Looking for the clock mechanism responsible for circatidal behavior in the oyster Crassostrea gigas

Valve activity rhythm of the oyster Crassostrea gigas is mainly driven by tides in the field, but in the laboratory, only a circadian clock mechanism has been demonstrated. In an attempt to reconcile these results, the mechanisms underlying the circatidal rhythm were studied in the laboratory under different entrainment or free-running regimes and in the field at Arcachon (44°39′N/1°09′W) in February–April 2011). Results confirm the existence of a circadian clock in C. gigas. Under entrainment regimes (12-h dark/12-h light photoperiod and tidal cycles simulated by a reversing current flow), oysters exhibited both circadian and circatidal cycles. Under free-running conditions (e.g., continuous darkness), the endogenous rhythm appeared to be circadian. There was no experimental evidence for an endogenous circatidal rhythm, even in oysters just transferred from the field, where a clear tidal cycle was expressed. There are two possible mechanisms to explain tidal behavior in C. gigas: an exogenous tidal cue that drives tidal activity and masks the circadian rhythm and an endogenous circatidal clock that is sensitive to tidal zeitgebers and runs at tidal frequency.     

Circatidal rhythm of an intertidal crab, Hemigrapsus sanguineus : synchrony with unequal tide height and involvement of a light-response mechanism

The circatidal rhythm of intertidal animals may reflect the inequality of the tides. In addition, a light-sensitive mechanism may be involved in their internal timing systems. To test these hypotheses, the larval release activity of the intertidal crab Hemigrapsus sanguineus was monitored under different light conditions in the laboratory. Under a 24-h light–dark (LD) cycle with the phase similar to the field, the activity coincided with the times of high tide in the field and showed a tidal rhythm. This rhythm free-ran in constant, dim-light conditions, suggesting that the timing is controlled by an endogenous clock. When the population was exposed to a 24-h LD cycle with the phase changed from that in the field, the tidal rhythm was phase-shifted; while the light cycle advanced in phase from the field caused a phase-advance of the rhythm, that delayed in phase induced a phase-delay of the rhythm. Thus, a light-response mechanism is definitely involved in circatidal timing systems. But the population rhythm showed a large variability among individuals, associated with the phase-shift, and the magnitude of the phase-shift did not accurately correspond to that of the light cycle. These results suggest that the light-response system can control the phase of the rhythm less stronger than that in estuarine crabs. Most releases occurred at higher high tides, but the release of some females obviously occurred at lower high tides. The larval release pattern thus could not be accounted for by a simple synchrony with higher high tides. Hatching of H. sanguineus occurred after a “hatching program” of 49.5 to 52.5 h. This program is initiated by some factor (hatching-program inducing stumuli: HPIS) transmitted from the female to the embryos. We speculated that this factor is effectively transmitted to the embryos when the habitat is exposed to air, i.e., at lower low-tide periods, and that once each embryo is stimulated, hatching occurs synchronously 2 d later during high tide. The release of HPIS is probably controlled by the circatidal clock of the female, and the 24-h LD cycle may participate in shifting this timing to the opposite low tide. Received: 14 January 1997 / Accepted: 18 February 1997     

Rhythmic activity responses of the fiddler crab Uca crenulata to artificial tides and artificial light

The fiddler crab Uca crenulata, from California (USA), was exposed to artificial tides in order to differentiate between the influence of a tidal cycle from that of a light-dark cycle on its locomotor activity. Most crabs could be well synchronized by tides, but the activity patterns of other crabs was merely exogenously reinforced. Under constant conditions, after tidal treatment, crabs showed bimodal or unimodal activity patterns. When a 24 h light-dark cycle and a 12.35 h tidal cycle acted as concurrent stimuli, their effectiveness in synchronizing the crabs' activity was variable, depending on the undividual. It is likely that crabs which tend to exhibit a unimodal activity pattern are more sensitive to a light stimulus, whereas crabs with a bimodal activity pattern respond preferably to the tides. Thus, responses of endogenous activity of U. crenulata to tides and light-dark cycles are similar, suggesting that the basic oscillarory mechanism for circadian and tidal activity may be the same.     

Relationships between intertidal zonation and circatidal rhythmicity in littoral gastropods

The presence and phase of circatidal rhythmicity was correlated with vertical zonation and other ecological factors. Ten species were studied in the field and in aktographs under controlled conditions in the laboratory. Retina plicata, Melanerita atramentosa, Bembicium nanum, Austrocochlea obtusa and Morula marginalba occupied mid- or upper-littoral zonations, and were subjected to regular tidal influence. They were active at high water and for a period after the ebb, possibly as these were the times of least desiccation. Each possessed a circatidal activity rhythm under non-tidal conditions. Midlittoral Amphinerita polita and lower-littoral Theliostyla albicilla were active at low water, possibly because their prefered habitats reduced desiccation and other selective forces, for example wave action and predation, determined the phase of the rhythmicity. Supra- and upper-littoral Nodilittorina pyramidalis and Melarapha unifasciata and lower- and infralittoral specimens of Bembicium auratum displayed no overt circatidal rhythmicity, possibly because they were not subjected to regular tidal action.     

Circannual variations of lizard circadian activity rhythms in constant darkness

The daily pattern of locomotor activity of Podarcis sicula in the field changes from unimodal in spring to bimodal in summer, becoming unimodal again in autumn. Short-term experiments in which P. sicula collected in different months were tested under constant conditions immediately after capture showed that the activity pattern typical of each season is retained in the lizard circadian locomotor rhythm. In constant conditions, the bimodal pattern is associated with a short free-running period (τ) of the circadian locomotor rhythm and a long circadian activity (α), while the unimodal pattern is associated with a long τ and short α. To test whether seasonal changes in circadian locomotor rhythms are driven by a circannual clock, we recorded locomotor activity of lizards over 12–15 months in constant temperature and darkness. The present results demonstrate, for the first time in a vertebrate, the existence of circannual changes in constant darkness of both τ and α. In most lizards, the longest τ along its circannual cycle is associated with a short α, and the shortest τ in the same cycle with a long α, so that the pattern of mutual association between τ and α is the same as in short-term experiments. Most lizards, however, stayed unimodal all the time. This shows that changes in activity pattern from unimodal to bimodal (and vice versa) are induced by seasonal changes in environmental factors, instead of being incorporated into a circannual rhythm. Circannual changes in τ and α of locomotor rhythms may adaptively predispose the circadian system of P. sicula to a change in activity pattern as soon as seasonal changes in the environment demand it. Received: 22 January 1999 / Received in revised form: 14 April 1999 / Accepted: 19 April 1999     

External synchronizers of tidal activity rhythms in the prawns Penaeus indicus and P. monodon

Phasing of persistent circatidal rhythmicity to an artificial tidal cycle was assessed in the prawns Penaeus indicus Milne Edwards and P. monodon (Fabricius) collected from the Vellar estuary, South India, in the period between June and December 1984. Simulated 6 h cycles of 20 and 30 S, and 6 h cycles of 20° and 30°C induced a persistent tidal rhythmicity after 20 cycles. The imposed 6 h cycles of 25 and 30 S, and 25° and 30°C induced tidal rhythms after 30 cycles. In both cases, re-established tidal activity rhythms were evident for at least 48 h — higher activity occurring during the higher salinity and lower temperature phases of the simulated tidal cycles. Artificial tidal cycles of still water and running water synchronized the tidal rhythm after 20 cycles. Combined 30 S, 20°C, for 6 h and 20 S and 30°C for 6 h established a persistent tidal rhythm after 10 cycles, whereas wave action had no influence on tidal synchronization. The influence of possible interactions of tidal rhythms and in situ tidal variables on circatidal activity is discussed.     

Seasonal changes of locomotor activity patterns in ruin lizards Podarcis sicula

The daily pattern of locomotor activity of the ruin lizard Podarcis sicula in its natural environment changes from unimodal in spring (with only one activity peak per day) to bimodal in summer (with two well-separated activity peaks per day) and it becomes unimodal again in autumn. In order to establish whether such seasonal changes in pattern might be at least in part controlled by endogenous temporal programs, lizards were collected at different times of the year and immediately after capture their locomotor behavior was tested in the laboratory under constant temperature (29°C) and in darkness. For some individuals tested in the laboratory the locomotor pattern previously expressed in the field was known. Seasonal differences in pattern have been unequivocally found to have an endogenous component, as most lizards in constant conditions retained the locomotor pattern shown in the field during the same season. Besides, in the bimodal lizards the freerunning period of locomotor rhythms () was significantly shorter and circadian activity time (a) longer than in the unimodal ones. Altogether the data are compatible with the idea that both the interdependent changes of and a and the changes in locomotor pattern occurring seasonally in the circadian activity rhythms of P. sicula would depend on changes in the phase relationship between mutually coupled oscillators which drive these rhythms. Correspondence to: A. Foà     

Relationship of daily and circatidal activity rhythms of the fiddler crab,<Emphasis Type="Italic"> Uca princeps</Emphasis>, to the harmonic structure of semidiurnal and mixed tides

Intertidal organisms may employ circatidal rhythms to track the tidal cycle, but tidal patterns may vary within a species range and necessitate adaptation to the local tides. Circatidal rhythms were examined in populations of the eastern Pacific fiddler crab Uca princeps (Smith) from four sites with differing tidal characteristics, La Paz (24°10N; 110°21W), San Blas (21°33N; 105°18W) and Manzanillo (19°6N; 104°24W), Mexico (lower amplitude, mixed semidiurnal tides) and Mata de Limon, Costa Rica (9°55N; 84°43W) (high-amplitude, semidiurnal tides). Local tides were characterized by harmonic constants of M₂, S₂, K₁, and O₁, partial tides that largely determine their semidiurnal and diurnal features. Rhythmic structure in continuously recorded locomotor activity of individual crabs held under laboratory conditions was described by cosinor and periodogram methods of time-series analysis. Both daily and circatidal rhythms were found in crabs studied in light–dark cycles set to local conditions at the time of collection. Crabs at all four sites shared a tendency toward bimodality, with a mid-morning activity peak and varying degrees of nocturnal activity. Circatidal rhythms closely matching the period of the 12.42-h M₂ partial tide were consistently present at all sites except Manzanillo. At Mata de Limon, the circatidal rhythm clearly dominated locomotor activity, but was strongly modulated by a daily rhythm in a repeating pattern at a semilunar interval. In contrast, the amplitude of the daily rhythm was higher than that of the circatidal rhythm in crabs from the three mixed tide sites on the Mexican coast, where the tidal pattern is dominated by a diurnal inequality arising from the diurnal K₁ and O₁ partial tides. These results suggest that populations of U. princeps use both daily and circatidal timing systems to track local forms of the tide generated by their M₂, S₂, K₁, and O₁ geophysical counterparts.Communicated by J.P. Grassle, New Brunswick     

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.     

Rhythmic motor activity responses of the California fiddler crab Uca crenulata to artificial light conditions

Uca crenulata, the California species of fiddler crab, was exposed to artificial light conditions to separate the influence of the light cycle from that of the tidal input on its rhythmic motor activity. Under both constant light and light-dark cycles, rhythmic activity was demonstrated in only 50% of the experimental crabs; the activity of the remaining 50% was random. Individuals exposed to constant light conditions after having been subjected to 24 h light-dark cycles demonstrated no significant difference in period length of their rhythmic activity from crabs investigated in constant light immediately after field collection. The mean period did not differ significantly from the tidal period of 24.8 h, but the variation was considerable. In artificial light-dark cycles, the activity rhythms were usually masked but, in some cases, synchronized. The results indicate that U. crenulata has an endogenous rhythm with a period close to the tidal cycle which may be synchronized by light as well as by tidal cues. The display of this endogenous rhythm, however, is poor.     

Endogenous swimming rhythms of blue crab, Callinectes sapidus , megalopae: effects of offshore and estuarine cues

Larvae of the blue crab Callinectes sapidus Rathbun develop on the continental shelf. The postlarval stage (megalopa) occurs near the surface and is transported shoreward by wind-driven surface currents. It then uses selective tidal stream transport for migration up an estuary. Endogenous swimming rhythms were measured under constant dark conditions in the laboratory in megalopae collected from the Newport River Estuary (North Carolina), the Delaware Bay, and offshore from the Newport River Estuary. Megalopae from all areas had a similar circadian activity rhythm, in which they swam during the time of the day phase in the field and were inactive at night. This rhythm predicts the presence of a reverse, diel, vertical-migration pattern offshore which would contribute to the location of megalopae near the surface during the day. The rhythm lacks obvious ecological significance in estuaries because it does not contribute to selective tidal stream transport and would increase vulnerability to visual predators during the day. Attempts to entrain a circatidal rhythm in swimming by cyclic and step changes in salinity were unsuccessful, as the circadian rhythm persisted. The rhythm also continued in the presence of the eelgrass Zostera marina, which is a site of settlement and metamorphosis in the field. Thus, megalopae enter estuaries with a solar day rhythm in activity. This rhythm, however, is not expressed, because light inhibits swimming during the day upon exposure to estuarine water. Since this light inhibition is removed in offshore waters, the rhythm would be expressed if, after entering an estuary, megalopae were transported back to offshore areas. Received: 19 December 1995 / Accepted: 2 August 1996     

Gezeitenrhythmische Nahrungsaufnahme und Kotballenablage einer terrestrischen Milbe (Oribatei: Ameronothridae) im marinen Felslitoral

Feeding, locomotory and defecatory activity of the algivorous orbatid mite Ameronothrus marinus have been measured under defined conditions in the laboratory. Of terrestrial origin, these mites inhabit the intertidal zone of rocky shores along the North and Baltic Seas. Under daylight cycles found at the sampling site and in the absence of tides, feeding and defecation follow an endogenous rhythm with a spontaneous period of 12.3 h; in the tideless littoral of the Baltic Sea these activities are synchronized with diurnal cycles. Feeding and defecation follow a definite sequential pattern, repeated on the average every 4 h, i.e., 3 times between two high tides. The phase of this tidal feeding-defecation rhythm is determined by the onset of the high tide, and different patterns of this rhythm can be observed according to the tidal zones. Inhabitants of the lower eulittoral exposed to longer periods of submersion start feeding earlier, take up more food at one time before the beginning of high tide, quickly defecate part of the food undigested after feeding, and later produce other faecal pellets which contain the real remains of digestion. Inhabitants of the upper eulittoral, exposed to shorter periods of submersion, start feeding later and extend their feeding activity over the whole period of low tide; the difference between faecal pellets with digested and undigested contents is indistinct. The occurrence of these different types of faecal pellets in inhabitants of the intertidal zone is interpreted as a compensatory physiological adaptation resulting from increasing periods of submersion.     

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.     

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.     

The tidal rhythm of extracellular digestion and the response to feeding in Ostrea edulis

Under natural conditions, on the shore, there is a tidal rhythm for changes in pH, length, and protein and amylase content of the crystalline style of Ostrea edulis L. When oysters were kept immersed and fed continuously for 2 weeks, in the laboratory, the rhythm of extracellular digestion was lost. Oysters were fed discontinuously for 2 weeks, in the laboratory, with a 6 h-on, 6 h-off feeding regime. During the feeding period, the changes in pH, size, and protein content of the style were similar to the changes observed in the field over the period of high tide. It is our hypothesis that the tidal rhythm of extracellular digestion in Ostrea is not endogenous, but is controlled by feeding activity.     

Endogenous circatidal rhythm in the Manila clam Ruditapes philippinarum (Bivalvia: Veneridae)

Manila clams, Ruditapes philippinarum, removed from their natural environment and maintained for 9 weeks in continuously immersed conditions exhibited a clear endogenous circatidal rhythm in oxygen consumption. The clams exhibited a semidiurnal rhythmicity in oxygen consumption after showing a diurnal pattern in the first few days (5 to 7 d) of the experiment. The results of the present study indicate that activity rhythms of clams are controlled not only by exogenous factors, but also by an endogenous circatidal periodicity. Received: 8 April 1998 / Accepted: 1 February 1999     

Activity rhythms of two cirolanid isopods from an exposed microtidal sandy beach in Uruguay

Activity rhythms of two cirolanid isopods, Excirolana armata and Excirolana braziliensis, were studied based on both seasonal field observations and laboratory experiments, at an exposed microtidal sandy beach in Uruguay. The natural emergence patterns of both species were observed in the field for 1 year, twice in each season, and correlated to sea level, expected tidal cycles and diel cycles. Laboratory experiments were carried out in order to detect endogenous rhythms of activity and observe how emergence of both species was affected by changes in light and/or sediment thixotropy. We also compared behavioral strategies of sympatric species that occupy different beach levels. Sea level (and thus swash zone position) during field sampling did not follow expected tidal cycles for most sampling occasions. E. armata was observed in activity most of the time, but activity only correlated with sea level on three out of eight occasions, and only once was correlated to expected tidal cycle. Laboratory results showed that emergence under constant conditions was rare; changes in sediment thixotropy stimulated emergence, but the response was not cyclical; light had little effect on this response. On the other hand, E. braziliensis was fairly scarce in the water column, but swimming individuals were observed always during the night. They displayed an endogenous circadian activity pattern in the laboratory which augmented in response to changes in sediment thixotropy. The natural light/dark cycle modulated both spontaneous and response emergence by increasing day/night differences in activity. In this study E. armata, a midlittoral species more exposed to sea level variations, seemed to rely entirely on different physical and/or biological cues to trigger emergence at the appropriate time. E. braziliensis, found mostly in the upper intertidal zone, emerged in a circadian rhythm, which was stimulated by changes in sediment thixotropy and reinforced by light cycles. The results of this study led us to conclude that on microtidal, unpredictable beaches, local physical and biological factors can combine to determine different activity strategies in organisms from different intertidal levels. Received: 23 March 2000 / Accepted: 30 August 2000     

Occurrence of lysozyme in the common cockle Cerastoderma edule and the effect of the tidal cycle on lysozyme activity

In 1983 a sample of cockles [Cerastoderma edule (L.)] was removed from the mid-tide level at Booterstown Strand, County Dublin, Ireland at 2-h intervals, for a period of 24 h. The gills, mantle and visceral mass tissues contained significant levels of lysozyme, with the gills containing the highest, weight-specific levels. A tidal rhythm of lysozyme activity in the gills, mantle and visceral mass was inversely related to the tidal cycle. The data were best described by a tidal curve shifted-5 hours out of phase, with maximum and minimum levels of activity occurring one hour after low and high tides, respectively. In marine bivalves, lysozyme may be involved in the extracellular digestion and absorption of bacteria during low tide. In this context the present use of bivalves as biological indicators of bacterial pollution is discussed.Constribution No. 6282 from the Woods Hole Oceanographic Institution     

Biological and physical aspects of migration in the estuarine amphipod Gammarus zaddachi

The amphipod Gammarus zaddachi (Sexton) conducts extensive migrations along estuaries from near the limit of tidal influence in winter to more downstream reaches (where reproduction occurs) in spring. A return migration then takes place, primarily by juveniles, until the seaward areas are depopulated in winter. The present study was conducted between 1988 and 1990 in the Conwy Estuary, North Wales. This represents the first investigation on this species in a strongly tidal estuary, where the amphipods appear to migrate vertically into the water column on flood or ebb tides to control horizontal transport and to maintain preferred distributions. The timing of vertical migration seems to be largely controlled by an endogenous circatidal swimming rhythm. Phasing of peak activity relative to the time of expected high tide varies with season; upstream migrants in the autumn showed peak activity at the time of expected high tide, while in the spring at the time of downstream migration the rhythm was phase-delayed, with peak activity during the expected ebb tide. Together with the season, position along the estuary also affected the timing of peak endogenous activity; downstream migrants, originally active on the ebb tide and experimentally displaced seawards, showed a phase-advance of the rhythm relative to the time of high tide. Salinity-preference behaviour also varied between different developmental stages, with ovigerous females (downstream migrants) showing no preference between fresh and saline water, and juveniles (upstream migrants) showing a significant preference for freshwater. The interactions of endogenous rhythmicity and salinity-preference behaviour are discussed as controlling factors of migration in this species.     

Rhythms in larval release of the sublittoral crab Neopanope sayi and the supralittoral crab Sesarma cinereum (Decapoda: Brachyura)

Ovigerous females of the subtidal xanthid crab Neopanope sayi (Smith) and the high intertidal grapsid crab Sesarma cinereum (Bosc) were collected during the summers of 1986 and 1987 in the Beaufort, North Carolina (USA), area and brought into the laboratory, where rhythms in larval release were monitored. When crabs with late-stage embryos were put under a 14 h light:10 h dark cycle in an otherwise constant-environment room, an apparent tidal rhythm in release of larvae was observed for both species, with N. sayi releasing near the time of day and night high tides, and S. cinereum releasing around the time of night high tides. The time of sunset relative to high tide was a complicating factor, since larval release for both species was often concentrated around sunset when evening high tides fell several hours before sunset. When a group of N. sayi and S. cinereum were brought into the laboratory and placed under constant lowlevel light for 5 d, the release rhythm of the population persisted, thus implying that the rhythm is endogenous. Larval release near the time of high tide and often at night is common among brachyurans living in tidal areas, regardless of specific adult habitat, suggesting a common functional advantage. Possibilities include transport of larvae from areas where predation and the likelihood of stranding and exposure to low-salinity waters are high, as well as a reduced probability of predation on adult females. Results of the present study suggest that the importance of release after darkness may increase with increasing tidal height of the adult.