K. 1995. Diet of copepods (Scopalatum vorax) associated with mesopelagic detritus (giant larvacean houses) in Monterey Bay, California. Marine Biology 122: 571–584.CrossRefGoogle Scholar
  87. Steinberg, D.K., M.W. Silver, C.H. Pilskaln, S.L. Coale, and J.B. Paduan. 1994. Midwater zooplankton communities on pelagic detritus (giant larvacean houses) in Monterey Bay, California. Limnology and Oceanography 39: 1606–1620.CrossRefGoogle Scholar
  88. Steinberg, D.K., M.W. Silver, and C.H. Pilskaln. 1997. Role of mesopelagic zooplankton in the community metabolism of giant larvacean house detritus in Monterey Bay, California, USA. Marine Ecology Progress Series 147: 167–179.CrossRefGoogle Scholar
  89. Strickland, J.D.H., and T.R. Parsons. 1968. A practical handbook of sea water analysis. Bulletin of the Fisheries Research Board of Canada 167: 310.Google Scholar
  90. Svensen, C., and J.C. Nejstgaard. 2003. Is sedimentation of copepod faecal pellets determined by cyclopoids? Evidence from enclosed ecosystems. Journal of Plankton Research 25: 917–926.Google Scholar
  91. Svensen, C., D. Vilicic, P. Wassmann, E. Arashkevich, and T. Ratkova. 2007. Plankton distribution and vertical flux of biogenic matter during high summer stratification in the Krka estuary (Eastern Adriatic). Estuarine, Coastal and Shelf Science 71(3–4): 381–390.CrossRefGoogle Scholar
  92. Taguchi, S. 1982. Seasonal study of faecal pellets and discarded houses of Appendicularia in a subtropical inlet, Kaneohe Bay, Hawaii. Estuarine, Coastal and Shelf Science 14: 545–555.CrossRefGoogle Scholar
  93. Tiselius, P., J.K. Petersen, T.G. Nielsen, M. Maar, E.F. Møller, S. Satapoomin, K. Tönnesson, S. Zervoudaki, E.D. Christou, A. Giannakourou, A. Sell, and C. Vargas. 2003. Functional response of Oikopleura dioica to house clogging due to exposure to algae of different sizes. Marine Biology 142: 253–261.Google Scholar
  94. Tragou, E., V. Zervakis, E. Papageorgiou, S. Stavrakakis, and V. Lykousis. 2005. Monitoring the physical forcing of resuspension events in the Thermaikos Gulf–NW Aegean during 2001–2003. Continental Shelf Research 25(19–20): 2315–2331.CrossRefGoogle Scholar
  95. Turner, J.T. 2002. Zooplankton fecal pellets, marine snow and sinking phytoplankton blooms. Aquatic Microbial Ecology 27: 57–102.CrossRefGoogle Scholar
  96. UNESCO. 1968. Zooplankton sampling. Paris: The UNESCO Press.Google Scholar
  97. Urban, J.L., C.H. McKenzie, and D. Deibel. 1992. Seasonal differences in the content of Oikopleura vanhoeffeni and Calanus finmarchicus faecal pellets: Illustrations of plankton food web shifts in coastal Newfoundland waters. Marine Ecology Progress Series 84: 255–264.CrossRefGoogle Scholar
  98. Ütermohl, H. 1958. Zur Vervollkommnumg der quantitativen Phytoplankton Methodik. Mitteilungen der Internationale Limnologische Vereingung 9: 1–38.Google Scholar
  99. Vargas, C., K. Tonnesson, A. Sell, M. Maar, E.F. Moller, S. Zervoudaki, A. Giannakourou, E.D. Christou, S. Satapoomin, J.K. Petersen, T.G. Nielsen, and P. Tiselius. 2002. Importance of copepods versus appendicularians in vertical carbon fluxes in a Swedish fjord. Marine Ecology Progress Series 241: 125–138.CrossRefGoogle Scholar
  100. Verardo, D.J., P.N. Froelich, and A. McIntyre. 1990. Determination of organic carbon and nitrogen in marine sediments using the Carlo Erba NA-1500 Analyzer. Deep Sea Research 37: 157–165.CrossRefGoogle Scholar
  101. Wassmann, P., Fonda Umani, S., Ypma, J.E., Reigstad, M., Cok, S., Salvi, C., Cauwet, G. 1999. Suspended biomass and vertical flux in the Gulf of Trieste (Northern Adriatic Sea). Two summer scenarios. Physical and biogeochemical processes in the Adriatic Sea. Ecosystem Research Report No. 32. The Adriatic Sea, 537–551.Google Scholar
  102. Welschmeyer, N. 1994. A method for the determination of chlorophyll a in the presence of chlorophyll b and pheopigments. Limnology and Oceanography 39: 1985–1992.CrossRefGoogle Scholar
  103. Zervakis, V., A.P. Karageorgis, H. Kontoyiannis, V. Papadopoulos, and V. Lykousis. 2005. Hydrology, circulation and distribution of particulate matter in Thermaikos Gulf (NW Aegean Sea), during September 2001–October 2001 and February 2002. Continental Shelf Research 25(19–20): 2332–2349.CrossRefGoogle Scholar
  104. Zervoudaki, S., E.D. Christou, T.G. Nielsen, I. Siokou-Frangou, G. Assimakopoulou, A. Giannakourou, M. Maar, K. Pagou, E. Krasakopoulou, U. Christaki, and M. Moraitou-Apostolopoulou. 2007. The importance of small-sized copepods in a frontal area of the Aegean Sea. Journal of Plankton Research 29(4): 317–338.CrossRefGoogle Scholar
  105. Zervoudaki, S., E.D. Christou, G. Assimakopoulou, H. Örek, A.C. Gucu, A. Giannakourou, P. Pitta, T. Terbiyik, N. Yυcel, T. Moutsopoulos, K. Pagou, S. Psarra, E. Özsoy, and E. Papathanassiou. 2011. Copepod communities, production and grazing in the Turkish Straits System and the adjacent northern Aegean Sea during spring. Journal of Marine Systems 86(3–4): 45–56.CrossRefGoogle Scholar

Copyright information

© Coastal and Estuarine Research Federation 2013

Authors and Affiliations

  • S. Zervoudaki
    • 1
  • C. Frangoulis
    • 1
  • C. Svensen
    • 2
  • E. D. Christou
    • 1
  • E. Tragou
    • 3
  • E. G. Arashkevich
    • 4
  • T. N. Ratkova
    • 4
  • Ι. Varkitzi
    • 1
  • E. Krasakopoulou
    • 1
    • 3
  • K. Pagou
    • 1
  1. 1.Institute of OceanographyHellenic Centre for Marine ResearchAnavissosGreece
  2. 2.Faculty of Bioscience, Fisheries and EconomicsUiT—The Arctic University of NorwayTromsøNorway
  3. 3.Department of Marine SciencesUniversity of the AegeanAegeanGreece
  4. 4.P.P. Shirshov Institute of OceanologyRussian Academy of SciencesMoscowRussia

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