Marine Biology

, Volume 149, Issue 3, pp 573–584 | Cite as

Diel vertical migration of Sagitta setosa as inferred acoustically in the Black Sea

  • Erhan MutluEmail author
Research Article


Swimming trajectories of chaetognaths Sagitta setosa Müller in the Black Sea were studied using an echosounder operating at 120 and 200 kHz and an acoustic Doppler current profiler (ADCP) operating at 150 kHz. S. setosa were acoustically discriminated with respect to vertical migration and swimming speed, according to dissolved oxygen (DO) concentration and the timing of migrations. S. setosa formed a concentration layer thicker than Calanus euxinus did (1–3 m). The migration was completed in about 2.5–4 h, upward migration starting before C. euxinus and downward migration after C. euxinus. Adult Sagitta swam fast only in the well-oxygenated layer (subsurface maximum DO). The DO was found to be a significant (p<0.05) variable by partial correlation between the speed and hydrographical parameters. This feature constituted an oxygen-dependent influence on S. setosa’s vertical swimming and distinguished S. setosa from C. euxinus. Chaetognaths migrated daily between the nearsurface and the oxycline or the suboxic zone (OMZ, see Fig. 3b for the layers characterized by DO). Whether the deepest depth limit of migration was the oxycline or the OMZ depended on the relative abundance of adult and immature (young) individuals in the concentration layer. In July and September, individuals belonging to a new generation did not migrate but stayed in subsurface water day and night.


Swimming Speed Vertical Migration Acoustic Doppler Current Profiler Diel Vertical Migration Concentration Layer 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.



This work was carried out within the NATO TU-Fisheries and Black Sea projects. The IMS-METU was funded by the Scientific and Technical Research Council of Turkey (TUBITAK); by the Scientific Affairs Division of NATO as part of the Science for Stability program; and by a project (METU-AFP-99-06-01-01) linked with other programs of TUBITAK/Turkey and NATO-SfP and a project funded by TUBITAK (YDABAG-100Y071). The hydrographical data were obtained from the Physical and Chemical Oceanography Department of the IMS-METU. I thank the crew of R.V. “Bilim” for assistance at sea. Dr. S. Besiktepe provided zooplankton data (April, September 1995 and June 1996; Besiktepe and Unsal 2000; Besiktepe 2001). I also thank Mr. V. Myroshnychenko for providing me with a PC program to extract echo intensity from ADCP data and Drs. Mark C. Benfield and Karen Fisher for their valuable comments and correction of English of the text. I am much indebted to anonymous referees for their constructive comments on the manuscript.


  1. Benoit-Bird KJ, Au WWL (2001) Target strength measurements of Hawaiian mesopelagic boundary community animals. J Acoust Soc Am 110(2):812–819CrossRefGoogle Scholar
  2. Besiktepe S (2001) Diel vertical distribution, and herbivory of copepods in the south-western part of the Black Sea. J Mar Syst 28:281–301CrossRefGoogle Scholar
  3. Besiktepe S, Unsal M (2000) Population structure, vertical distribution and diel migration of Sagitta setosa (Chaetognatha) in the south-western part of the Black Sea. J Plankton Res 22:669–683CrossRefGoogle Scholar
  4. Bone Q, Duvert M (1991) Locomotion and buoyancy. In: Bone Q, Kapp H, Pierrot-Bults AC (eds) The biology of Chaetognaths, Oxford University Press, New York, pp 32–44Google Scholar
  5. Brierley AS, Ward P, Watkins JL, Goss C (1998) Acoustic discrimination of Southern Ocean zooplankton. Deep Sea Res II 45(7):1155CrossRefGoogle Scholar
  6. Childress JJ, Cowles DL, Favuzzi JA, Mickel TJ (1990) Metabolic rates of benthic deep-sea decapod crustaceans decline with increasing depth primarily due to the decline in temperature. Deep-Sea Res 37:929–949CrossRefGoogle Scholar
  7. Cushing DH (1951) The vertical migration of planktonic crustacea. Biol Rev 26:158–192CrossRefGoogle Scholar
  8. David PM, Guerin-Ancey O, Van Cuyck JP (1999) Acoustic discrimination of two zooplankton species (mysid) at 38 and 120 kHz. Deep-Sea Res I 46:319–333CrossRefGoogle Scholar
  9. Drits AV, Utkina SV (1988) Feeding of Sagitta setosa in layer of daytime plankton accumulation in the Black Sea. Oceanology 28:781–785Google Scholar
  10. Duvert M, Salat C (1990) Ultrastructural studies on the fins of chaetognaths. Tissue Cell 22(6):853–863CrossRefGoogle Scholar
  11. Erkan F, Gucu AC, Zagorodnyaya J (2000) The diel vertical distribution of zooplankton in the southeast Black Sea. Turk J Zool 24:417–428Google Scholar
  12. Feigenbaum D (1991) Food and feeding behaviour. In: Bone Q, Kapp H, Pierrot-Bults AC (eds) The biology of Chaetognaths, Oxford University Press, New York, pp 45–54Google Scholar
  13. Foote KG, Stanton TK (2000) Acoustical methods, ICES zooplankton methodology manual. Chap 6. Academic, New York, pp 223–258CrossRefGoogle Scholar
  14. Greze VN (1970) The biomass and production of different trophic level in the pelagic communities of South Seas. In: Steele JS (ed) Marine food chains. Oliver & Boyd, Edinburgh, pp 458–467Google Scholar
  15. Hammer WM, Hammer PP, Strand SW (1994) Sun-compass migration of Aurelia aurita (Scyphozoa): population retention and reproduction in Saanich Inlet, British Columbia. Mar Biol 119: 347–356CrossRefGoogle Scholar
  16. Kideys AE, Romanova Z (2001) Distribution of gelatinous macrozooplankton in the southern Black Sea during 1996–1999. Mar Biol 139:535–547CrossRefGoogle Scholar
  17. Mironov GN (1960) The feeding of plankton predators. II. The feeding of Sagittae. Trudy Sevastopol’ Biol Sta 13:78–88 (in Russian)Google Scholar
  18. Monger BC, Chinna-Chandy S, Meir E, Billings S, Greene CH, Wiebe PH (1998) Sound scattering gelatinous zooplankters Aequorea victoria and Pleurobrachia bachei. Deep-Sea Res II 45:1255–1271CrossRefGoogle Scholar
  19. Mutlu E (1996) Target strength of the common jellyfish (Aurelia aurita): a preliminary experimental study with a dual-beam acoustic system. ICES J Mar Sci 53(2): 309–311CrossRefGoogle Scholar
  20. Mutlu E (2001) Distribution and abundance of moon jellyfish (Aurelia aurita) and its zooplankton food in the Black Sea. Mar Biol 138(2):329–339CrossRefGoogle Scholar
  21. Mutlu E (2003a) Acoustical identification of the concentration layer of a copepod species, Calanus euxinus. Mar Biol 142:517–523CrossRefGoogle Scholar
  22. Mutlu E (2003b) Diel vertical distribution of zooplankton in the Black Sea (April 1995). In: Yilmaz A (Ed), Proceedings of second international conference on oceanography of the Eastern Mediterranean and Black Sea: similarities and differences of two interconnected basins, TUBITAK, Ankara, pp 753–760Google Scholar
  23. Mutlu E (2005) An intercomparison of the contribution of zooplankton and nekton taxa to the near-surface acoustic structure of three Turkish Seas. Mar Ecol 26: 17–32CrossRefGoogle Scholar
  24. Mutlu E, Bingel F (1999) Distribution and abundance of ctenophores and their zooplankton food in the Black Sea. I. Pleurobrachia pileus. Mar Biol 135:589–601CrossRefGoogle Scholar
  25. Niermann U, Bingel F, Ergun G, Greve W (1998) Fluctuation of dominant mesozooplankton species in the Black Sea, North Sea and the Baltic Sea: Is a general trend recognizable? Tr J Zool 22:63–81Google Scholar
  26. Ozsoy E, Unluata U (1997) Oceanography of the Black Sea: a review of some recent results. Earth-Sci Rev 42:231–272CrossRefGoogle Scholar
  27. Pearre S Jr (1979) On the adaptive significance of vertical migration. Limnol Oceanogr 24:781–782CrossRefGoogle Scholar
  28. Pearre S Jr (1991) Growth and reproduction. In: Bone Q, Kapp H, Pierrot-Bults AC (eds) The biology of Chaetognaths, Oxford University Press, New York, pp 61–75Google Scholar
  29. Petipa TS, Pavlova EV, Mironov GN (1970) The food web structure, utilization and transport of energy by trophic levels in the planktonic communities. In: Steele JS (ed) Marine food chains. Oliver & Boyd, Edinburgh, pp 419–457Google Scholar
  30. Poyarkov SG (1989) Pecularities of water hydrochemical structure in relation to its stratification. In: Vinogradov ME, Flint MV (eds) Structure and productional characteristics of planktonic populations in the Black Sea. Nauka, Moskow, pp 10–23 (in Russian)Google Scholar
  31. RDI (1999) Transect–User’s manual. RD Instrument, P/N 951-6092-00, San Diego, 170 ppGoogle Scholar
  32. Reeve MR, Walter MA, Ikeda T (1978) Laboratory studies of ingestion and food utilization in lobate and tentaculate ctenophores. Limnol Oceanogr 23(4):740–751CrossRefGoogle Scholar
  33. Stanton TK, Chu D (2000) Review and recommendations for the modeling of acoustic scattering by fluid-like elongated zooplankton: euphausiids and copepods. ICES J Mar Sci 57:793–807CrossRefGoogle Scholar
  34. Stanton TK, Chu D, Wiebe PH (1996) Acoustic scattering characteristics of several zooplankton groups. ICES J Mar Sci 53:289–295CrossRefGoogle Scholar
  35. Stanton TK, Chu D, Wiebe PH (1998b) Sound scattering by several zooplankton. II. Scattering models. J Acoust Soc Am 103:236–253CrossRefGoogle Scholar
  36. Stanton TK, Chu D, Wiebe PH, Martin LV, Eastwood RL (1998a) Sound scattering by several zooplankton. I. Experimental determination of dominant scattering mechanisms. J Acoust Soc Am 103:225–235CrossRefGoogle Scholar
  37. Stanton TK, Wiebe PH, Chu D, Benfield MC, Scanlon L, Martin L, Eastwood RL (1994) On acoustic estimates of zooplankton biomass. ICES J Mar Sci 51:505–512CrossRefGoogle Scholar
  38. Svetlichny LS, Hubareva ES, Erkan F, Gucu AC (2000) Physiological and behavioral aspects of Calanus euxinus females (Copepoda: Calanoida) during vertical migration across temperature and oxygen gradients. Mar Biol 137:963–971CrossRefGoogle Scholar
  39. Thuesen EV, Childress JJ (1993) Enzymatic activities and metabolic rates of pelagic chaetognaths: lack of depth-related deckines. Limnol Oceanogr 38(5):935–948CrossRefGoogle Scholar
  40. Tugrul D, Basturk O, Saydam C, Yilmaz A (1992) Changes in the hydrochemistry of the Black Sea inferred from the water density profiles. Nature 359:137–139CrossRefGoogle Scholar
  41. Vinogradov MYe, Flint MV, Shushkina EA (1985) Vertical distribution of mesoplankton in the open area of the Black Sea. Mar Biol 89:95–107CrossRefGoogle Scholar
  42. Vinogradov MYe, Musayeva EI, Semenova TN (1990) Factors determining the position of the lower layer of mesoplankton concentration in the Black Sea. Oceanology 30:217–224Google Scholar
  43. Vinogradov MYe, Sapozhnikov VV, Shushkina EA (1992) The Black Sea ecosystem. Nauka, MoscowGoogle Scholar
  44. Wiebe PH, Greene CH, Stanton TK, Burczynski J (1990) Sound scattering by live zooplankton and micronekton: empirical studies with a dual-beam acoustical system. J Acous Soc Am 88:2346–2360CrossRefGoogle Scholar
  45. Wiebe PH, Greene CH (1994) The use of high frequency acoustics in the study of zooplankton spatial and temporal patterns. In: Proceedings of the NIPR symposium on polar biology. NIPR, Tokyo, pp 133–157Google Scholar
  46. Wiebe PH, Mountain DG, Stanton TK, Greene CH, Lough G, Kaartvedt S, Dawson J, Copley N (1996) Acoustical study of the spatial distribution of plankton on Georges Bank and the relationship between volume backscattering strength and the taxonomic composition of the plankton. Deep-Sea Res II 43:1971–2001CrossRefGoogle Scholar
  47. Yilmaz A, Tugrul S, Polat C, Ediger D, Coban Y, Morkoc E (1998) On the production, elemental composition (C, N, P) and distribution of photosynthetic organic matter in the southern Black Sea. Hydrobiologia 363:141–145CrossRefGoogle Scholar
  48. Zenkevitch L (1963) Biology of the Seas of the USSR. George Allen and Unwin Ltd, LondonGoogle Scholar

Copyright information

© Springer-Verlag 2005

Authors and Affiliations

  1. 1.Institute of Marine SciencesMiddle East Technical UniversityMersinTurkey

Personalised recommendations