Seasonal and Lunar Variation in the Emergence Time of a Population of Uca lactea annulipes (Milne-Edwards, 1837) at a Shore in Kuwait

This study monitored the endogenous emergence time of the fiddler crab Uca lactea annulipes (Milne-Edwards, 1837) in the field, for the first time, at an intertidal shore in Kuwait, from 1997 to 2001. The results revealed a significant cyclic change in the median emergence time as the season progressed from winter, through spring and summer, to autumn (.44, 1.29, 3.12, and 1.1?h prior to the dead-low tide, respectively). The data also revealed a significant shift in the median emergence time according to moon phase (2.27?h at new moon versus 2.56?h at full moon prior to the dead-low tide). (Author correspondence: l.almusawi@ropme.org)     

Dancing to the rhythms of geological time: the biorhythm capabilities of Polychaeta in a geological context

Summary

Errant polychaete worms in the Orders Eunicida (family Eunicidae) and Phyllodocida (families Nereidae and Polynoidae) exhibit a highly developed biorhythmic capability. The Pacific palolo worms are well known for a precisely timed annual breeding event in which mass spawning occurs at a particular time of day, on one day per year, that day having a fixed relationship to the lunar period. Nereidae and Polynoidae exhibit photoperiodic responses that determine the breeding season by regulation of oocyte growth. Nereis virens shows short-term cycles of foraging activity; automated recording of these patterns has revealed four distinct behaviour rhythm phenotypes: circadian, tidal, lunidian and arrhythmic, the last phenotype being expressed during the photoperiod induced growth diapause. The Eunicids and Phyllodocids are represented in the fossil record by scolecodonts, their fossilised jaws. There was a major radiation of these polychaetes during the Ordovician and the earliest suggested polychaete fossils are from the Cambrian. The simultaneous expression of tidal and circadian rhythmicity is characteristic of intertidal animals and it is likely that this complex behavioural repertoire was found in the ancestors of modern terrestrial forms, such as tetrapods and arthropods, prior to their emergence onto land during the Carboniferous and Silurian periods at least 400 Ma. The period of the earth's rotation, and hence day length and tidal period, has long been known to be increasing, and additionally the moon to be retreating from the earth, due to the phenomenon of tidal friction caused by the gravitational interactions between the moon and the earth. These changes are significant over a geological time scale. Consequently, the length of day was substantially less (and the number of days in a year more) than at present in the Cambrian and Ordovician periods. Recent theoretical analysis of the period of the earth's rotation suggests that the day length prior to a critical period (t _crit ) around 1800 Ma may have been stable, with a length of only 4 h. At that time a period of more rapid change in the dynamics of the rotation was initiated. The implications of this theory for the evolution of the biological clock are discussed.     

The rate of predation by fishes on hatchlings of the green turtle (Chelonia mydas)

This study addresses the need for empirical data on the survival of sea turtle hatchlings after entry into the sea by (1) developing a method for measuring marine predation; (2) estimating predation rates while crossing the reef; and (3) investigating the effect of environmental variables on predation rates. Predation rates were quantified by following individual hatchlings, tethered by a 10m monofilament nylon line, as they swam from the water's edge towards the reef crest. Predation rates under particular combinations of environmental variables (tide, time of day, and moon phase) were measured in separate trials. Predation rates varied among trials from 0 to 85% with a mean of 31% (SE=2.5%). The simplest logistic regression model that explained variation in predation contained tide and moon phase as predictor variables. The results suggest that noctural emergence from the nest is a behavioral adaptation to minimize exposure to the heat of the day rather than a predator-escape mechanism. For the green turtle populations breeding in eastern Australia, most first year mortality is caused by predation while crossing the reef within the first hour of entering the sea.     

Temporal scales of variation in settlement and recruitment of the mussel Perna perna (Linnaeus, 1758)

Population dynamics of many intertidal organisms are strongly affected by the abundance and distribution of larvae arriving on the shore. In particular, not only absolute numbers of settlers but also the degree of synchronisation of settlement can have a strong influence on whether density-dependent or density-independent processes shape adult shape populations. Temporal variation in rates of settlement and recruitment of the mussel Perna perna on the south coast of South Africa was investigated using a nested spatial design at different temporal scales. Variability in settlement at spring tides was examined at two temporal scales: lunar (to investigate the effect of state of the moon on settlement) and tidal (to investigate the influence of state of the tide on mussel settlement). Recruitment over neap tides was examined at one temporal scale, fortnight (to investigate the effect of date on mussel recruitment).Strong temporal variation was evident for both settlement and recruitment, but not at all time scales. Distinct peaks of settler/recruit abundance were observed during the lunar and neap tide studies. Recruitment intensity differed over the course of the year, and pulsing of recruitment was generally synchronised among locations. However, the strength of pulsing differed dramatically among locations, giving a significant interaction between fortnight and location. The finest temporal scale, investigated in the tidal study, did not reveal a significant effect of the state of the tide on settlement. The state of the moon (new or full) was not significant as a main factor (p = 0.052), although generally more settlers arrived on the shore during new moon. Phase of the moon appeared to have an effect on settler abundances, but only when and where densities were high.     

Factors affecting recruitment of glass eels into the Grey River, New Zealand

The arrival pattern of glass eels of the shortfin eel Anguilla australis and longfin eel Anguilla dieffenbachii was studied over a two successive migration seasons in the Grey River, South Island, New Zealand. Fishing was carried out on selected nights during September to November, for 3 h per night during 2000 and 2001. The number of shortfin glass eels exceeded longfins in both years; earliest glass eels (September) were larger than later glass eels (November), and longfins larger than shortfins, but there were significant differences in size between years for both species. Environmental variables affecting recruitment differed between years, but common variables for both years were sampling date, time after high tide (incoming tide preferred), time after sunset and moon phase; the influence of moon phase appeared to be expressed both through tides (spring tides preferred) and moonlight. During the year of higher discharge variability, both increasing discharge and increased turbidity were also important.     

The effect of estradiol benzoate on synchrony of estrus and fertility in cattle after removal of a progesterone-releasing intravaginal device.

The aim was to determine the effect of estradiol benzoate (EDB) given after removal of a progesterone-releasing intravaginal device (PRID) at either emergence or dominance of a follicle wave, on the interval to estrus, variation in its onset and pregnancy rate in heifers. Heifers (n=186) were assigned randomly to four treatments in a 2 x 2 factorial design; emergence or dominance of a follicle wave at PRID removal, with or without 0.5 mg EDB 24 h after PRID removal. Ovarian ultrasonography was performed to confirm follicular status; data from heifers of undeterminable follicular status were excluded (n=36). Mean size of the largest follicle of the new wave at PRID removal was smaller (P < 0.01) in heifers given EDB at emergence (6.3 +/- 0.09 mm) compared with those given it at dominance (10.9 +/- 0.30 mm). The onset of estrus was earlier (P < 0.01) in heifers given EDB at dominance (median 42 h, range 13 h) compared with those not given EDB at dominance (median 43 h, range 42 h). The median interval to estrus was decreased (P < 0.01) in heifers given EDB at emergence (median 48 h, range 73 h) compared with those not given EDB at emergence (median 66 h, range 45 h). Variation in onset of estrus was reduced (P < 0.05) in heifers given EDB compared with those not given EDB. The pregnancy rate was not affected when EDB was given at dominance, however, it was decreased (P < 0.05) when given at emergence (23 of 40 vs 26 of 32, respectively). To determine the effect of EDB on follicular dynamics in heifers treated with EDB at emergence, heifers (n=37) were assigned to two treatments: at emergence with or without EDB and their ovaries were examined daily using ultrasonography. Follicular dynamics were not different (P > 0.05) in EDB-heifers compared with untreated controls. Mean serum estradiol was greater (P < 0.01) in EDB-treated heifers compared with controls. In conclusion, 0.5 mg EDB given 24 h after PRID removal to heifers decreased the interval to estrous onset at emergence or dominance, decreased variation in onset of estrus and decreased pregnancy rates when given at emergence of a follicle wave.     

Spawning timing of the cleaner wrasse, Labroides dimidiatus, on a warm temperate rocky shore

We observed spawning behaviors of the haremic cleaner wrasse Labroides dimidiatus at a rocky reef in southern Japan. Females released pelagic eggs daily from June to September after a spawning ascent by pairs. When the high tide occurred between noon and evening (quarter moon to new or full moon), they spawned around the time of the tide providing fast offshore currents. For the rest of the lunar cycle, spawnings occurred during late afternoon independent of the state of the tide at the time of day. This spawning pattern is quite different from that of conspecifics on coral reefs that always spawn around the high tide at all lunar stages to avoid potential egg predators. The wrasse ascended high in the water column, probably because of its predator-immune characters as a cleaner. The steep slope of the study site seemed to contribute to releasing gametes far above aggregating planktivores. Thus, gametes were not subject to heavy predation. Current direction might not always affect survival of spawned pelagic eggs on the temperate reefs so crucially as it does on coral reefs. We conclude that spawning during unfavorable tidal conditions during the daytime may be better than spawning during the best tidal condition in the crepuscular period when predation pressure on adults will be high. Intraspecific variation in the timing and location of spawning of the wrasse may result from different conditions for larval survival. Received: February 6, 2000 / Revised: June 16, 2000 / Accepted: July 17, 2000     

Semilunar rhythm in the zoea-release activity of the land crabs Sesarma

Summary The number of the land crabs Sesarma which released zoeae in a river was counted and recorded for 2 years. The number of crabs releasing zoeae reached a maximum at around every syzygy and decreased to a minimum at around every half moon, showing a semilunar rhythm (Fig. 2). This semilunar rhythm, when examined temporally, showed a peculiar pattern accurately synchronized with the lunar cycle. On the day of and during 4–5 days prior to the syzygy, the peak of zoea-release activity came just after sunset. A few days later, the peak gradually shifted to later in the evening and became flattened until temporal concentration was no longer observed. About 2 days after the half moon, the peak appeared again just after sunset (Figs. 3, 4 and 5). It is not likely that the tide itself affects the semilunar rhythm of Sesarma, but it is supposed that the lunadian factor is involved in it. The adaptive significance of this semilunar rhythm may be interpreted to mean that zoeae released in the river just after the time of spring high tides will successfully arrive at the sea and that the lunadian modification of the peak in the temporal structure will also ensure that the zoeae will be released at the time of high tides and will have a better chance of arriving at the sea than they would otherwise do.Contributions from Shimoda Marine Research Center, The University of Tsukuba, No 331     

OBSERVATIONS ON THE DISTRIBUTION,EMERGENCE AND BEHAVIOUR OF CENTRAL AFRICAN CHAOBORIDAE

SUMMARY

The distribution of four species of Chaoboras over selected parts of Zimbabwe is given. The effect of temperature on the duration of the larval life cycle is discussed in relation to the generation time and the lunar periodicity of the adult emergence period. There were apparently two generations of larvae present in the habitat at any one time, although these generations were not distinct due to the variations in the time taken by the larvae to complete development at any temperature. The emergence was synchronized to the lunar cycle, but the actual moon phase at which emergence occurred was variable, as the two populations under observation both changed from new moon emergences to full moon emergences during the study. Some observations on the behaviour of adult Chaoboras edulis are given.     

Time of day affects heart rate recovery and variability after maximal exercise in pre-hypertensive men

ABSTRACT

Previous literature suggests that human behaviour and physiology are somehow altered by the moon-cycle, with particular emphasis on poorer sleep quality and increased aggressive behaviour during full moon. The latter variables can negatively impact athletes’ recovery and increase the likelihood of injury resulting from collision with another athlete. Therefore, the current study aimed to investigate the association between the lunar cycle and injury risk in professional football players (soccer). We monitored injuries and player exposure in the premier professional league in Qatar during four consecutive seasons (2013–2014 through 2016–2017). Acute (sudden-onset traumatic) injuries (n = 1184; 587 from contact with another player and 597 without player contact) recorded during matches and training were classified according to the lunar cycle characteristics on the date of injury: (i) moon illumination, (ii) lunar distance from earth and (iii) tidal coefficient, acquired from the lunar calendar and tide tables. We used a Poisson regression model to examine the relationship between injury risk and lunar cycle characteristics. We did not detect any association between injury risk and moon illumination, earth-to-moon distance or tidal coefficient, not for all acute injuries, nor for contact and non-contact injuries when examined separately. The findings suggest that the full moon or new moon or the gravitational pull have no effect on football injuries. Thus, organisers need not consult moon or tide tables when planning future event schedules.     

Emergence behavior of a tide pool fish <Emphasis Type="Italic">Praealticus tanegasimae</Emphasis> (Teleostei; Blenniidae) on subtropical reefs

Praealticus tanegasimae is a rockpool blenny that occurs around the higher edge of the intertidal zone in subtropical waters of the western Pacific. Terrestrial emergence from tide pools was confirmed in the blenny during daytime low tide periods on Kuchierabu-jima Island, southern Japan. They stayed in the pools, putting their heads above the water surface, and then jumped out with a tail flip. The duration for each terrestrial emergence ranged from 4 to 665 s (median 31 s), during which they never showed reproductive-related behaviors or active feeding behavior on land. The emergence behavior was often followed by hopping toward neighboring pools, and suggested that the goal of the behavior was to change pools for escape from unfavorable water conditions.     

Lunar phase influence on the glycemia of worker honeybees

The variations of the concentrations of trehalose and glucose, the major components of the haemolymph of emerging worker honeybees, have been examined with respect to the lunar phase. Two series of experiments were performed independently, from April to June and from July to September respectively. The results revealed significant peakings for both carbohydrates at full moon and at new moon, and also a minor peak at last quarter. The data obtained for trehalose in the July-September period are consistent with the existence of a circaseptan subcomponent of period tau congruent to 132h.     

Rhythms in the biting behaviour of a mosquito Armigeres subalbatus

Summary The biting cycle of Armigeres subalbatus is distinctly crepuscular, exhibiting two peaks of activity, a smaller one at dawn and a larger one at dusk. The biting cycle is entrained to natural light-dark cycles and the time interval from dawn to dawn or dusk to dusk peaks is exactly 24 h and from dawn to dusk or dusk to dawn is about 12 h measured at 50% level. This rhythm manifests itself day after day without any marked qualitative change.The rate of change of light intensity may determine the onset of crepuscular biting. The sudden increase (up to ca. 17 lx) or decrease (down to ca. 4 lx) in the intensity of ambient light at the time of sunrise or sunset coincides with the peak of the biting activity.The density of the population of the host-seeking females fluctuates in relation to the phases of the moon, increasing with the full moon phase and decreasing with the new moon phase.Even though the density of the population is greater outdoors than indoors both at ground levels and in the first floor, the peak of activity occurs at the same time in all the places. A vertical stratification of biting activity was also noticed.     

Activity and feeding of Dotilla fenestrata (Brachyura: Ocypodidae) in a warm,temperate South African estuary

The activity of and consumption of organic material by the sand-bubbler crab Dotilla fenestrata was studied over neap and spring tides on a sheltered sand bank close to the mouth of the warm, temperate Kowie Estuary, South Africa. Crabs emerged from their burrows only after the tide receded, and it was light. Time to emergence therefore varied from about 30 minutes to three hours after exposure, depending on the time of low water in the early morning vs at midday. General activity of the crab population was longer on a spring tide (about five hours) than on a neap tide (about three hours). Maximum densities of active crabs were 140 m^–2 and 41 m^–2 on spring and neap tides, respectively. After emergence, crabs spent 60% to 80% of their time feeding. In the Kowie Estuary, D. fenestrata produced between 7 and 12 pseudofaecal pellets, average weight 0.0358 g per pellet, per minute. These pellets had a significantly lower organic and chlorophyll a content than the substratum, and it was estimated that crabs removed about 25% of the organic content from the sediment.     

Genetic adaptation in emergence time ofClunio populations to different tidal conditions

Summary 1. The emergence times of intertidalClunio-species (Diptera, Chironomidae) are correlated with special tidal conditions in such a way that the immediately following reproduction of the short-lived imagos can take place on the exposed habitat.2. If the habitat of aClunio-species is situated in the middle tidal region and exposed twice a day by the tidal cycle (T = 12.4 h), a tidal rhythm of emergence with an average period of 12.4 hours may result (example:Clunio takahashii).3. If the habitat is located in the lower tidal zone, exposed only at about the time of the spring tides, a semilunar rhythm of emergence is expected (examples:Clunio marinus andClunio mediterraneus). These semilunar rhythms are correlated with certain conditions of low tide which occur at the coastal locations every 15 days at about the same time of day. The semilunar rhythm is therefore exactly characterized by two dates: the lunar emergence time (a few successive days around full and new moon) and the diurnal emergence time.4. According to experimental investigations on the control of the emergence rhythm, the midges are able to determine both dates in advance.5. Coastal populations differ in their lunar and diurnal emergence times. These differences correspond to the time of low tide which exists at each location during the emergence days of the semilunar rhythm.6. Crossbreeding between stocks of different populations showed that the differences in diurnal emergence time are gene-controlled.

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Genetische Adaptation der Schlüpfzeiten vonClunio-Populationen an verschiedene Gezeitenbedingungen

Kurzfassung Die Schlüpfzeiten der in der Gezeitenzone lebendenClunio-Arten (Diptera, Chironomidae) sind mit bestimmten Wasserstandsbedingungen synchronisiert, und zwar derart, daß die unmittelbar anschließende Fortpflanzung der kurzlebigen Imagines auf dem trockengefallenen Habitat stattfinden kann. Wenn das Habitat einerClunio-Art in der mittleren Gezeitenzone liegt und parallel zu dem halbtägigen Gezeitenzyklus (T = 12,4 h) zweimal täglich auftaucht, dann kann sich eine 12,4stündige Schlüpfperiodik einstellen (Beispiel:Clunio takahashii). Wenn das Habitat in der unteren Gezeitenzone liegt und nur um die Zeit der Springtiden auftaucht, dann ist eine 15tägige (semilunare) Schlüpfperiodik zu erwarten (Beispiele:Clunio marinus undC. mediterraneus). Diese 15tägige Schlüpfperiodik ist synchronisiert mit bestimmten Niedrigwasserbedingungen, die an einem Küstenort alle 15 Tage jeweils um die gleiche Tageszeit auftreten. Sie wird daher durch zwei Daten eindeutig gekennzeichnet: (1) die lunaren Schlüpftage (wenige aufeinanderfolgende Tage um Voll- und Neumond) und (2) die tägliche Schlüpfzeit. Wie experimentelle Untersuchungen über die Steuerung der Schlüpfperiodik zeigten, können die Tiere beide Daten richtig vorausbestimmen. Die einzelnen Küstenpopulationen unterscheiden sich allerdings in Anpassung an die örtlichen Gezeiten- und Standortbedingungen recht auffällig in ihren lunaren und täglichen Schlüpfzeiten. Kreuzungsversuche zwischen Laboratoriumsstämmen verschiedener Populationen belegen, daß die Unterschiede in der täglichen Schlüpfzeit genkontrolliert sind.

     

Endogenous rhythms and entrainment cues of larval activity in the horseshoe crab Limulus polyphemus

The American horseshoe crab, Limulus polyphemus (Linnaeus), typically inhabits estuaries and coastal areas with pronounced semi-diurnal and diurnal tides that are used to synchronize the timing of spawning, larval hatching, and emergence. Horseshoe crabs spawn in the intertidal zone of sandy beaches and larval emergence occurs when the larvae exit the sediments and enter the plankton. However, L. polyphemus populations also occur in areas that lack significant tidal changes and associated synchronization cues. Endogenous activity rhythms that match predictable environmental cycles may enable larval horseshoe crabs to time swimming activity to prevent stranding on the beach. To determine if L. polyphemus larvae possess a circatidal rhythm in vertical swimming, larvae collected from beach nests and the plankton were placed under constant conditions and their activity monitored for 72 h. Time-series analyses of the activity records revealed a circatidal rhythm with a free-running period of ≈ 12.5 h. Maximum swimming activity consistently occurred during the time of expected falling tides, which may serve to reduce the chance of larvae being stranded on the beach and aid in seaward transport by ebb currents (i.e., ebb-tide transport). To determine if agitation serves as the entrainment cue, larvae were shaken on a 12.4 h cycle to simulate conditions during high tide in areas with semi-diurnal tides. When placed under constant conditions, larval swimming increased near the expected times of agitation. Thus, endogenous rhythms of swimming activity of L. polyphemus larvae in both tidal and nontidal systems may help synchronize swimming activity with periods of high water and inundation.     

Interspecific variation in spawning time and male mating tactics of the parrotfishes on a fringing coral reef at Iriomote Island,Okinawa

Spawning time and male mating tactics of parrotfishes (family Scaridae) were investigated on a fringing coral reef at Iriomote Island, Okinawa. Spawning was observed in 14 species, and more frequently in more abundant species such as Chlorurus sordidus, Scarus rivulatus and Chlorurus bowersi. At the reef-edge spawning site, C. bowersi spawned at high tide, C. sordidus spawned both at high tide and in the early morning, whereas Calotomus carolinus and most of the Scarus species such as S. rivulatus spawned only in the early morning, mostly 0630–0830 h. Spawning only in the early morning irrespective of tide phase and moon age has seldom been reported from the scarid species of other localities. It is suggested that spawning in the early morning would be adaptive in species such as S. rivulatus, which migrated considerable distances (ca. 500 m) to the inshore feeding sites, in order to minimize feeding losses due to migration. For male mating tactics, pair spawning by territorial TP (terminal phase) males occurred in all 14 species, and streaking and group spawning by nonterritorial small IP (initial phase) males were seen more frequently in more abundant species. Moreover, group spawning by nonterritorial TP males, which were larger than the IP males but smaller than the territorial TP males, frequently occurred in S. rivulatus. Such mating tactics of TP males have not been reported from Scaridae.     

Behavioural basis of intertidal zonation in Eurydice pulchra Leach

Eurydice pulchra Leach freshly collected at neap and spring tides exhibits an endogenous circatidal rhythm of swimming when kept without sand in constant conditions in laboratory aquaria. When provided with sand in the laboratory, tidally-phased swimming may occur spontaneously at some times of the lunar month, predominantly after the times of highest spring tides but not at neaps. In these cases, therefore, there appears to be clearly demonstrated for the first time in a marine animal a circasemilunar pattern of emergence from the substratum which is coupled to a circatidal rhythm of swimming; the isopods in the laboratory emerge from the sediment only when the swimming rhythm and emergence rhythm are at their maxima. In the field at neap tides some isopods emerge to swim at high tide, dependent upon their responses to water agitation, their feeding state and responses to light. At and just after spring tides in the field large numbers of the isopods swim at high tide, triggered by the endogenous circasemilunar rhythm of emergence which in the field is probably cued by the effects of increased water agitation. The main function of the endogenous circatidal rhythm of swimming appears to be to permit the isopods to return to their preferred zone in the sand before the fall of the tide. The coupled circasemilunar rhythm of emergence which induces greater swimming at spring tides reduces the risk of animals being stranded high on the shore during neap tides.     

Diel biting activity of Culex (Melanoconion) caudelli in Trinidad, West Indies

The diel biting periodicity of the arbovirus vector Culex (Melanoconion) caudelli Dyar and Knab (Diptera; Culicidae) in the Aripo-Wallerfield forest, Trinidad, was studied by collecting mosquitoes attracted to mouse-baited traps at 2-hourly intervals during eight 24 h periods. Biting females of Cx caudelli were collected during all night-time hours, 18.00-06.00 hours, with an overall unimodal pattern, i.e. one well-defined peak between 22.00 and 04.00 hours. This contrasts with previous reports that Cx caudelli is diurnally active. During the four moon phases, the period of peak biting activity varied from 22.00 to 24.00 hours at fullmoon, from 22.00 to 04.00 hours at first quarter, and from 24.00 to 02.00 hours at new moon and last quarter phases. The number of mosquitoes collected varied significantly with moon phases, the highest (152/24 h) being collected in the first lunar quarter and the lowest (71/24 h) in the last lunar quarter. Cloud cover and rainfall had no significant effect on the diel biting periodicity of Cx caudelli.     

Activity patterns, feeding and burrowing behaviour of the crab Ucides cordatus (Ucididae) in a high intertidal mangrove forest in North Brazil

Activity patterns, feeding and burrowing behaviour of the economically important semi-terrestrial mangrove crab Ucides cordatus (Ucididae, L. 1763) was studied in a high intertidal Rhizophora mangle forest stand in Bragança, North Brazil. Video observations in the rainy and dry season were conducted over 24 h cycles at different lunar phases to investigate the behaviour of these litter-feeding crabs outside their burrows. During the rainy season, crabs stayed inside their burrows for 79% and 92% of the time during day and night, respectively. Time spent for feeding, burrowing and other activities outside their burrows was significantly longer during the day with 9.9% (night: 1.7%) and at waning and waxing moon with 9% (full and new moon: 0.9%). At neap tides (no tidal inundation) foraging and feeding activities outside burrows were clearly light-dependent, increasing at dawn and decreasing at dusk. Highest activities during daytime relate to the visual localisation of food. During the dry season, crabs spent less time inside burrows at neap tides than during the rainy season (80% and 91%, respectively). However, time spent for feeding activities was similar during both seasons. During almost all observation periods crabs collected leaf litter, but rarely fed on it outside burrows. At neap tides nearly all available litter was collected, suggesting that the U. cordatus population is litter-limited during these times. At spring tides (regular tidal inundation) the surface activity of U. cordatus was tide-dependent. Crabs closed their burrow entrances 2-3 h before flooding and re-emerged as soon as the tide retreated. During the day, burrow maintenance was the second most frequent behaviour after feeding. Agonistic interactions were regularly observed and were mainly related to burrow defence. The mean foraging radius of the crabs was only 19 cm (max: 1 m) underneath high Rhizophora mangle trees where crab densities were high. The results point to a high competition for burrows and show that U. cordatus is territorial. It is concluded that several exogenous factors, in particular light, leaf litter availability, flooding of burrows and the presence of conspecifics are important in controlling the crabs' activity patterns.