In the current study, behavioral responses of several species of calanoid copepods to mimics of oceanographic structure were observed and evaluated in the context of foraging and aggregation. Zooplankton distributions in oceanic habitats are often attributed to physical forcing; however, physical factors only act to drive ecological patterns at large scales (m to km). At fine to intermediate scales (cm to m) zooplankton behavior is believed to govern observed distributions, but the mechanisms and ecological significance of these behaviors are not well understood. In a water column, biological activity is often concentrated into one or a few regions, called thin layers, on the order of a meter thick, and zooplankton, such as copepods, must be able to reliably locate and exploit these patches for survival. Thin layers commonly are associated with oceanic structure such as flow gradients, fluid density jumps, or chemical composition gradients. Utilization of mechanosensory or chemosensory cues associated with thin layers may increase foraging success, thus translating into a significant ecological advantage. A laboratory apparatus was developed to create isolated and combined thin layer properties. Copepods then were exposed to laboratory mimics of thin layers. All of the tested species of copepods exhibited behavioral responses associated with area-restricted search behavior to one of the physical gradients (flow velocity or fluid density), but not both. Similar responses were observed for chemical exudate layer experiments and included increased proportional residence times, swimming speeds, and turn frequency. Food layers induced feeding responses from all tested species (increased proportional residence time, decreased swimming speed). Responses to various combinations of gradients were not fully synergistic, but suggested that some copepods employ a cue hierarchy to locate food-rich areas. Velocity or density gradients acted as initial cues for narrowing search regions, while chemical exudates induced responses that strengthened or removed the initial reactions. A simple foraging model was used to illustrate how such behavioral changes can lead to observed aggregations at larger temporal and spatial scales. Consequently, these results suggest that individual responses to oceanographic structure may have far reaching influence on population dynamics, succession, and biodiversity in coastal and pelagic ecosystems.

Copepod species are distributed throughout the ocean by many factors, including chemical, biological, and physical effects. Turbulence in the ocean has been suggested as a factor that vertically partitions some species of copepod. Copepods may seek calmer waters by sinking to deeper levels as the surface waters become more turbulent, or may maintain their position in turbulent waters. The goal of this study is to determine the behavioral effects of turbulence on three species of copepod, Calanus finmarchicus, Acartia tonsa, and Temora longicornis. Experiments consisted of exposing each of the species to stagnant water plus four levels of turbulence intensity. The experiments were conducted in a laboratory apparatus that mimics oceanic turbulence. The turbulence characteristics have been previously characterized by particle image velocimetry (PIV), that show the turbulence to be nearly isotropic and homogeneous in the observation region. Behavior responses were quantified via several measures, including the number of animals phototactically aggregating per minute, the number of escape events, the swimming speed, and the net-to-gross-displacement ratio. There are important conclusions about the effect of laboratory turbulence on copepods. The size of the copepod has a significant effect on its aggregation and swimming capability with increasing turbulence. The smaller copepods had less ability to overcome a strong flow field, and they were more likely to be advected by the stronger flow fields. Swim style also can influence how a copepod reacts to increased turbulence. If the copepod is a hop and sink traveler, then the copepod continues to hop and sink more than its cruising counterparts as turbulence increases. The cruise and sink travelers did not alter the number of escapes in response to turbulence.