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Larvae of coastal gastropods sink in turbulence and may use nearshore turbulence as an initial settlement cue. Our objective was to quantify the relationship between turbulence and the proportion of sinking larvae for competent mud snail veligers (Ilyanassa obsoleta). We exposed larvae to a range of field-relevant turbulence conditions ( ϭ 8.1 ϫ 10 Ϫ3 to 2.7 ϫ 10 0 cm 2 s Ϫ3 ) in a grid-stirred tank, holding other factors constant. We used a video plankton recorder to record larval movements in still water and in turbulence. Larval trajectories and velocity measurements were extracted using video-image analysis. We also measured turbulent flow velocities independently, using laser Doppler velocimetry. To interpret empirical measurements in terms of larval behavior, we developed a threecomponent, normal mixture model for vertical velocity distributions of larvae in turbulence. The model was fitted to observed larval velocities by maximum likelihood, to estimate the proportions of sinking, hovering, and swimming larvae. Over the range of turbulence intensities found in typical coastal habitats, the proportion of sinking larvae increased exponentially (r 2 ϭ 0.89) with the log of the turbulence dissipation rate. The net mean behavioral velocity of the larvae shifted from positive to negative when the dissipation rate reached ϳ10 Ϫ1 cm 2 s Ϫ3 . By sinking when they enter turbulent, shallow water, competent larvae could improve their chances of settling in favorable coastal habitats.Very little is known about how larval behavior in the plankton affects patterns of larval supply and settlement of benthic invertebrates. Much work has been done to describe larval behavior during the exploration of substrates, when larvae can sometimes select settlement sites over small scales (millimeters to centimeters). Less progress has been made on understanding the behavioral contribution while larvae are transported through the water column to benthic habitats. Under some hydrodynamic conditions, larvae could settle more successfully if they responded to waterborne cues by sinking toward the benthos. If the swimming velocity and gravitational sinking velocity differ by a factor of two or more, behavioral changes can significantly affect larval sink-1 Corresponding author (hfuchs@whoi.edu). AcknowledgmentsY. Yamashita helped collect and culture the larvae. We are grateful to J. Sisson for assistance with LDV measurements and to J. H. Trowbridge for guidance on spectral analysis. B. Raubenheimer and E. A. Terray also gave advice on flow data analysis. We thank C. DiBacco for generously sharing his culturing expertise and supplies, V. R. Starczak for advising us on the experimental design, and S. P. McKenna for introducing us to the turbulence tank. S. M. Gallager provided video equipment, software, and advice on particle tracking.

SUMMARYOyster larvae (Crassostrea virginica) could enhance their settlement success by moving toward the seafloor in the strong turbulence associated with coastal habitats. We characterized the behavior of individual oyster larvae in grid-generated turbulence by measuring larval velocities and flow velocities simultaneously using infrared particle image velocimetry. We estimated larval behavioral velocities and propulsive forces as functions of the kinetic energy dissipation rate ε, strain rate γ, vorticity ξ and acceleration α. In calm water most larvae had near-zero vertical velocities despite propelling themselves upward (swimming). In stronger turbulence all larvae used more propulsive force, but relative to the larval axis, larvae propelled themselves downward (diving) instead of upward more frequently and more forcefully. Vertical velocity magnitudes of both swimmers and divers increased with turbulence, but the swimming velocity leveled off as larvae were rotated away from their stable, velum-up orientation in strong turbulence. Diving speeds rose steadily with turbulence intensity to several times the terminal fall velocity in still water. Rapid dives may require a switch from ciliary swimming to another propulsive mode such as flapping the velum, which would become energetically efficient at the intermediate Reynolds numbers attained by larvae in strong turbulence. We expected larvae to respond to spatial or temporal velocity gradients, but although the diving frequency changed abruptly at a threshold acceleration, the variation in propulsive force and behavioral velocity was best explained by the dissipation rate. Downward propulsion could enhance oyster larval settlement by raising the probability of larval contact with oyster reef patches.

Hydrodynamic signals from turbulence and waves may provide marine invertebrate larvae with behavioral cues that affect the pathways and energetic costs of larval delivery to adult habitats. Oysters (Crassostrea virginica) live in sheltered estuaries with strong turbulence and small waves, but their larvae can be transported into coastal waters with large waves. These contrasting environments have different ranges of hydrodynamic signals, because turbulence generally produces higher spatial velocity gradients, whereas waves can produce higher temporal velocity gradients. To understand how physical processes affect oyster larval behavior, transport and energetics, we exposed larvae to different combinations of turbulence and waves in flow tanks with (1) wavy turbulence, (2) a seiche and (3) rectilinear accelerations. We quantified behavioral responses of individual larvae to local instantaneous flows using twophase, infrared particle-image velocimetry. Both high dissipation rates and high wave-generated accelerations induced most larvae to swim faster upward. High dissipation rates also induced some rapid, active dives, whereas high accelerations induced only weak active dives. In both turbulence and waves, faster swimming and active diving were achieved through an increase in propulsive force and power output that would carry a high energetic cost. Swimming costs could be offset if larvae reaching surface waters had a higher probability of being transported shoreward by Stokes drift, whereas diving costs could be offset by enhanced settlement or predator avoidance. These complex behaviors suggest that larvae integrate multiple hydrodynamic signals to manage dispersal tradeoffs, spending more energy to raise the probability of successful transport to suitable locations.

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