Swimming behavior of juvenile anchovies (Anchoa spp.) in an episodically hypoxic estuary: implications for individual energetics and trophic dynamics

Diel swimming behaviors of juvenile anchovies (Anchoa spp.) were observed using stationary hydroacoustics and synoptic physicochemical and zooplankton profiles during four unique water quality scenarios in the Neuse River Estuary, NC, USA. Vertical distribution of fish was restricted to waters with DO greater than 2.5 mg O-2 l(-1), except when greater than 70% of the water column was hypoxic and a subset of fish were occupying water with 1 mg O-2 l(-1). We made the prediction that an individual fish would select a swim speed that would maximize net energy gain given the abundance and availability of prey in the normoxic waters. During the day, fish adopted swim speeds between 7 and 8.8 bl s(-1) that were near the theoretical optimum speeds between 7.0 and 8.0 bl s(-1). An exception was found during severe hypoxia, when fish were swimming at 60% above the optimum speed (observed speed = 10.6 bl s(-1), expected = 6.4 bl s(-1)). The anchovy is a visual planktivore; therefore, we expected a diel activity pattern characteristic of a diurnal species, with quiescence at night to minimize energetic costs. Under stratified and hypoxic conditions with high fish density coupled with limited prey availability, anchovies sustained high swimming speeds at night. The sustained nighttime activity resulted in estimated daily energy expenditure over 20% greater than fish that adopted a diurnal activity pattern. We provide evidence that the sustained nighttime activity patterns are a result of foraging at night due to a lower ration achieved during the day. During severe hypoxic events, we also observed individual fish making brief forays into the hypoxic hypolimnion. These bottom waters generally contained higher prey (copepod) concentrations than the surface waters. The bay anchovy, a facultative particle forager, adopts a range of behaviors to compensate for the effects of increased conspecific density and reduced prey availability in the presence of stratification-induced hypoxia.