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     

The contribution of seagrass beds (<Emphasis Type="Italic">Zostera noltii</Emphasis>) to the function of tidal flats as a juvenile habitat for dominant,mobile epibenthos in the Wadden Sea

Since the substantial loss of subtidal eelgrass (Zostera marina L.) in the 1930s, seagrass beds in the Wadden Sea are limited to the intertidal zone and dominated by Z. noltii Hornem. This study deals with the effect of vegetated tidal flats on quantities of mobile epifauna and proves empirically the function of seagrass canopies as a refuge for marine animals remaining in the intertidal zone at ebb tide. Drop-trap samples were taken in the Sylt-Rømø Bight, a shallow tidal basin in the northern Wadden Sea, on vegetated and unvegetated tidal flats during July and August 2002, and during the entire growth period of Z. noltii from May to September in 2003. The species composition in Z. noltii and bare sand flats showed minor differences since only two isopod species (Idotea baltica and I. chelipes) occurred on Z. noltii flats exclusively. Juvenile shore crabs (Carcinus maenas L.), brown shrimps (Crangon crangon L.) and common gobies (Pomatoschistus microps Krøyer) were also found abundantly on bare sand flats. However, the results showed significantly higher abundances and production of these dominant species on vegetated tidal flats. Additionally, the analyses of faunal size classes indicated higher percentages of small individuals in the seagrass bed during the entire sampling period. Despite drastic diurnal fluctuations of dissolved oxygen at low tide, faunal density in the residual water layer remaining in seagrass canopies at ebb tide was found to be consistently higher than that found in artificially created tide-pool units. Although species composition of mobile epifauna did not basically differ between vegetated and unvegetated tidal flats, Z. noltii beds are considered to contribute quantitatively to the function of tidal flats, as an extended juvenile habitat for some of the most important species of the Wadden Sea food web.Communicated by O. Kinne, Oldendorf/Luhe     

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     

Zonation of the interstitial polychaeteStygocapitella subterranea (Stygocapitellidae) in European sandy beaches

The zonation ofStygocapitella subterranea Knoellner (Polychaeta, Stygocapitellidae) in sandy beaches is compared for several localities along the European coasts. In tidal beaches the species occupies a zone near average high tide level (AHTL), whereas in non-tidal beaches it is found at a fairly long distance from the water line. The regularities observed inS. subterranea indicate that the interstitial fauna may prove well suited for the establishment of a general scheme of zonation in sandy beaches analogous to that for rocky shores proposed byStephenson andStephenson (1949).     

Niche segregation in sandy beach animals: an analysis with surface-active peracarid crustaceans on the Atlantic coast of Spain

Several species of sandy beach invertebrates regularly switch between burial in the sand during the day and surface activity at night to feed on stranded organic matter. Because all species consume essentially the same type of food (i.e., wrack) deposited over a restricted area, the potential for competition exists. Conversely, spatial and temporal segregation of surface activity behaviour is predicted to allow for niche separation. Here, we tested whether such behavioural niche separation occurs in three species of sympatric crustaceans (the talitrid amphipods Talitrus saltator (Montagu 1808) and Talorchestia brito Stebbing 1891, and the oniscoidean isopod Tylos europaeus Arcangeli 1938). We also assessed whether surface activity is modulated by weather, sea conditions and moon phases. Surface activity was measured over three consecutive moon phases on an exposed beach on the Atlantic Coast of Spain using pitfall traps along three transects from the foredunes to the swash. Adults of the amphipod T. saltator and the isopod T. europaeus overlapped spatially and temporally in their surface activity for most of the night. By contrast, the activity of T. brito was strongly disjunct in time and concentrated into significantly shorter bouts around dawn or dusk. Niche differentiation was also evident along the space axis, where T. brito occurred distinctly lower on the beach. Although several environmental factors (e.g., air and sand temperature, humidity, atmospheric pressure, wind speed and wind direction) were correlated with surface activity, this was highly variable amongst species, life stages, and synodic phases, and did not obscure the fundamental mosaic of spatio-temporal heterogeneity amongst species. Nocturnal feeding on beach wrack creates the potential for competition amongst sympatric crustaceans which can, however, be mitigated by behaviour patterns that separate consumers in time and space.     

Modeling the interaction between tides and storm surges for the Taiwan coast

An unstructured grid, two-dimensional hydrodynamic model was established and applied to the coast of Taiwan to investigate the tide-surge interaction. Tidal elevations at the open boundaries coupled with a global ocean tidal model and the meteorological conditions using a cyclone model are used to drive the model. The model was calibrated and verified with the observed tidal levels at six tidal stations for seven typhoon events to ascertain the capability and feasibility of the model. The results show reasonable agreement between the simulated and observed tidal levels. The validated model was then applied to probe the influence of tide-surge interaction on phase, water levels, and storm surge height. We found that the tide-surge interaction influenced both the magnitude and timing of the surge, which depended on the typhoon path. The storm surge heights at different tidal stations were significantly influenced by wind stresses and directions. The water level rise due to the storm surge during high tide was greater at neap tide than at spring tide. Changing tidal ranges altered the prediction of the surge enough to induce the changes in peak water levels.     

Egg release and settlement patterns of dioecious and hermaphroditic fucoid algae during the tidal cycle

The timing of gamete release by fucoid algae, although known to be restricted to calm days is not clearly understood within a circadian time scale. The need for externally fertilizing species to avoid gamete dilution suggests that in wave-exposed areas spawning may occur during particular tidal phases. However, this may differ between mating systems, as selfing species may be less affected by gamete dilution. In this study, two different approaches were used to determine when egg release occurs during the tidal cycle in two sister species with different mating systems. First, egg settlement of Fucus vesiculosus (dioecious) and Fucus spiralis (selfing hermaphrodite) was quantified on removable substrates (egg settlement disks) every day for 2 months and settlement patterns were used to statistically estimate the radius of a circle that would encompass 99% of each patch of settled eggs (the egg dispersal radius). Also, egg release was quantified every 2 h during the tidal cycle. A significantly larger egg dispersal radius (P < 0.02) was found for F. spiralis than F. vesiculosus, and this difference was somewhat site dependent with a greater difference between species in exposed sites. The egg dispersal radius was negatively correlated with significant wave height and positively correlated with sea surface temperature for both the species (P < 0.05), with a greater effect of both the factors for F. spiralis than for F. vesiculosus. Egg release during the tidal cycle was variable between species and experiments, with F. vesiculosus releasing more eggs, later in the day, and at a lower tide, than F. spiralis, which released fewer eggs, throughout the day and at all tides. The dioecious species, F. vesiculosus, may have developed a specific adaptation for timing the egg release to periods when emersed in exposed habitats to avoid rapid dilution of gametes that require outcrossing for fertilization. On the other hand, egg release for F. spiralis, which can self-fertilize, occurred both when emersed and immersed, suggesting this species has developed less synchrony with specific environmental factors. Site dependence also suggests local wave conditions can modulate timing of release. The data are consistent with the relaxation of the selective constraints of water motion on fertilization success in a selfing hermaphrodite, relative to an obligate outcrossing species. Results support the idea that species with different mating systems evolve different sensitivities to environmental cues for gamete release with specific implications for inbreeding and successful external fertilization in the ocean.     

Monitoring spatio-temporal variation in molluscan grazing pressure in seasonal, tropical rock pools

In Hong Kong, low-shore rock pools support a patchy mosaic of different successional stages of crustose coralline algae. Echinoid and molluscan grazers are abundant in these low-shore rock pools and spatial and temporal variation in grazing pressure is thought to play an important role in structuring these assemblages. Direct records of graze marks using a modified wax-disc method were used to monitor spatial (different pools) and temporal (periods, 3 or 4 days each; spring vs. neap tides; summer vs. winter) variations in grazing pressure. Radula marks of Chlorostoma argyrostoma, Lunella coronata and Nerita albicilla were common in all pools. Grazing pressure was highly variable with tidal state and season but also within these two factors. Lunella and Nerita exerted highest grazing pressure at neap tides during summer, whilst Chlorostoma exerted highest grazing pressure in winter. Although overall patterns of temporal variation were similar and consistent in all pools, there was between pool variation in grazing pressure and individual species showed different patterns. Within the seasonal and tidal variations, large period-to-period differences were found for all species, illustrating small-scale temporal variation. Fine-scale spatial and temporal patterns of variation in grazing pressure of different species, therefore, contribute to the highly dynamic nature of these low-shore rock pools.     

Nocturnal feeding migrations of Nerita plicata,N. undata and N. textilis (Prosobranchia: Neritacea) on the rocky shores at Mkomani,Mombasa, Kenya

The vertical zonation of the three common rocky shore neritids at Mkomani, Mombasa, Kenya, Nerita plicata Linnaeus, N. undata Linnaeus, and N. textilis Dillwyn, as a function of feeding migrations and of size, was studied from 28 February to 24 March 1983. These snails perform feeding migrations at night starting at around mid-ebb tide and return to their resting positions with the flood tide. They remain in their resting positions throughout the day until the next nocturnal ebb tide. The direction of migration is sizerelated, with the larger snails of each species moving in the opposite vertical direction to the smaller ones, so that the populations as a whole exhibit no statistically significant net vertical displacement. The larger individuals of two of the species, N. plicata and N. undata, invariably move downwards to their feeding levels, while the smaller individuals move upwards; the larger individuals of N. textilis display a different pattern of migration, moving downwards on and around spring-tide days and upwards on and around neap tide days, while the smaller individuals move in the opposite directions. N. textilis rest above their feeding level around spring tides, and below that level around neap tides. It is demonstrated how these nocturnal migratory feeding rhythms are integrated into the spring-neap and seasonal cycles of the snails' daytime resting positions. The adaptive significance of these migrations is also discussed.     

Feeding and swimming of lysianassid amphipods in a shallow cold-water bay

The potential for dispersal by lysianassid amphipods and their localization to carrion in a shallow cold-water bay in the Middle Saint Lawrence Estuary were assessed by means of endobenthic sampling, SCUBA observations, measures of swimming speeds, and by exposure of bait (50–100 g of fish) in traps. Seventy-five to 99.9% of animals attracted to traps were lysianassid amphipods belonging to five species. Lysianassid species were spatially segregated in the Bay at low tide but all were more or less dispersed at high tide. Second cohortAnonyx sarsi Steele and Brunel,Boeckosimus edwardsi andOnisimus littoralis (Krøyer) were more dispersed than the small first cohort individuals. Second cohortA. sarsi were crawlers or low (0–0.5 m off the bottom) suprabenthic swimmers in the day, but upper (0.5–2 m) suprabenthic swimmers at night. In contrast, first cohortA. sarsi were crawlers or low suprabenthic swimmers day-and-night, whileOrchomenella pinguis (Boeck) followed this swimming pattern at night but were generally akinetic in the day. Mean swimming speeds ofA. sarsi (13.6 cm s^-1) andOn. littoralis (12.1 cm s^-1) were 2 to 3 times greater than those ofOr. pinguis (7.4 cm s^-1) andPsammonyx nobilis (Stimpson) (4.4 cm s^-1). Catchability coefficients (i.e. ratio number of individuals per trap:endobenthic abundance) were 74 (A. sarsi), 8 (On. littoralis), 7 (Or. pinguis), and 0.7 (P. nobolis) m² of bottom. Gut content analysis indicated thatA. sarsi fed mostly on large carrion, whileOn. littoralis were markedly opportunistic, andOr. pinguis andP. nobilis relied on detritus, algae, and small crustaceans.     

Plasticity in the temporal organization of behaviour in the limpet <Emphasis Type="Italic">Cellana grata</Emphasis>

The behaviour of intertidal consumers is often tightly constrained to tidal movements, although activity patterns can vary within these constraints. Spatio-temporal variability in behaviour of a limpet, Cellana grata, was analysed over different tidal conditions (spring and neap tides) and during different times of the year (one summer and one winter) at sites in Hong Kong. Activity was generally dictated by tidal movements, being concentrated when animals were awash. Plasticity in behaviour was observed, with some limpets anticipating activity during the summer period and delaying activity during winter time. Limpets were active for a time equal, or slightly less, than time awash. As the time awash exceeded ~14–16 h, however, activity duration decreased. Within this general pattern, tidal variation as well as variation among times of the year was noted, with the lowest dependence on time awash being recorded during winter neap tides. Limpets showed a slight preference for being active during nighttime, which was particularly evident when animals were emersed during the summer period. Although the basic activity in C. grata is constrained to a specific temporal window, this limpet is able to modulate its foraging strategies and resting height, according to local, daily changes in environmental conditions.     

Behavior and ecology of Old World Luperini beetles of the genusAulacophora (Coleoptera: Chrysomelidae)

Summary The attraction of Old World leaf beetles in the genusAulacophora to kairomones and parakairomones which are effective lures for New WorldDiabrotica andAcalymma were investigated. Beetles captured on sticky traps baited with single and multicomponent lures were no different from the control traps for two species ofAulacophora. Yellow colored traps and squash blossoms are attractive toAulacophora beetles which detect sub-microgram quantities of cucurbitacins on silica gel. Leaf feeding behavior and flight activity data are correlated with varietal preference of threeAulacophora species. The common response byDiabrotica andAulacophora to cucurbitacins reinforces the two groups' coevolutionary association with the Cucurbitaceae. The apparent lack of a common response toCucurbita blossom volatiles suggests recent evolutionary pathways are substantially different for these two groups of beetles.     

Abundance of estuarine crab larvae is associated with tidal hydrologic variables

Abundances of brachyuran megalopae and juveniles were measured throughout consecutive tidal cycles during six 2 to 3 d sampling periods in summer 1992, and associated with rates of change of tidal hydrologic variables in the Newport River Estuary. Current speeds and rates of pressure change fitted sinusoidal (tidal) models well; however, rates of salinity and temperature change did not. Analysis of plankton samples taken during spring and neap tides showed peak abundances during nighttime rising tides for all taxonomic groups: Callinectes sapidus, Uca spp., Xanthidae, and Pinnixa spp. megalopae, and Pinnotheres spp. juveniles. Megalopal and juvenile abundances from time-intensive sampling were related to rates of changes in the hydrologic variables using stepwise logistic regression. No hydrologic variable accounted well for the presence of Uca spp. megalopae. Megalopal presence was best predicted by current speed for Pinnixa spp. megalopae, and rates of changes in pressure for xanthid megalopae and Pinnotheres spp. juveniles, and salinity for C. sapidus megalopae. These variables might act as cues causing megalopae to ascend into the water column at a particular point in the flooding tide, and subsequently descend to or near the bottom prior to ebb flow. In this way, larvae which develop on the continental shelf or lower estuary undergo transport up the estuary by behaviorally altering their swimming activity and depth concurrent with tidal changes.     

Turbulence measurements in a small subtropical estuary under king tide conditions

In natural waterways and estuaries, the understanding of turbulent mixing is critical to the knowledge of sediment transport, stormwater runoff during flood events, and release of nutrient-rich wastewater into ecosystems. In the present study, some field measurements were conducted in a small subtropical estuary with micro-tidal range and semi-diurnal tides during king tide conditions: i.e., the tidal range was the largest for both 2009 and 2010. The turbulent velocity measurements were performed continuously at high-frequency (50Hz) for 60?h. Two acoustic Doppler velocimeters (ADVs) were sampled simultaneously in the middle estuarine zone, and a third ADV was deployed in the upper estuary for 12?h only. The results provided an unique characterisation of the turbulence in both middle and upper estuarine zones under the king tide conditions. The present observations showed some marked differences between king tide and neap tide conditions. During the king tide conditions, the tidal forcing was the dominant water exchange and circulation mechanism in the estuary. In contrast, the long-term oscillations linked with internal and external resonance played a major role in the turbulent mixing during neap tides. The data set showed further that the upper estuarine zone was drastically less affected by the spring tide range: the flow motion remained slow, but the turbulent velocity data were affected by the propagation of a transient front during the very early flood tide motion at the sampling site.     

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.     

Rhythmicity of larval release in three species of intertidal brachyuran crabs (Crustacea: Brachyura) from Inhaca Island (Mozambique)

 A study of rhythmicity of larval release in three species of intertidal brachyuran crabs, based on laboratory and field experiments, was undertaken at Inhaca Island, southern Mozambique, using Leptodius exaratus and Macrophthalmus grandidieri from December 1994 to January 1995, and Arcotheres palaensis from April to July 1995. L. exaratus and M. grandidieri showed a semi-lunar cycle in larval release. The release of larvae for L. exaratus, a species having conspicuous larvae, occurred in the first half of the night, after the post-crepuscular high tide, which suggests maximisation of protection of larvae from visual predation. The larval release activity matched the late spring and early neap tides. Results from the field were similar to those from the laboratory. M. grandidieri, having inconspicuous larvae, did not show a pattern related to the light–dark cycle and hatched during spring tides (around full and new moons) to maximise larval dispersion. A. palaensis, living inside the host mussel which inhabits the lowest section of the intertidal zone, did not show a relation with moon phase, tidal or light–dark cycles. Received: 16 February 1999 / Accepted: 8 December 1999     

Annual reproductive cycle of the chiton Acanthopleura japonica

Annual reproductive cycle of the chiton Acanthopleura japonica (Lischke) was studied by observing both gonad index and histological features from October 1980 to October 1981. No significant changes in each factor were observed from October to April. Increased values of gonad index and spermatogenesis or vitellogenesis in gondas were observed from May to September. In the same time, gonad index decreased and sperms or oocytes in gonalds disappeared in the days around spring tide, and they were recovered around neap tide. Suggested spawning around spring tide from May to September is also supported by the occurrence of released eggs in the days around the new and full moons.     

Responses of three mouse species to deterrent chemicals in the monarch butterfly. II. Taste tests using intact monarchs

Summary Peromyscus melanotis is the only one of three mouse species that eats monarch butterflies at their overwintering sites in Mexico. I tested two hypotheses: 1)P. aztecus avoids monarchs because of a bitter taste aversion to cardiac glycosides (CGs) and an inability to reject CG-rich body parts; 2)Reithrodontomys sumichrasti avoids monarchs principally because of a bitter taste aversion to the CGs. None of the species are sensitive to the toxic effects of ingested CGs. Feeding responses of laboratory-reared mice of each species to monarchs with low, medium and high CG concentrations were compared. BothP. aztecus andR. sumichrasti ate significantly fewer of all three types of monarchs thanP. melanotis. ForP. aztecus andR. sumichrasti, the number of monarchs eaten decreased with increasing CG concentration, whereas forP. melanotis, the number remained constant.Peromyscus melanotis andR. sumichrasti developed a feeding technique for rejecting the CG-laden cuticular material, which reduced the bitterness of ingested monarch material. However,R. sumichrasti displayed the technique significantly less often thanP. melanotis; andP. aztecus never developed it. I conclude that high taste sensitivity to CGs and less versatile food handling preventP. aztecus andR. sumichrasti from overcoming the monarch's chemical defenses.     

Feeding and burrowing ecology of two East African mangrove crabs

The behavior and ecology of two mangrove crabs,Sesarma meinerti De Man, 1887 andCardisoma carnifex (Herbst, 1794) were investigated at the beginning of the rainy season (October–November 1988) at Mida Creek, Kenya. Both species occupy upper intertidal levels, above mean high-water neap, and completely overlap in their zonation. Each burrow lasts ca. 3 wk, with no significant difference between the species. Both are more active around dusk and dawn and also follow a similar trend in their foraging activity, but differ in that burrowing is mainly diurnal inC. carnifex and nocturnal inS. meinerti. A hierarchy of food preference, established by offeringC. carnifex andS. meinerti leaves of five mangrove species, proved similar for both, withBruguiera gymnorhiza ranking first andAvicennia marina last, but significant only forC. carnifex. A rough estimate of the amount of litter consumed by these two species and of the soil mixed up by their burrowing activity indicates that they play a role of primary importance in the ecology of East African mangroves.     

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.