The “pelagic larvaton” and its role in the biology of the World Ocean,with special reference to pelagic larvae of marine bottom invertebrates

Ecological subdivision of marine organisms is often based on two characteristics: presence in a defined environment, and types of locomotion (degree of free active movement) in such an environment. The use of these characteristics results in a simple scheme: (1) Inhabitants of the boundary surface “ocean-atmosphere” (a zone including not only the surface film but also the thin subsurface water layer below it and the air layer just above it, i.e., pleuston and neuston). (2) Inhabitants of the deeper water layers of the ocean i.e., excluding the zone mentioned under (1): (a) passively drifting forms with very limited locomotory capacity, moving practically in the vertical plane only (plankton); (b) actively moving forms which migrate both vertically and horizontally (nekton). (3) Inhabitants of the “bottom”-benthos (level-bottom of oceans and coastal waters, tidal zones up to the upper supralittoral, different types of drifting and floating substrata, e.g. ship bottoms, harbour structures, buoys, driftwood, sargassum, whales, etc.). This simple scheme is essentially based on characteristics of adults. If developmental stages are considered, pelagic larvae of bottom invertebrates, eggs and larvae of fishes and other forms, usually present only temporarily in the plankton, neuston, and pleuston, can be distinguished as “mero-plankton”, “mero-neuston” and “mero-pleuston”, from the permanent “holo”-components of these groups. Division into “mero”-subgroups opposes all these larvae to those of planktonic, neustonic and pleustonic forms developing within the “parental” groups and their environments. However, the last category of larvae in the light of world-wide distribution of the seasonal reproductive pattern of marine invertebrates and some other organisms — especially in temperate and high latitudes — can also be rated to some degree as “mero”-(not “holo”-) components. The present paper proposes to unite all larvae of marine invertebrates (and of other organisms) undergoing pelagic development into one biological group, the “pelagic larvaton”. The main characteristic for all forms of this group is the presence of one and the same life-cycle stage in one and the same environment. All forms of the “pelagic larvaton” are, to various degrees, biologically different from their respective adult forms. Even the pelagic larvae of the holoplanktonic species exhibit some differences. Within the “pelagic larvaton”, 3 subgroups can be distinguished on the basis of their ecological peculiarities;

1. Larvae undergoing their whole development in an environment different from that inhabited by their parents and belonging to a group different from that of their parental forms; e.g. the pelagic larvae of bottom invertebrates which develop in the plankton, neuston or pleuston.
2. Larvae undergoing development in the same general pelagic environment, but in “non-parental” ecological groups; e.g. larvae of nektonic species developing in the plankton, neuston or pleuston; larvae of planktonic species in the neuston or pleuston; larvae of neustonic and pleustonic species in the plankton.
3. Larvae undergoing development in the “parental” groups; e.g. larvae of planktonic species in the plankton, of neustonic species in the neuston, or of pleustonic species in the pleuston.

In contrast to the 5 ecological groups: benthos, plankton, nekton, neuston and pleuston, the “pelagic larvaton” represents rather a biological than an ecological group. The “pelagic larvaton” comprises the 5 ecological groups and maintains the permanent turnover of organic substances between water and bottom. This group short-circuits the interrelations between the 5 ecological groups in all possible combinations. The existence of the “pelagic larvaton” presents another illustration of the unity of the biological nature of the oceans. The present paper also discusses the specific distributional patterns of the pelagic larvae of bottom invertebrates and their biological role in the seas.     

The influence of pollution on pelagic larvae of bottom invertebrates in marine nearshore and estuarine waters

At present, human activities on marine shores, nearshore waters and estuaries, as well as in rivers which discharge into the seas, are influencing nearshore and estuarine waters so strongly that the resultant changes affect the breeding and spawning of bottom invertebrates which inhabit these waters. The majority of bottom invertebrate species from the shallow shelf of all seas and oceans, and from all except the highest latitudes, pass through a pelagic larval phase during development. Thus, survival and maintenance of such species are determined by survival and distribution of larvae during their planktonic phase and their settling on the substrate. Consequently, an attempt to summarize all available data on the influence of pollution in marine and estuarine waters, not only on adult bottom invertebrates, but also on their pelagic larvae, will be of practical interest from both scientific and economic points of view. This paper represents a first step towards that goal. Despite the fact that all aspects of pollution-industrial wastes (including heated effluents producing thermal pollution), domestic sewage, oil and oil products, oil-spill removers and oil-emulsifiers, various pesticides, synthetic surfactants, etc.—exert harmful effects on pelagic larvae of bottom invertebrates under experimental conditions, in natural environments, free-swimming larvae are influenced only slightly by these pollutants. Pollution becomes a grave danger for larvae when they are settling on the substrate, as much higher concentrations of different pollutants may be present on the substrate than in the water mass above it (the pollution of every nearshore water mass usually varies greatly from place to place). Pelagic larvae of bottom invertebrates inhabiting comparatively clean parts of polluted regions are dispersed throughout all such regions by local currents, tidal oscillations, eddies and other small-scale water movements. They represent, therefore, the potential source for re-establishment of the normal composition of the bottom communities in these regions, after the abatement of pollution by means of natural causes or by man's own improvements.     

Types of larval development in marine bottom invertebrates,their distribution and ecological significance: a re-evaluation

The distribution of various types of larval development among marine bottom invertebrates has been discussed on the basis of ecological evidence by Thorson (1936, 1946, 1950, 1952) and Mileikovsky (1961b, 1965). The information at hand is reviewed anew in this paper and is re-evaluated in the light of modern pertinent literature. The interrelationships between certain larval types and their distribution are not as rigid and direct as originally assumed. This can be proved even by the copy book example of the distribution of the various forms of development among species of the coastal gastropod genus Littorina. Especially among species with wide distributional areas, local populations may exhibit greater diversity in larval types than has previously been thought. Different types of larval development have now become known to exist in different populations of opisthobranch gastropods and lamellibranchs, i.e., in invertebrate groups in which such variability had been ruled out by Thorson. Variability in the type of larval development within given species — as a function of geographical, seasonal and other environmental parameters —is also more common in other marine bottom invertebrates than formerly considered. Marine bottom invertebrates are characterized not only by the 3 main different types of larval development proposed by Thorson (pelagic, direct, viviparous), but also by a fourth type: demersal (free non-pelagic) development. This fourth type occurs at all water depths and in all geographic zones of the oceans. The most important of the 4 types is pelagic (planktotrophic) development. Thorson's rule (decrease in numbers of species possessing pelagic development from the Equator towards the Poles, and from shallow-shelf waters to greater oceanic depths) is well substantiated by new data. However, one correction is necessary: pelagic development is not completely absent in the abyssal zone, as was proposed by Thorson (1950, and later), but is represented in it by at least several species belonging to various groups of invertebrates, and is also fairly common in the bathyal zone. A detailed analysis of the distributional pattern of the different types of development of marine bottom invertebrates must further take into consideration asexual reproduction with all its different modifications. Asexual reproduction in benthonic invertebrates is ecologically significant because of its common occurrence in nature; in numerous species it is also important as a biological supplement to sexual reproduction. The vast majority of species inhabiting the shallow-shelf zone and, partly, the higher levels of the slope zone of ocean areas located roughly between the polar circles, reveals development by means of planktotrophic larval stages. In the highest latitudes and on the slopes to abyssal depths—characterized by low water temperatures, scarcity of food, increasing hydrostatic pressure and other environmental peculiarities—other types of larval development prevail and, progressively, replace pelagic development with increasing latitude or depth. The distributional patterns of the various types of development among marine bottom invertebrates form one of the most important factors determining the basic distributional dynamics of the whole benthos in all oceans, both in the geological past and at the present time.Dedicated to the memory of Professor G. Thorson —founder of modern reproductive and larval ecology of marine bottom invertebrates.     

Speed of active movement of pelagic larvae of marine bottom invertebrates and their ability to regulate their vertical position

Pelagic larvae of marine bottom invertebrates are able to perform different types of active vertical movements in marine nearshore and estuarine waters. The speed of these active movements is in the order of magnitude of common representatives of marine micro-and mezo-holozooplankton performing daily vertical migrations, and only slightly less than that of holozooplankton. Only a few morphological types of larvae, characterized by weak ciliary movement apparatus (amphiblastulae of sponges, ophioplutei) have a speed of movement less than 1 cm·min^-1. All other types of larvae, which possess strong ciliary movement apparatus or swimming appendages, are characterized by quick active movement from more than 1 to 60 cm·min^-1 in lamellibranch larvae, and up to 117 cm x min^-1 in decapod larvae. Because of their ability to produce sufficient speed of active movement, pelagic larvae of bottom invertebrates are able to control their vertical distribution in marine nearshore and estuarine waters in all neritic regions, including those with very strong tidal currents, except where local water stratification inhibits their active vertical movement.     

Circatidal activity rhythms in ovigerous blue crabs, <Emphasis Type="Italic">Callinectes</Emphasis><Emphasis Type="Italic">sapidus</Emphasis>: implications for ebb-tide transport during the spawning migration

Female blue crabs (Callinectes sapidus Rathbun) with mature embryos have a spawning migration in which they: (1) undergo ebb-tide transport for movement seaward from estuaries, (2) release their larvae, and (3) reverse direction by undergoing flood-tide transport for up-estuary movement. The study determined whether ebb-tide transport during the spawning migration is based upon an endogenous rhythm in vertical migration. Under constant conditions in a rectangular container, which limited horizontal and vertical movements, females with young and mature embryos had circatidal rhythms (periods=12.11-12.95 h) in migratory restlessness (swimming activity) and egg maintenance behavior (abdominal pumping). However, the rhythms were out of phase, as migratory restlessness occurred during the expected time of ebb tide in the field, and egg maintenance behavior, during the time of flood tide. Under constant conditions in vertical columns (1.32 m high), crabs with mature embryos had a circatidal rhythm (periods=12.2-13.7 h) in which they had frequent bouts of swimming to the surface of the column during the expected time of ebb tide in the field and remained on the bottom during the time of flood tide. This rhythm was not present in crabs with young embryos and disappeared after larval release. Thus, an endogenous rhythm in vertical migration does underlie the ebb-tide transport behavior of ovigerous blue crabs with mature embryos during their spawning migration.     

Mechanisms of cross-shelf transport of crab megalopae inferred from a time series of daily abundance

From 20 July 1982 to 19 July 1984, crab megalopae were trapped daily from the Scripps Institution of Oceanography pier. Pachygrapsus crassipes, Portunus xantusii, Cancer spp., Hemigrapsus spp., and Majid crab megalopae dominated the catch. Yearly, seasonal, and daily variations in the magnitude of the catch were observed. Yearly and seasonal variations were probably due to a strong El Niño event that occurred during the study and to the timing of spawning and duration of the larval phase, respectively. Daily variation was correlated with oceanographic processes that can transport larvae to shore. Catch of some taxon during some seasons correlated with wind stress suggesting that transport was wind driven. The correlations were, however, weak and the sign of the correlation varied between years. The maximum daily tidal range was significantly correlated (cross-correlations and cross Fourier analysis) to both daily seawater temperature anomalies (surface and bottom) and daily catch of crab megalopae in all taxa enumerated. Significant correlations between tidal range and temperature anomalies suggest that temperature anomalies were primarily due to the shoreward transport of warm and cold water by the internal tides. The consistent and relatively strong relationship between tidal range and catch of megalopae (the cross-Fourier analysis suggests that from 20 to >90% of the variation in catch can be attributed to variation in the tidal range) suggest that much of the shoreward transport of megalopae was via the internal tides. Shoreward transport of larvae by internal tides may be due to internal cold bores or convergences over large tidally generated internal waves (solitons). Peak catches of megalopae, however, were often not associated with cold anomalies suggesting that transport was due to moving convergences over internal waves.     

Circatidal swimming behavior of brachyuran crab zoea larvae: implications for ebb-tide transport

Larvae of the blue crab Callinectes sapidus and fiddler crab Uca pugilator are exported from estuaries and develop on the continental shelf. Previous studies have shown that the zoea-1 larvae of some crab species use selective tidal-stream transport (STST) to migrate from estuaries to coastal areas. The STST behavior of newly hatched larvae is characterized by upward vertical migration during ebb tide followed by a descent toward the bottom during flood. The objectives of the study were (1) to determine if newly hatched zoeae of U. pugilator and C. sapidus possess endogenous tidal rhythms in vertical migration that could underlie STST, (2) to determine if the rhythms persist in the absence of estuarine chemical cues, and (3) to characterize the photoresponses of zoeae to assess the impact of light on swimming behavior and vertical distribution. Ovigerous crabs with late-stage embryos were collected from June to August 2002 and maintained under constant laboratory conditions. Following hatching, swimming activity of zoeae was monitored in darkness for 72 h. U. pugilator zoeae displayed a circatidal rhythm in swimming with peaks in activity occurring near the expected times of ebb currents in the field. Conversely, C. sapidus zoeae exhibited no clear rhythmic migration patterns. When placed in a light field that simulated the underwater angular light distribution, C. sapidus larvae displayed a weak positive phototaxis at the highest light levels tested, while U. pugilator zoeae were unresponsive. Swimming behaviors and photoresponses of both species were not significantly influenced by the presence of chemical cues associated with offshore or estuarine water. These results are consistent with predictions based on species-specific differences in spawning and the proximity of hatching areas to the mouths of estuaries. U. pugilator larvae are released within estuaries near the adult habitat. Thus, ebb-phased STST behavior by zoeae is adaptive since it enhances export. Selective pressures for a tidal migration in C. sapidus larvae are likely weaker than for U. pugilator since ovigerous females migrate seaward prior to spawning and hatching occurs near inlets and in coastal waters.     

Temporal and spatial use of stranded wrack by the macrofauna of a tropical sandy beach

During two field studies the macrofauna associated with wrack stranded on a tropical sandy shore was analysed. During the first period all animals present in small wrack deposits were randomly collected with cores. During the second field study active animals were sampled in larger wrack deposits for an entire lunar period, and tube traps were used. After a comparison between the two studies, only the data from the tube traps were examined from a temporal and spatial point of view. The aim of the present work was to gain information on how, when and where wrack was colonised by invertebrates, keeping in mind cyclical aspects such as lunar, diel and tidal phases. Predatory taxa, such as Staphylinoidea and the amphipods Talorchestia martensii, were most abundant, whereas Diptera larvae and Tenebrionidae were scarce. Successional changes of beach wrack colonisation were evident throughout a semi-lunar period, with molluscs invading wrack during the first days of deposition and histerids during the last ones. Strictly nocturnal or diurnal surface-active species were found to be active in the wrack during both the day and the night. The analysis of the tidal component of species activity in the wrack showed that some species moved at ebbing tide, and others moved at rising tide. Differences were found also in the mean hours of tidal activity, calculated separately during day and night periods. The study of the zonation of each species showed that in some cases wrack deposits were closely followed by the fauna as their position changed during the semi-lunar phase. In other cases differences occurred between species zonation during day and night periods and tidal phases. Cluster analysis indicates that most species exploit the wrack in different ways (as refuges and/or feeding site), both in space and in time. Received: 11 July 1999 / Accepted: 20 January 2000     

Diel and tidal cycles regulate larval dynamics in salt marshes and mangrove forests

Variation in the release and recruitment of larvae of estuarine invertebrates affects the distribution and abundance of adults, as well as trophic interactions in both the plankton and the benthos. Larval release and supply are often timed to environmental cycles such as the diel and tidal cycles. Here, we determined using plankton tows whether the abundance of larvae spanning salt marsh and mangrove habitats across the intertidal landscape varied with diel and tidal cycles. Using three different sampling designs across two sites and within each of two estuaries over a 12-month period, we covered a range of spatial and temporal scales. This allowed us to test the general prediction that densities of meroplankton in the water column would be greater during nocturnal ebb tides than during other phases of the diel or tidal cycle. As predicted, nocturnal ebb tides yielded the highest densities of meroplanktonic larvae and were dominated by first-stage crab zoeae and this finding was most pronounced in the salt marsh. Throughout the course of the year, greater numbers of meroplankters consistently occurred during the ebb tide compared with the flood tide. The densities of other taxa (e.g. gastropods and polychaetes) showed no clear trends with diel or tidal cycles. This study highlights the effects of these pervasive physical cycles on the timing of larval release and supply in the salt marsh–mangrove complex, and emphasises their contribution to the trophic interactions and the dynamics of benthic populations within estuaries.     

Flow and transport characteristics at an Epinephelus guttatus (red hind grouper) spawning aggregation site in St. Thomas (US Virgin Islands)

This study characterizes the flow field at a spawning site located at the shelf break of a Caribbean island for the Epinephelus guttatus (red hind grouper) in relation to this species spawning events. In order to understand the oceanographic dynamics targeted by the fishes, current measurements were profiled throughout the water column for almost a year at the spawning site. The characteristics of the flow field and its evolution after spawning were investigated by using a numerical ocean model that resolved the observed tide and simulated the island scale flow where passive, neutrally buoyant virtual particles were released for 10 days to trace the flow pathways.Observed currents during the spawning period revealed that the flow was vertically sheared, to the south and weakest at the bottom, and to the west or east at the surface. The tidal analysis revealed that the flow at the time of spawning was directed across and on-shelf, although weaker close to the bottom. The model showed that the initial on-shelf transport was counteracted by the bottom flow directed to the shelf break, where virtual particles were entrained by the downwelling flow. A significant percent of particles resided less than two hundred meters deep, in the vicinity of the chlorophyll maximum and returned to the shelf break, close to the release location within 8-10 days. This journey was largely controlled by the timing between downwelling at the spawning site and upwelling further east at the shelf break, which was driven by the coupling between wind and tide induced vertical movements at the shelf break and deeper. The release location, vertical rotation of its flow field, and its transport properties were shown to be relatively resilient to the passage of transient sub-mesoscale eddies as well as to acute mesoscale flow reversals, suggesting that physical retention is maximized in the area surrounding the spawning site.     

Reproductive cycle,larval development,juvenile growth and population dynamics of Patiriella pseudoexigua (Echinodermata: Asteroidea) in Taiwan

A field study was conducted at Wanlitung, southern Taiwan, in 1986–1089, to determine the reproductive cycle, development mode, growth rate and population dynamics of the small seastar Patiriella pseudoexigua (Dartnall), which occurs in highly stressful and disturbed intertidal pools in this area. An inverse relationship between gonad index and pyloric-caccum index was only recorded immediately prior to spawning. A short, well-synchronized seasonal spawning occurs in October. When reared at 25 °C, lecithotrophic larvae develop directly, lack a bipinnaria stage, and metamorphose completely on the seventh day after fertilization. The growth curves of field juveniles are linear, those of laboratory-reared juveniles are sigmoid. Juveniles appear in tide pools in spring-early summer of each year. Adults spawn mainly in late fall, enabling spawning to occur in time for the larvae to benefit from the environmentally favorable winter season. Populations in high-tidal pools decrease in later summer, but remain more stable in lowtidal pools and lagoons.     

Analysis of lipid components for determining the condition of anchovy larvae,Engraulis mordax

It has been presumed that intertidal spawning by Limulus polyphemus minimizes the loss of egges to subtidal predators; however, this strategy involves considerable risks. Massive beach strandings of adults accompany seasonal spawning migrations of crabs along Cape May in Delaware Bay, (USA). At least 190000 horseshoe crabs, approximating 10% of the adult population, died from beach stranding along the New Jersey shore of Delaware Bay during the 1986 (May to June) spawning season. Abnormalities of the telson (which is used in righting behavior) were significantly more common among stranded crabs than among individuals actively spawning on the intertidal beach. The number of stranded crabs per day was not correlated with tidal height or environmental variables (wind speed, wave height) which characterized the conditions at spawning. A complex suite of factors, including the size of the available spawning population, tidal and weather conditions, and beach slope, influence the number stranded during the breeding season. Horseshoe crab stranding results in a large loss of gravid females from the population, and may represent a major input of organic matter to intertidal sandy beaches in certain regions of Delaware Bay.     

Diurnal and lunar patterns of larval recruitment of Brachyura into a mangrove estuary system in Ranong Province, Thailand

The present study followed the temporal recruitment pattern of brachyuran larvae in a mangrove tidal creek on the Andaman Sea coast of Ranong Province, Thailand, based on the assumption that the processes governing recruitment are important for the overall population dynamics of mangrove brachyuran crabs. Plankton net samples were taken on five occasions: on two new moon spring tides, one waxing moon neap tide, one full moon spring tide and one waning moon neap tide during October and November 1997. In addition collectors for larval crab megalopae were employed every 3 days through one dry season and one wet season (March–October 1998). Both the plankton net samples and collector samples revealed four major brachyuran groups in three families: Ocypodidae, Grapsidae and Portunidae. The grapsid group was further separated into two morphotypes which were identified as Metaplax and sesarmid species. Identified group mean numbers per cubic metre were ocypodids 3.0, sesarmids 0.8 and Metaplax 0.5, while portunid megalopae were very scarce (≪0.1 m⁻³). Further analysis of plankton net samples showed that when considering the parameters date, depth, current direction and the diel cycle, Metaplax and ocypodids distribute differently in the tidal and lunar cycle. Metaplax recruitment dominates on flood tides and on bottom layers, followed by middle and surface layers. Conversely, ocypodid abundance varied significantly with date only. Notably recruitment was not dependent on the diel cycle for either group. The collector samples of megalopae showed that recruitment of ocypodids, Metaplax and sesarmids occurred on full and new moon spring tides, while portunid megalopae preferred to settle on full moon spring tides. Since tidal currents were related to the lunar cycle megalopa groups are also cross-correlated with tidal amplitude, except for the portunid group. It is concluded that megalopae recruit in a similar manner to what has been found in other regions of the world, except that the abundance of ocypodids and Metaplax is not influenced by the diel cycle. Received: 14 February 2000 / Accepted: 24 November 2000     

Larval supply of shorefishes to nursery habitats around Lee Stocking Island,Bahamas. II. Lunar and oceanographic influences

Plankton nets were moored in tidal channels around Lee Stocking Island to monitor larval supply of fishes to nursery habitats that fringe Exuma Sound, Bahamas. Sampling was conducted continuously through two consecutive winters of 1990–1991 and 1991–1992. Meteorological and hydrographic data were also recorded during these periods. Three of the six most abundant taxa (leptocephali, Bothidae and Ophidiidae) showed strong lunar associations, with periodicities between 28 and 30 d. Labrids were the only family to show evidence of any semi-lunar activity. The time series for the families Clupeidae and Tetraodontidae appeared to be stochastic. Cross-correlation analyses were used to examine coupling of larval replenishment and physical processes. Four of six taxa examined showed significant relationships between larval supply and onshore winds at short time lags during 1990–1991. In 1991–1992, however, taxa showed stronger associations with the northwest, long-shore wind component than with onshore winds. Coherency in larval supply among several taxa was remarkably strong. This implies either that the magnitude of spawning is linked among taxa, or that planktonic processes are affecting a number of taxa in very similar ways. The similarities in the magnitude of supply events among taxa found here may be related to complex interactions between larval behavior and the physical environment.     

On predation of pelagic larvae and early juveniles of marine bottom invertebrates by adult benthic invertebrates and their passing alive through their predators

The dynamic quantitative balance between prey and predator invertebrate species inhabiting the same shallow-shelf (sublittoral level bottom) benthic communities was first discussed by Thorson (1953). Thorson considered the exact timing of larval settlement of prey and predator species possessing pelagic development and temporal supression of the adult predators' feeding activities during reproduction at the time of the preys' settlement to constitute the major factors which facilitate survival of the prey species in such communities. However, information obtained demonstrates that Thorson's “mechanism of balance between predator and prey species of benthic communities” is not always effective in securing survival from predation not only of the prey's spat but even sometimes of the predator's spat also. Because of this, the “mechanism” can not be rated as universally effective in all situations. Analysis of the data so far published demonstrates that, in marine benthic communities, especially in shallow-shelf waters, it is not uncommon for gametes, larvae, or early juveniles of different prey species to pass alive through suspension (filter)-feeding and deposit-feeding adult invertebrates preying on them. Sometimes development can even continue after excretion by predators. The hypothesis of Voskresensky (1948) and Goycher (1949) of the importance of this phenomenon for the maintenance and recruitment of the mussel Mytilus edulis and other filter-feeding lamellibranchs of nearshore waters preying on their own and other lamellibranch pelagic larvae must be rejected on the basis of accumulated data on their feeding and general biology and on the adverse influence of the mucous of their faecal pellets and pseudofaeces on the larvae excreted by them alive. The data considered here demonstrate that, although the passing alive of larvae and spat of benthic invertebrates through benthic predators is not uncommon in shallow-shelf bottom-communities, it plays no important role in the processes of maintenance and recruitment of the species and communities involved nor of the marine benthos as a whole. The actual ecological significance of predation on pelagic larvae and bottom spat of benthic invertebrate prey species by all three main trophic groups of marine benthos (suspension or filter-feeders, deposit-feeders, carnivores) and its importance to predator-prey dynamics in marine benthic communities remains open to debate until more reliable quantitative data become available.     

Endogenous swimming rhythms underlying the spawning migration of the blue crab, Callinectes sapidus: ontogeny and variation with ambient tidal regime

Spawning female blue crabs, Callinectes sapidus, use ebb-tide transport (ETT) to migrate seaward. In estuaries with semi-diurnal tides, ETT in ovigerous blue crabs is driven by a circatidal rhythm in vertical swimming in which crabs ascend into the water column during ebb tide. The ontogeny of this rhythm was examined by monitoring swimming behavior of females before the pubertal molt, females that had recently undergone the molt but had not yet produced a clutch of eggs, and ovigerous females from an estuary with strong semi-diurnal tides. To assess variation in swimming rhythms with ambient tidal regime, swimming rhythms of ovigerous females from semi-diurnal (Beaufort, North Carolina), diurnal (St. Andrew Bay, Florida), and non-tidal (South River, North Carolina) estuaries were compared. Experiments were conducted during the summers of 2006–2008. Female crabs prior to oviposition had variable endogenous swimming rhythms (circadian, circatidal, or circalunidian). Ovigerous females from estuaries with semi-diurnal and diurnal tides had pronounced circatidal or circalunidian rhythms with swimming during the time of ambient ebb tide. Swimming rhythms of several ovigerous crabs switched between circatidal and circalunidian during the ~5-day observation period. Ovigerous crabs from a non-tidal estuary had a circadian rhythm with vertical swimming around the time of sunset. These results suggest that, while endogenous swimming rhythms are present in some female blue crabs prior to oviposition, rapid seaward movement via ETT in tidal estuaries begins following oviposition of the first clutch of eggs.     

Persistent locomotor rhythmicity in the prawns Penaeus indicus and P. monodon

The locomotory activity of adult Penaeus indicus Milne Edwards and P. monodon (Fabricius) collected from the Vellar estuary, South India, in 1983 was monitored continuously for a period of 40 d in the laboratory. Both species exhibited persistent circatidal and circadian rhythmicity, the former displayed peak activity during predicated times of high water in their original habitat. The latter exhibited peak activity during periods corresponding to in situ night-time, with almost no activity during the day, except at the time of high tide. The tidal and daily rhythms were evident for only 3 to 4 d and 5 to 6 d, respectively, under laboratory conditions. The amplitude of the rhythms displayed variations in each semilunar period, reflecting the synchrony and dissynchrony of the tidal and daily rhythms. Activity was correlated with phases of the moon, peaking at times of new and full moon, and showing a scattered pattern with smaller peaks during the first and third quarters.     

Endogenous swimming rhythms in estuarine crab megalopae: implications for flood-tide transport

Up-estuary migration of crab larvae to adult habitats is thought to be accomplished by selective tidal transport in which late-stage larvae enter the water column on flood tides and remain on or near the bottom on ebb tides. This study measured endogenous rhythms in swimming by the last larval stage (megalopa) of blue crabs Callinectes sapidus and fiddler crabs Uca spp. Previous field studies found that megalopae of both species were only abundant in the estuarine water column on nocturnal rising tides. Megalopae were collected from the Newport River Estuary, North Carolina (34°41N; 76°40W) during August–September 1992 and swimming activity was recorded for 4.5 to 7 d under constant conditions with a video system. Rhythms exhibited by both genera in the laboratory were not identical to those recorded in the field. Uca spp. displayed a circatidal rhythm, with maximum swimming occurring near the time of high tide in the field. Rhythm amplitude increased when crushed oyster shells were present, which suggested that megalopae bury or cling to the substrate during quiescent periods. In contrast, C. sapidus had a circadian rhythm in which maximum swimming coincided with the day phase in the field. In most trials, the activity of blue crab megalopae was unrelated to the expected tidal cycle. It was concluded that a tidal rhythm in swimming was the behavioral basis of flood-tide transport for fiddler crab larvae. The endogenous rhythm in blue crabs does not participate in transport, which probably results from behavioral responses to environmental cues associated with flood tide.