Thin Layers: Physical and Chemical Cues Contributing to Observed Copepod Aggergations
Woodson, Clifton Brock
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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.