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     

Hatching rhythms in Uca pugilator (Decapoda:Brachyura)

Gravid female Uca pugilator (Bosc) maintained in conditions of 14h L: 10h D light regime and constant temperature without simulated tides showed a significant positive correlation between time of hatch and time of the nighttime high tide. Light: Dark experiments showed no entrainment by LD cues. Larval hatching was delayed until dark when high tide occurred between 1900 and 2130 hrs. Hatch time was a positive linear function of the time of the nighttime high tide when high tide occurred after 2130 hrs. Time of hatch was delayed by the light period on days lacking a nighttime high tide. Experiments conducted under constant light (LL) and temperature, without simulated tides, indicated a persistent free-running rhythm of approximately 25 h 03 min, suggesting an endogenous timer. Rhythmic larval release on the nighttime high tide may be adaptive as a predator-avoidance mechanism for adult gravid females.Contribution No. 366 from the Belle W. Baruch Institute for Marine Biology and Coastal Research, University of South Carolina     

Effects of salinity on endogenous rhythm of the Manila clam, Ruditapes philippinarum (Bivalvia: Veneridae)

The Manila clam Ruditapes philippinarum, an intertidal bivalve, was exposed to different salinity regimes (from 31.0–31.7‰ down to 20‰, 15‰, 10‰, 5‰), and the endogenous rhythm in its oxygen consumption was studied using an automatic intermittent-flow respirometer. When exposed to salinities reduced from 31.5‰ to 20‰ and 15‰ under otherwise constant conditions, the clams recovered a clear endogenous circatidal rhythm in their oxygen-consumption rate after having dampened periods of 12 h and 48 h, respectively. At salinities less than 10‰, however, the oxygen-consumption rate was depressed greatly at the beginning of the experiment for about 36 h and then increased to a level higher than normal, but the rhythm of oxygen consumption was not recovered. The results of this study indicate that the Manila clam, a euryhaline organism, cannot maintain a normal metabolic activity at a salinity lower than 15‰. All clams were dead after exposure at a salinity of 5‰ for 7 days. Received: 28 February 2000 / Accepted: 26 August 2000     

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     

Intraspecific parasitism and split sex ratios in a monogynous and monandrous ant (Leptothorax nylanderi)

Sex ratios were bimodally distributed in a population of the monogynous and monandrous ant Leptothorax nylanderi during each of 3 study years. The population-wide investment ratios suggested worker control of sex allocation. Nest-level variation in the proportional investment in virgin queens was not affected by the presence or absence of a queen and only slightly by collecting year, but was correlated with nest size, total sexual investment and, unexpectedly, with differences in nestmate relatedness: small, low-investment nests and nests with several worker lineages produced male-biased sex ratios. Colonies containing several worker lineages arise from usurpation of mature colonies by unrelated founding queens and the fusion of unrelated colonies under strong nest site limitation. In contrast to facultatively polygynous and polyandrous species of social insects, where workers can maximize their inclusive fitness by adjusting sex ratios according to the degree of relatedness asymmetry, workers in mixed colonies of L. nylanderi do not benefit from manipulating sex allocation, as here relatedness asymmetries appear to be the same as in homogeneous colonies. Received: 7 December 1999 / Received in revised form: 29 February 2000 / Accepted: 13 March 2000     

Natterer’s bat (Myotis nattereri Kuhl, 1818) hawks for prey close to vegetation using echolocation signals of very broad bandwidth

We present a hitherto unknown prey perception strategy in bats: Myotis nattereri (Vespertilionidae, Chiroptera) is able to perceive prey by echolocation within a few centimeters of echo-cluttering vegetation, by using frequency-modulated search signals of very large bandwidth (up to 135 kHz). We describe the species’ search behavior and echolocation repertoire from the field and from experiments in a flight tent. In the field, bats varied signal parameters in relation to their distance from vegetation and usually flew close to vegetation. In the flight tent, M. nattereri detected and localized prey by echolocation alone as close as 5 cm from vegetation. Apparently, the bats were able to tolerate some overlap between prey and clutter echoes. Passive prey cues (vision, olfaction, prey-generated sounds) were not used in prey perception. The bats selected prey by size. The animals performed aerial catches and produced approach sequences typical for aerial hawking bats, but were able to do so within a few centimeters of the substrate. M. nattereri thus has access to silent, suspended prey very close to vegetation (e.g., spiders, and caterpillars on threads). Received: 29 September 1999 / Received in revised form: 12 February 2000 / Accepted: 12 February 2000     

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.     

Mechanisms of selective attention in grasshopper choruses: who listens to whom?

In many rhythmically signaling species of acoustic insects and anurans, males form choruses at typical population densities. Recent findings that females may prefer leading calls indicate that the timing of a male's signals relative to those of neighboring choruses is an important component of mate attraction. Within a chorus, however, males cannot time their calls such that they lead all neighbors. Hence, they are expected to show selective attention toward only a subset of the group. We used field playback experiments to determine the incidence of and mechanism(s) responsible for selective attention in choruses of the territorial grasshoppers Ligurotettix coquilletti and L. planum. Our data revealed significant selective attention in both species and are inconsistent with either fixed-threshold or fixed-number mechanisms. Rather, regulation of selective attention by a sliding-threshold mechanism is supported. We discuss these results in the context of the evolution of chorus structure. Received: 24 September 1997 / Accepted after revision: 28 February 1998     

Effects of sudden changes in salinity on endogenous rhythms of the spotted sea bass Lateolabrax sp.

The endogenous rhythm of oxygen consumption in juvenile spotted sea bass (Lateolabrax sp.) was measured to test the effects of sudden changes in salinity on the metabolic activity. Mean oxygen consumption rates of this euryhaline fish decreased by 13.5 to 16.0% and 25.3 to 36.4% when they were transferred from 31.5 to 15‰ seawater and to fresh water (0‰), respectively. The maximum rate of oxygen consumption was observed between 18:00 and 19:00 hrs local time, 1 to 2 h before sunset, even though they were kept in constant darkness. The peaks of oxygen consumption occurred in 23.2- and 23.3-h intervals, which correspond with a circadian rhythm, as revealed by maximum entropy spectral analysis. A markedly weakened rhythm in oxygen consumption occurred from 8 to 10 d after onset of the experiments. This study indicates that spotted sea bass can withstand sudden drops in salinity from 31.5‰ to fresh water, and yet maintain a regular though somewhat dampened endogenous rhythm of oxygen consumption. Received: 16 June 1997 / Accepted: 3 February 1998     

Ontogenetic differences in the circadian locomotor activity rhythm of the talitrid amphipod crustacean Orchestoidea tuberculata

 Adult talitrid amphipods often display a nocturnal activity pattern, which has been shown in a number of cases to be partly controlled by an endogenous rhythm of circadian period. Juvenile talitrids have been little studied in the past, but evidence from field studies indicates that they may display a different diel pattern of locomotor activity from that of adults. This study presents evidence from the field in south–central Chile that juvenile Orchestoidea tuberculata Nicolet show crepuscular activity peaks, contrasting with the nocturnal peaks shown by adult conspecifics. Under constant conditions in the laboratory, both juveniles and adults exhibit endogenous rhythms of circadian periodicity. However, the phasing of activity differs in juveniles and adults, corresponding to the differences observed in the field. It is suggested that differing phasing of peak locomotor activity may be a mechanism to reduce contact between juvenile and adult talitrids. Experimental evidence indicates a significant negative influence on survival of juveniles in the presence of adult conspecifics by intraspecific predation. Other factors influencing the timing of surface activity are also discussed. Received: 3 August 1999 / Accepted: 26 May 2000     

Correlations of food distribution and patch size with agonistic interactions in female vervets (Chlorocebus aethiops) and patas monkeys (Erythrocebus patas) living in simple habitats

Food distribution is hypothesized to be important in determining the nature of female relationships within social groups of primates. When food limits female reproductive success, spatially clumped foods are expected to produce strong, linear dominance hierarchies within groups, whereas more spatially dispersed foods are expected to produce weaker or non-existent dominance hierarchies. The association between food distribution and competitive relationships presumably occurs because clumped foods are usurpable but dispersed foods are not. We examined the spatial distribution of food patches (trees) and patch size relative to feeding behavior and agonistic interactions in vervets and patas monkeys, two closely related and sympatric species that nonetheless differ in the strength of the female dominance hierarchy. Food patches of both patas monkeys and vervets were small in size and randomly distributed in Acacia drepanolobium habitat. In contrast, in A. xanthophloea woodland, the habitat type that was exclusively used by vervets, food patches were larger and more spatially clumped. These similarities and differences between and within species were correlated with similarities and differences in the strength and linearity of their dominance hierarchies. Patas monkeys and vervets in A. drepanolobium habitat had dominance hierarchies that were weakly defined because there were relatively few agonistic interactions between females. By contrast, in A. xanthophloea habitat, vervets had a stronger, linear dominance hierarchy characterized by a higher rate of agonistic interactions over food. The covariation of agonistic interactions with patch size is discussed in relation to depletion time, another characteristic that may covary with food distribution, and resource renewal rate, an important determinant of agonistic interactions in insectivorous birds, fishes, insects, and mammals. Received: 18 February 2000 / Revised: 5 September 2000 / Accepted: 26 September 2000     

Flood tide transport of blue crab, Callinectes sapidus, postlarvae: behavioral responses to salinity and turbulence

Blue crab (Callinectes sapidus) postlarvae (megalopae) use flood tide transport to move upstream in estuaries during nocturnal flood tides. The megalopae have a endogenous diel rhythm in activity that is inconsistent with this tidally timed behavior. Thus, it is hypothesized that this behavior is regulated by behavioral changes in response to exogenous cues associated with tidal currents. In a laboratory flow tank, blue crab megalopae were exposed to simultaneous changes in salinity and turbulence to simulate tides in an estuary. On simulated flood tides, megalopae ascended upon exposure to a salinity increase, remained swimming during times of high turbulence, and descended at times of low turbulence. Turbulence stimulated swimming for several hours, approximating the duration of tidal currents in estuaries. Swimming was inhibited by decreasing salinity on simulated ebb tides. These results support a model for regulation of flood tide transport by blue crab megalopae as follows: (1) blue crab megalopae are stimulated to swim into the water column by increasing salinity associated with flood tide; (2) megalopae remain swimming during flood tide in response to high levels of turbulence; (3) megalopae descend at the end of flood tide, when current speed and turbulence decline to low levels; and (4) megalopae are inhibited from swimming on ebb tides by the associated salinity decrease. This is the first model for regulation of flood tide transport in a species lacking a tidal rhythm in activity.     

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.     

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.     

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     

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.     

Responses of estuarine crab megalopae to pressure,salinity and light: Implications for flood-tide transport

The megalopal larval stage of many estuarine brachyuran crabs appears to return to adult habitats by undergoing rhythmic vertical migrations which result in saltatory up-estuary transport on flood tides. Larval ascent into the water column during rising tides may be cued by changing hydrologic variables. To test this hypothesis, we investigated the responses of field-caught megalopae of the blue crab Callinectes sapidus and the fiddler crab Uca spp. to constant rates of pressure and salinity change under laboratory conditions. For both genera, pressure changes resulted in increased movement (barokinesis) and upward migration in the test chamber, with C. sapidus megalopae having a lower response threshold (2.8×10^-2 mbar s^-1) than Uca spp. larvae (5×10^-2 mbar s^-1). Similarly, larvae ascended in response to increasing salinity, with C. sapidus larvae being more sensitive. Larvae were negatively phototactic and failed to respond to pressure increases at light levels above 1.0×10¹⁵ and 1.0×10¹³ photons m^-2 s^-1 for C. sapidus and Uca spp. megalopae, respectively. Such responses are thought to explain the low abundances of larvae in the water column during daytime flood tides. Nevertheless, threshold sensitivities to increasing pressure for both genera were above levels experienced during floodtide conditions in the field. Similarly, it is unlikely that increasing salinity is sufficient to induce ascent in Uca spp. postlarvae. However, rates of salinity increase during midflood tide typically reach levels necessary to induce an ascent in C. sapidus megalopae. These results are consistent with the hypothesis that fiddler crab megalopae utilize an endogenous activity rhythm for flood-tide transport, while blue crab megalopae rely upon external cues, especially salinity changes, to time their sojourns in the water column.     

Diel vertical migration of the marine copepod<Emphasis Type="Italic"> Calanopia americana</Emphasis>. II. Proximate role of exogenous light cues and endogenous rhythms

The marine copepod Calanopia americana Dahl undergoes twilight diel vertical migration (DVM) in the Newport River estuary, North Carolina, USA, in synchrony with the light:dark cycle. Copepods ascend to the surface at sunset, descend to the bottom around midnight, and make a second ascent and descent before sunrise. Behavioral assays with C. americana in the laboratory during fall 2002/2003 and summer 2004 investigated aspects of three hypotheses for the proximate role of light in DVM: (1) preferendum hypothesis (absolute irradiance), (2) rate of change hypothesis (relative rates of irradiance change), and (3) endogenous rhythm hypothesis. Results suggest that C. americana responds to exogenous light cues consistent with its DVM pattern; changes in absolute irradiance evoked swimming responses that would result in an ascent at sunset and descent at sunrise, while relative rates of irradiance decrease at sunset (–0.0046 s^–1) evoked an ascent response, and relative rates of irradiance increase at sunrise (0.0042 s^–1) evoked a descent response. Furthermore, C. americana expressed an endogenous rhythm in vertical migration that was positively correlated with field observations of twilight DVM. Collectively, these results indicate that both exogenous light cues and endogenous rhythms play a proximate role in twilight DVM of C. americana, providing redundancy in the causes of its vertical migration.Communicated by J.P. Grassle, New Brunswick     

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     

Endogenous control of timing of metamorphosis in megalopae of the shore crab Carcinus maenas

Batches of hundreds of freshly collected megalopae of the shore crab Carcinus maenas (L.) showed persistent circatidal rhythms of moulting to the juvenile crab stage when maintained in constant laboratory conditions. Peaks of moulting occurred around expected times of high tides, with few megalopae moulting at other times. In larvae collected offshore, the highest tidally-timed peak of metamorphosis occurred during the second to fifth expected times of high tide, and metamorphosis of 50% of each batch took about 22 h or longer. In contrast, in larvae collected at the water's edge, 70% metamorphosed during the first expected episode of high tide, within 6 to 8 h after collection. However, although inshore megalopae moulted before offshore ones, the tidal timing of moulting remained unaltered whether megalopae were collected at neaps or springs, from the water's edge or farther offshore, in the presence or absence of natural substratum, and under various light–dark and salinity regimes. Metamorphosis of C. maenas megalopae around the times of high tides may enhance settlement into the upper intertidal zone. Early juveniles of the crab apparently prefer that zone as they are most abundant there and, unlike adults, do not undertake up-and-down-shore migration with tides. The present finding demonstrates, for the first time, endogenous physiological timing of circatidal periodicity in the metamorphic moult of crab megalopae, suggesting that endogenous factors, as well as exogenous ones should be taken into account in considering the process of settlement by crab megalopae. Received: 21 February 1996 / Accepted: 27 November 1996