We summarize what is known of the biophysical interactions that control vertical migration and dispersal of decapod larvae, asking the following main questions: How common is vertical migration in decapod crustacean larvae? What is the vertical extent of the migrations? What are the behavioural mechanisms that control vertical migrations? How does vertical migration interact with the physics of the ocean to control the dispersal of larvae? These questions are analysed by first giving a synopsis of the physical processes that are believed to significantly affect horizontal transport, and then by describing migration patterns according to taxon, to ecological category based on the habitat of adults and larvae, and to stage within the larval series. Some kind of vertical migration has been found in larval stages of virtually all species that have been investigated, irrespective of taxonomic or ecological category. Most vertical migration schedules have a cyclic nature that is related to a major environmental cyclic factor. Tidal (ebb or flood) migration and daily (nocturnal and twilight) migration are the two types of cyclic migration that have been identified. In general, all species show some type of daily migration, with nocturnal migration being the most common, whereas tidal migrations have only been identified in species that use estuaries during part of their life cycle. Moreover, there are several examples indicating that the phasing and extent of migration both change throughout ontogeny. Reported ranges of vertical displacement vary between a few metres in estuaries and several tens of metres (sometimes more than 100 m) in shelf and oceanic waters. Vertical movements are controlled by behavioural responses to the main factors of the marine environment. The most important factors in this respect are light, pressure and gravity, but salinity, temperature, turbulence, current and other factors, also influence behaviour. Many of these factors change cyclically, and the larvae respond with cyclic behaviours. The type of response may be endogenous and regulated by an internal clock, as in the case of some tidally synchronised migrations, but in most cases it is a direct response to a change in an environmental variable, as in diel migration. The reaction of the larvae to exogenous cues depends both on the rate of change of the variable and on the absolute amount of change. A series of dispersal types, involving different spatial and temporal scales, have been identified in decapod larvae: retention of the larval series within estuaries; export from estuarine habitats, dispersal over the shelf, and reinvasion of estuaries by the last stage; hatching in shelf waters and immigration to estuaries by late larvae or postlarvae; complete development on the shelf; and hatching in shelf waters, long-range dispersal in the ocean, and return to the shelf by late stages. In all of these cases, vertical migration behaviour and changes of behaviour during the course of larval development have been related to particular physical processes, resulting in conceptual mechanisms that explain dispersal and recruitment. Most decapod larvae are capable of crossing the vertical temperature differences normally found across thermoclines in natural systems. This ability may have significant consequences for horizontal transport within shelf waters, because amplitude and phase differences of the tidal currents across the thermocline may be reflected in different trajectories of the migrating larvae.

Interactions between behaviour and physical forcing in the control of horizontal transport of decapod crustacean larvae.

Blue crabs Callinectes sapidus Rathbun were sampled in a lower Chesapeake Bay seagrass bed and adjacent tidal marsh creek from October 1982 through December 1986, using a drop net and suction sampler. Suction sampling was 88 % efficient and provided more accurate estimates of juvenile blue crab abundance, that were at least an order of magnitude greater than those obtained by conventional trawl gear For data analysis, C. sapidus were separated into 4 size classes by carapace width-54.25 mm (recently settled 1st and 2nd juveniles); 4.25 to 11.00 mm (3rd through 7th juvenile crabs); > 11.00 to 525.00 mm (8th through 12th juvenile crabs); and >25.00 mm (older juveniles and adults). Seasonal and annual cycles of C. sapidus abundance were observed in both habitats with annual differences most pronounced in the grassbed. Densities of C. sapidus were significantly greater in the grassbed in all but 2 of the 48 sampling dates. Settlement of the new year class appeared to be by megalopae and occurred from August through December A pulse of small C. sapidus C4.25 mm was observed in the grassbed from early to mid-September, whereas in the marsh creek this pulse occurred 2 to 4 wk later and consisted of 3rd stage and generally larger crabs (>4.25 mm). The small numbers of 1st and 2nd stage juvenile crabs ( 25 mm throughout the study period did not differ significantly among years (1983 to 1986) even though there were significant differences in the abundance of smaller sized individuals between 1983-84 and 1985-86. This suggests mortality within or emigration from grassbeds around this size. A possible ontogenetic shift in habitat use by crabs 11 to 25 mm may reflect a refuge in size from predation.

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Ultilization of a seagrass meadow and tidal marsh creek by blue crabs Callinectes sapidus. I. Seasonal and annual variations in abundance with emphasis on post-settlement juveniles

The transport of plaice larvae from the open sea towards shallow, mainly inshore, nursery areas along the North Sea coast was studied for 5 years in the Easter Scheldt and the Wadden Sea. Metamorphosing larvae (stages 4 and 5) entered the nursery areas in March and April. Larva density was significantly correlated with the density of 0-group plaice at the end of the following summer. Significant factors determining the larval densities were date of sampling, location, tide, depth, and did period. Larvae usually were most abundant in the bottom stratum, but during nighttime flood tides they moved into midwater and surface strata. Flood catches exceeded the ebb catches of pelagic larvae for all stations. The results suggest that plaice larvae accomplish passive but selective horizontal transport by swimming up from the seabed during flood tides and remaining on the seabed during ebb tides.

Selective Tidal Transport of North Sea Plaice Larvae Pleuronectes platessa in Coastal Nursery Areas

Cannibal–prey dynamics in young juveniles and postlarvae of the blue crab

For marine benthic invertebrates with meroplanktonic larvae, the relative importance of hydrodynamics and swimming behaviour in determining larval dispersal in the water column, particularly at sma...

Behaviour in flow: perspectives on the distribution and dispersion of meroplanktonic larvae in the water column

Results of a three-year survey of the occurrence of Callinectes sapidus larvae in the mouth of Delaware Bay indicated that stage I zoea larvae were most abundant insurface water as compared to mid-depths and near bottom. The major peak in abundance of stage I zoea larvae occurred in early August with a secondary peak in early September. Peaks in abundance of megalopae occurred five weeks after the respective peaks in zoeal abundance. Zoea stages II–VIII were not collected in the bay mouth. Results of sampling every 3 h over consecutive tidal cycles showed that stage I zoea larvae were most common in the water column on ebbing tidal currents. Megalopae were most common in the water column on flooding tidal currents, suggesting a tidally related, vertical migration. It was concluded that stage I zoea larvae are flushed from the estuary and undergo development on the continental shelf. Megalopae are then transported back to inshore waters by a combination of winds and currents and invade the estuary by means of migration into the water column on flooding tidal currents and migration to the bottom on ebbing tidal currents.

C.C. Valenti

Papers

Dispersal and recruitment of blue crab larvae in Delaware Bay, U.S.A.