Estuaries and Coasts

, Volume 34, Issue 6, pp 1194–1204 | Cite as

Cues, Not an Endogenous Rhythm, Control the Dusk Peak in Water-Column Entry by Benthic Copepods



Individuals of some benthic species swim out of or away from the sediment surface into the water column, i.e., they emerge. Individuals of both emergent and nonemergent benthic species can be entrained by near-bottom flows. Both emergence and entrainment are of interest, e.g., for their roles in benthopelagic coupling, but the controlling factors are poorly understood. Our experiments with benthic copepods from contrasting environments showed that a factor (or factors) associated with the onset of darkness, rather than an endogenous rhythm, controls their dusk emergence. In addition, we argue that entrainment and emergence can interact in at least two ways: (1) Light-induced changes in oxygenation of the sediment pore water may affect the entrainment flux of benthic copepods and (2) if large numbers of individuals are entrained in the time leading up to sunset, few will remain in the sediment to be part of the dusk peak in emergence.


Emergence Benthos Behavior Oxygen Sulfide 



D. Oliff built the watertight cases for the light meter and the data logger. C. Hasemann, B. C. Hippolyte, I. Hoedl, G. Hurst, M. Pöhn, L. Sedlacek, K. Suderman, M. Teasdale, and A. Vopel helped in the laboratory or the field. The FSU Academic Diving Program provided equipment and training. The European Union Program “Training and Mobility of Researchers,” the Alfred Wegener Institute for Polar and Marine Research, the Kristineberg Marine Research Station of The Royal Swedish Academy of Science, the Klubban Biological Station of Uppsala University, the Florida State University Coastal and Marine Laboratory, and the Faculty of Health and Environmental Sciences of Auckland University of Technology provided funding and/or equipment.

Conflict of Interest

We do not have financial interests in the organizations that sponsored the research. We have full control of all primary data and agree to allow the journal to review the data if requested. No conflicts of interest exist.


  1. Armonies, W. 1988a. Active emergence of meiofauna from intertidal sediment. Marine Ecology Progress Series 43: 151–159.CrossRefGoogle Scholar
  2. Armonies, W. 1988b. Physical factors influencing active emergence of meiofauna from boreal intertidal sediment. Marine Ecology Progress Series 49: 277–286.CrossRefGoogle Scholar
  3. Armonies, W. 1994. Drifting meio- and macrobenthic invertebrates on tidal flats in Königshafen: A review. Helgolander Meeresuntersuchungen 48: 299–320.CrossRefGoogle Scholar
  4. Broecker, W.S., and T.H. Peng. 1974. Gas exchange rates between air and sea. Tellus 26: 185–190.CrossRefGoogle Scholar
  5. Buffan-Dubau, E., and J. Castel. 1996. Diel and seasonal vertical distribution of meiobenthic copepods in muddy sediments of a eutrophic lagoon (fishponds of Arcachon Bay). Hydrobiologia 329: 69–78.CrossRefGoogle Scholar
  6. Chandler, G.T., and J.W. Fleeger. 1983. Meiofaunal colonization of azoic estuarine sediment in Louisiana: Mechanisms of dispersal. Journal of Experimental Marine Biology and Ecology 69: 175–188.CrossRefGoogle Scholar
  7. Cohen, J.H., and R.B. Forward Jr. 2002. Spectral sensitivity of vertically migrating marine copepods. The Biological Bulletin 203: 307–314.CrossRefGoogle Scholar
  8. Cohen, J.H., and R.B. Forward Jr. 2005. Diel vertical migration of the marine copepod Calanopia americana II. Proximate role of exogenous light cues and endogenous rhythms. Marine Biology 147: 399–410.CrossRefGoogle Scholar
  9. Cubit, J. 1969. Behavior and physical factors causing migration and aggregation of the sand crab Emerita analoga (Stimpson). Ecology 50: 118–123.CrossRefGoogle Scholar
  10. Foy, M.S., and D. Thistle. 1991. On the vertical distribution of a benthic harpacticoid copepod: Field, laboratory, and flume results. Journal of Experimental Marine Biology and Ecology 153: 153–163.CrossRefGoogle Scholar
  11. Hicks, G.R.F. 1986. Distribution and behaviour of meiofaunal copepods inside and outside seagrass beds. Marine Ecology Progress Series 31: 159–170.CrossRefGoogle Scholar
  12. Jeroschewski, P., C. Steuckart, and M. Kühl. 1996. An amperometric microsensor for the determination of H2S in aquatic environments. Analytical Chemistry 68: 4351–4357.CrossRefGoogle Scholar
  13. Jørgensen, B.B., and N.P. Revsbech. 1985. Diffusive boundary layers and the oxygen uptake of sediments and detritus. Limnology and Oceanography 30: 111–123.CrossRefGoogle Scholar
  14. Li, Y.H., and S. Gregory. 1974. Diffusion of ions in sea water and in deep-sea sediments. Geochimica et Cosmochimica Acta 38: 703–714.CrossRefGoogle Scholar
  15. Palmer, M.A. 1984. Invertebrate drift: Behavioral experiments with intertidal meiobenthos. Marine Behaviour and Physiology 10: 235–253.CrossRefGoogle Scholar
  16. Palmer, M.A. 1988. Dispersal of marine meiofauna: A review and conceptual model explaining passive transport and active emergence with implications for recruitment. Marine Ecology Progress Series 48: 81–91.CrossRefGoogle Scholar
  17. Palmer, M.A., and R.R. Brandt. 1981. Tidal variation in sediment densities of marine benthic copepods. Marine Ecology Progress Series 4: 207–212.CrossRefGoogle Scholar
  18. Revsbech, N.P. 1989. An oxygen microelectrode with a guard cathode. Limnology and Oceanography 34: 472–476.CrossRefGoogle Scholar
  19. Service, S.K., and S.S. Bell. 1987. Density-influenced active dispersal of harpacticoid copepods. Journal of Experimental Marine Biology and Ecology 114: 49–62.CrossRefGoogle Scholar
  20. Teasdale, M., K. Vopel, and D. Thistle. 2004. The timing of benthic copepod emergence. Limnology and Oceanography 49: 884–889.CrossRefGoogle Scholar
  21. Thistle, D., G.L. Weatherly, and S.C. Ertman. 1995. Shelf harpacticoid copepods do not escape into the seabed during winter storms. Journal of Marine Research 53: 847–863.CrossRefGoogle Scholar
  22. Thistle, D., L. Sedlacek, K.R. Carman, J.W. Fleeger, and J.P. Barry. 2007. Emergence in the deep sea: Evidence from harpacticoid copepods. Deep-Sea Research I 54: 1008–1014.Google Scholar
  23. Trüper, H.G., and H.G. Schlegel. 1964. Sulphur metabolism in Thiorhodaceae 1 Qualitative measurements in growing cultures of Chromatium okenii. Antonie van Leeuwenhoek 30: 225–238.CrossRefGoogle Scholar
  24. Van Gemerden, H., C.S. Tughan, R. De Wit, and R.A. Herbert. 1989. Laminated microbial ecosystems on sheltered beaches in Scapa Flow, Orkney Island. FEMS Microbiology Ecology 62: 87–102.Google Scholar
  25. Visscher, P.T., J. Beukema, and H. Van Gemerden. 1991. In situ characterization of sediments: Measurements of oxygen and sulfide profiles with a novel combined needle electrode. Limnology and Oceanography 36: 1476–1480.CrossRefGoogle Scholar
  26. Vopel, K., J. Dehmlow, and G. Arlt. 1996. Vertical distribution of Cletocamptus confluens (Copepoda, Harpacticoida) in relation to oxygen and sulphide microprofiles of a brackish water sulphuretum. Marine Ecology Progress Series 141: 129–137.CrossRefGoogle Scholar
  27. Walters, K. 1988. Diel vertical migration of sediment-associated meiofauna in subtropical sand and seagrass habitats. Journal of Experimental Marine Biology and Ecology 117: 169–186.CrossRefGoogle Scholar
  28. Walters, K. 1991. Influences of abundance, behavior, species composition, and ontogenetic stage on active emergence of meiobenthic copepods in subtropical habitats. Marine Biology 108: 207–215.CrossRefGoogle Scholar
  29. Walters, K., and S.S. Bell. 1986. Diel patterns of active migration in seagrass meiofauna. Marine Ecology Progress Series 34: 95–103.CrossRefGoogle Scholar

Copyright information

© Coastal and Estuarine Research Federation 2011

Authors and Affiliations

  1. 1.Department of Earth, Ocean, and Atmospheric ScienceFlorida State UniversityTallahasseeUSA
  2. 2.School of Applied SciencesAuckland University of TechnologyAucklandNew Zealand
  3. 3.Department of Earth, Ocean, and Atmospheric Sciences/OceanographyFlorida State UniversityTallahasseeUSA

Personalised recommendations