Semilunar spawning cycles ofFundulus similis (Cyprinodontidae)

Synopsis The longnose killifish,Fundulus similis (Cyprinodontidae), spawns with a semilunar periodicity during an extended breeding season (March through August) along the Alabama (U.S.A.) Gulf coast. Spawning characteristically occurs during the 3–6 days of the ascending and spring tides of each semilunar tidal cycle. Such spawning cycles appear to be a relatively common adaptation within the reproductive strategies of intertidal cyprinodontids.     

HABITAT DIFFERENCES IN THE TIMING OF REPRODUCTION OF THE INVASIVE ALGA SARGASSUM MUTICUM (PHAEOPHYTA,SARGASSACEAE) OVER TIDAL AND LUNAR CYCLES1

Sargassum muticum (Yendo) Fensholt is an invasive species that is firmly established on intertidal and subtidal rocky shores of Europe and the Pacific coast of North America. Local success and spread of S. muticum is thought to rely on its reproductive potential that seems dependent on exogenous factors like tidal and lunar cycles. This study is the first to compare the reproductive patterns (periodicity of egg expulsion and embryo settlement) of this invader in two different habitats: the middle and low intertidal. The combination of monthly, daily, and tidal samples at triplicate sites within each habitat showed a semilunar periodicity of egg expulsion and embryo settlement coincident with increasing tidal amplitude just before full and new moons. In both habitats, duration of each egg expulsion event was ～1 week, and embryo settlement occurred during the first daily low tide and with the incoming high tide during spring tides. However, both expulsion and settlement started 1–2 d earlier, expulsion saturation was faster, and settlement was higher in the mid‐ compared to the low intertidal. Our results suggest that the exact timing of gamete expulsion and embryo release of S. muticum responds to local factors, including tidal cues, which result in differences between mid‐ and low‐intertidal habitats.     

Contrasting reproductive traits in two species of mangrove‐dwelling littorinid snails in a seasonal tropical habitat

Previous studies on the reproductive biology of littorinid snails have focused on rocky shore species, investigating how these gastropods can achieve maximal reproductive success, as well as on processes of sexual selection. This study documented differences in the reproductive traits of two mangrove‐dwelling littorinids, Littoraria ardouiniana and L. melanostoma, in Hong Kong. Reproductive activity of both species was most intense during the summer months. Mating pairs of the two species generally occurred in the tree canopies. Few false mating pairs (same sex or heterospecific pairs: <10%) were recorded, and members of both species showed size‐assortative mating. Littoraria ardouiniana had a shorter reproductive season but a higher intensity of mating and higher seasonal fecundity, than did L. melanostoma. Members of both species showed bi‐lunar periodicities of egg or larval release, synchronized with spring tides. Fecundity showed a strong positive relationship with body size in L. ardouiniana, but not in L. melanostoma. Females of L. ardouiniana released entire broods of larvae in a single brief event, whereas females of L. melanostoma released fewer eggs over 1–8 d. Release of larvae in L. ardouiniana involved a series of short bursts and was much faster than the trickle release of eggs in L. melanostoma. The contrasting reproductive traits in these two species represent different strategies to optimize reproductive success in mangrove habitats.     

Female reproductive cycles in two subspecies of the tropical lizard Mabuya striata

Summary Female reproductive cycles were examined in two subspecies of the live-bearing lizard Mabuya striata from Central Africa, an area with distinct rainy and dry seasons. The low altitude M. s. striata was reproductive throughout the year apart from a brief period at the start of the rainy season, and probably produced three clutches a year. Most females of the high altitude M. s. punctatissima were non-reproductive in the rainy season, came into reproductive condition in the early dry season, and gave birth in the late dry season. For some females, there was a second reproductive cycle starting in the late dry season with birth in the rainy season. For the low altitude M. s. striata it appears that availability of moisture may affect reproduction. For the high altitude M. s. punctatissima it appears that the low temperatures and short hours of sunshine of the early dry season constrain reproduction, and that reproductive cycles are timed to avoid birth occurring in the early dry season.     

Flux of decapod larvae and juveniles at a station in the lower Canal de Mira (Ria de Aveiro,Portugal) during one lunar month

Summary

Emigration and immigration of decapod larvae from estuaries depend on timing of larvae occurrence in the water column relative to the tidal, tidal amplitude and day cycles. The phase relation of these natural cycles varies with tidal regime and geographically, resulting in different time-patterns of hatching of first stage larvae and of presence of late stage larvae in the water column. Vertical migration behaviour according to phase of tide also controls transport inside estuaries. These mechanisms were investigated in a field study conducted on the northwest coast of Portugal where neap ebb tides occur during the night around the quarters of the moon. Flux of decapod larvae through one sampling station was measured during one lunar month at the Canal de Mira (Ria de Aveiro) in the spring of 1990. The sampling programme was comprised of a set of 25-h fixed station studies, separated by 25-h intervals during which no sampling took place. Plankton samples were collected with a pump every hour at three depths. Current velocity and direction at the standard depths, as well as height of the water column, were also measured every hour. Hourly instantaneous flux of larvae through a 1-m-wide vertical section of the Canal de Mira was calculated for the most abundant forms. A total number of 13 combinations of species and larval stages were analyzed, belonging to the families Atelecyclidae, Pirimelidae, Portunidae, Pilumnidae, Grapsidae, Palaemonidae, Crangonidae and Thalassinidae. Patterns of net larval flux along the lunar month could be grouped into three types. Type 1 includes first zoeas that were consistently exported to the sea. Type 2 comprises late zoeas, megalops and juveniles that were consistently imported into the estuary. First zoeas that were imported during some of the 25-h studies but were exported during the others were included in Type 3; in species of this type import periods appeared to alternate with export periods according to lunar phase. Flux of Type 1 larvae followed a semi-lunar pattern. Release activity of Type 1 zoeas took place during the night and started during neap tides around the quarters of the moon, but maximum releases occurred 3–4 h after high tide of average amplitude tides, 3–4 days after the quadratures. These observations agree with the hypothesis that hatching is timed to occur on ebb tides of the largest possible amplitude so that larvae are easily dispersed from areas with a high density of predator fishes. However, based on other observations on the Portuguese coast, it cannot be ruled out that hatching might depend on a minimum number of hours of darkness experienced by the females. Larvae included in Type 2 comprise forms that may have been retained inside the estuary for the entire larval phase, as well as one form that was imported from shelf waters. No semi-lunar pattern of import was detected in this last form. Fluctuations of net flux observed in Type 3 larvae, as well in other forms that were not included in any of the types, were more difficult to explain. These larvae were first zoeas of species belonging to different taxonomic, morphological and ecological groups and may show a diversity of adaptations to the way of life of the adults. Imports and exports of larvae depended not only on time-patterns of abundance, but also on time-patterns of larval vertical distribution. As a general rule, larval stages showed patterns of depth distribution that were consistent with vertical migration rhythmic behaviours synchronized with the tidal cycle. Though the effect was not always statistically significant, first-stage larvae were closer to the surface during ebb, especially during the night, enhancing seaward transport. On the contrary, later zoeal stages, megalops and juveniles were usually closer to the surface during flood, suggesting migration to the water column during this phase of the tide and landward transport.     

Combined effects of tidal and diurnal cycles on spawning of the puffer,Takifugu niphobles (Tetraodontidae)

Synopsis Spawning in the pufferTakifugu niphobles was observed at Tomioka in western Kyushu, Japan, and the factors affecting the spawning time were estimated using a multiple regression analysis. Spawning in this puffer occurred in the intertidal beach during evening rising tides around the full and new moons. Both the time of commencement and ending of a day's spawning were related mainly to the tidal cycle, and occurred later when the high tides occurred later. However, the termination of a day's spawning was also affected by the diurnal cycle; spawning ended earlier when sunset was earlier. Besides, a day's spawning tended to be concentrated within a shorter time span when the interval between the commencement of spawning and the sunset was shorter. Thus the spawning time of the puffer is regulated not only by the tidal but also by the diel cycle, and it is suggested that the adaptive significance of the spawning reactions to both cycles should be examined separately.     

Spring-neap cycle as a major driver of temporal variations in feeding of intertidal fishes: Evidence from the sea catfish Sciades herzbergii (Ariidae) of equatorial west Atlantic mangrove creeks

On macrotidal coasts, short- to medium term variations in feeding of intertidal fishes are influenced by several interacting time scales. To identify the driver of major variations in the feeding habits of intertidal fish, we used the pemecou sea catfish Sciades herzbergii (Ariidae), an abundant intertidal benthic second order consumer, as a model species. We analyzed the influence of the spring-neap and the day-night cycle on intertidal abundance, stomach fullness, diet composition and food consumption of S. herzbergii using block nets set at slack high tides in two mangrove creeks in north Brazil. At spring tides, intertidal abundance, stomach fullness, and total daily consumption of S. herzbergii were on average 8.8, 1.9, and 3.8 times higher than at neap tides, respectively. At spring tides, Uca spp. and Grapsidae (mostly Pachygrapsus gracilis) dominated the diet, irrespective of the time of day. Other important food items were Insecta and the semiterrestrial crab Ucides cordatus. At neap tides, Capitellidae contributed to the diet of nightly inundations while no specimens were caught at daytime. Creek location had no effect on any variable. Results from our study area and evidence from other studies suggest that the spring-neap tide pulse is likely the major driver of short- to medium term variations in feeding of intertidal fishes. This has important implications for feeding-related issues on macro- and probably also on mesotidal coasts: (i) juvenile fishes may have fortnightly growth spurts, (ii) intertidal prey populations may suffer regular fluctuations in mortality, (iii) studies of the feeding ecology of intertidal fishes should cover the combined effects of the tidal, diel, lunar and seasonal cycles, and (iv) the modeling of food webs should consider the differences between highly dynamic spring tide and quieter neap tide conditions.     

Seasonal and tidal abundance patterns of decapod crustacean larvae in a shallow inlet (SW Spain)

Planktonic larvae of decapod crustaceans were collected monthlyfrom July 1991 to June 1992 by pumping during nocturnal floodand ebb tides to establish seasonal larval abundance patternsin an inlet of the Bay of Cdiz. Additional 24 h series of sampleswere collected seasonally (July 1991, October 1991, January1992 and May/June 1992) during spring and neap tides to analyselarval abundance in relation to the main environmental cycles(diel, tidal and lunar phases) and vertical position in thewater column. First zoeae were the most abundant stage for mostspecies, representing 97.6% of all individuals collected. ZoeaI abundance was higher in spring and swmner and, on most samplingoccasions, there was a net output from the inlet to the bay.Five species (Liocarcinus arcuatus and Liocarcinus vernalis,Uca tangeri, Diogenes pugilator and Panopeus africanus) represented60% of total individuals caught. The seasonal occurrence offirst zoeae of the most abundant species indicated two differentreproductive patterns: species with a short reproductive periodand species spawning year round. Zoea I of several species (Panopeusafricanus, Uca tangeri, Pachygrap sus marmoratus, Processa spp.)were significantly more abundant during ebb tides and theirlater larval stages were scarcely collected, suggesting thatthese larvae are released in the inlet and exported to the bay.Conversely, a net input of first zoeae was observed for otherspecies (D.pugilator and Pinnotheres pinnotheres), but theirlater larval stages were also scarcely collected. Such importationcould be a larval rhythm artifact due to release of larvae inthe bay that drifted into the inlet by tidal currents. The crabIlia nucleus, whose later larval stages were collected frequently,was the only species that seemed to complete its life cyclewithin the bay. These results suggest that the studied inletwas primarily used by decapods as an adult habitat and spawningground, while larval development occurred in open sea. Sincevertical migration was not observed for exported larvae, thetidal synchronization of female release seemed to be the mostprobable mechanism of larval exportation. There were no significantdifferences between larval release during spring and neap tides.     

Lunar synchronization of the monthly reproductive rhythm in the sea urchin Centrostephanus coronatus Verrill

The monthly reproductive rhythm in the diadematid sea urchin Centrostephanus coronatus Verrill at Santa Catalina Island, California, was studied in the summer of 1973 and the results are compared with data for the summer of 1969. In the summer of 1973 the more extreme spring tides coincided with the new moon, while in the summer of 1969 the more extreme spring tides coincided with the full moon. The reproductive rhythm in both years was closely synchronized with lunar phases and not with the monthly tidal cycles; spawning occurred near the third lunar quarter in both years. These observations suggest that this monthly reproductive rhythm is synchronized by monthly changes in moonlight, and not by monthly tidal changes.     

Physiological aspects of sound communication in crickets (Gryllus campestris L.)

Summary An extremely elaborate performance, involving endogenous timing with both tidal and lunar frequencies, has been recorded in the locomotor activity of an adult specimen of the intertidal isopod,Excirolana chiltoni. During two months of observation, under constant, non-tidal conditions, this animal showed a persistent tidal rhythm in its swimming activity. Bursts of activity were initially well synchronized with times of tide crest on the shore. The average free-running period of the tidal rhythm was about 24 h 55 min, i.e. about 5 minutes longer than the average period of the tides; thus, the loss of synchrony with the concurrent tides was very gradual. The amount of activity per burst showed a conspicuous pattern of variation, a periodic amplitude modulation which paralleled, in detail, the complex lunar cycle of changes in height of high tide. The free-running period of the bimodal, circa-lunar rhythm of amplitude modulation was one or two days longer than the natural 29-day lunar cycle of tide heights.Each feature of this recording has been qualitatively replicated in activity records from other individuals of this species. Freshly-collectedExcirolana generally show spontaneous bursts of activity at times of tide crest, bursts which are repeated as a persistent tidal rhythm, the period of which commonly departs by only a few minutes from that of the natural tidal cycle. Superimposed on the tidal rhythm is an endogenous monthly pattern of amplitude modulation, which alters the amount of activity per burst. This circa-lunar rhythm has a free-running period between about 26 and 33 days, and generally leads to maximum activity on days of highest of high tides. The net result of the tidal and lunar rhythms is an activity pattern which permits the isopods to recapitulate, in great detail, certain significant ecological aspects of the mixed, semi-diurnal tidal regime of California.The experimental data are not compatible with the hypothesis that uncontrolled environmental factors, such as vibrations from waves, were responsible for the rhythmic behavior of the animals. Neither do the data support the hypothesis that beats between the observed tidal rhythm and a hidden daily or circadian rhythm were responsible for the observed circa-lunar rhythm. Furthermore, the pattern of the circa-lunar rhythm cannot be accounted for by a single monthly oscillation in the excitability of the animals, such as might be mediated by changes in the level of an excitatory or inhibitory hormone in the circulatory system.     

Larval release rhythms and tidal exchange in the estuarine mudprawn, Upogebia africana

The mudprawn, Upogebia africana is common in intertidal regions of many South African estuaries. The life cycle is complex, incorporating a marine phase of development during the larval stages. Breeding peaks are in summer and first-stage larvae are released into the plankton at night. Maximum release activity and export to the marine environment follow a semi-lunar cycle synchronized to the time when high water in the estuary is crepuscular. This occurs after peak spring tidal amplitude. Estuarine reinvasion by postlarvae is also nocturnal, and maximum return occurs after neap's when low water at sea occurs around sunset. Rhythmic cycles of larval export and postlarval estuarine reinvasion are therefore asynchronous during the lunar cycle and are best explained by the timing of the change in light intensity relative to high and low water respectively. If maximum activity rhythms of Stage 1 and postlarvae are independent of tidal amplitude, then timing of maximum release and reinvasion during the lunar cycle would alter as the time of sunset shifts between solstices. Much of southern Africa experiences a semi-arid type climate and most estuaries close off from the sea for varying periods owing to sandbar development across tidal inlets. Larvae do not metamorphose if trapped in estuaries and recruitment ceases. Thus, mudprawn populations are directly affected by tidal inlet dynamics. In extreme cases populations become locally extinct if inlets remain closed for extended periods.     

Circadian components of semilunar and lunar timing mechanisms

The timing of semilunar as well as lunar reproductive rhythms has been analyzed in different geographic populations of the intertidal chironomid Clunio. In stocks of three populations differing in period and phase relationship with the lunar month, these long-term rhythms were synchronized in the laboratory by using artificial moonlight cycles of 30 days in otherwise 24-hr light-dark (LD) cycles (0.4 lux during 4 successive nights every 30 days in LD 12:12). In LD cycles of various periods, a strong synchronization was only possible in LD 12:12 and LD 11:11, whereas in LD 10:10 and LD 15:15 the synchronization by the 30-"day" moonlight cycle was weak or even absent. The study demonstrates a limited range of circadian periods for entrainment of the long-term rhythms. It is concluded that an LD cycle with a period near 24 hr is an essential zeitgeber condition for semilunar and lunar timing in this marine insect. Further, it is suggested that the underlying physiological timing mechanism of Clunio consists of a circadian function for the perception of the monthly moonlight zeitgeber cycles that entrain the endogenous, temperature-compensated oscillator of the circasemilunar (or circalunar) period. The long-term oscillator triggers the metamorphosis of the insect, and thereby determines the time of its eclosion and reproduction on the shorelines, in correlation with days of spring tides recurring about every 14-15 days.     

Reproducing on Time When Temperature Varies: Shifts in the Timing of Courtship by Fiddler Crabs

Many species reproduce when conditions are most favorable for the survival of young. Numerous intertidal fish and invertebrates release eggs or larvae during semilunar, large amplitude, nocturnal tides when these early life stages are best able to escape predation by fish that feed near the shore during the day. Remarkably, some species, including the fiddler crabs Uca terpsichores and Uca deichmanni, maintain this timing throughout the year as temperature, and thus the rate of embryonic development, vary. The mechanisms that allow such precision in the timing of the production of young are poorly known. A preliminary study suggested that when temperature decreases, U. terpsichores mate earlier in the tidal amplitude cycle such that larvae are released at the appropriate time. We tested this idea by studying the timing of courtship in U. terpsichores and U. deichmanni as temperature varied annually during two years, at 5 locations that differed in the temperature of the sediment where females incubate their eggs. Uca terpsichores courted earlier at locations where sediment temperature declined seasonally but not where sediment temperature remained elevated throughout the year. In contrast, clear shifts in courtship timing were not observed for U. deichmanni despite variation in sediment temperature. We discuss other mechanisms by which this species may maintain reproductive timing. These two species are likely to be affected differently by changes in the frequency and intensity of cold periods that are expected to accompany climate change.     

A Lunar Clock Changes Shielding Pigment Transparency in Larval Ocelli of Clunio marinus

Living in the tidal zones of the sea requires synchronization with the dominant environmental influences of tidal, solar, and lunar periodicity. Endogenous clocks anticipate those geoclimatic changes and control the respective rhythms of vital functions. But the underlying mechanisms are only partly understood. While the circadian clocks in animals are investigated employing neurobiological, molecular, and genetic approaches, clocks with a lunar periodicity have been studied with reference to development and behavior only. Sites of their pacemakers, zeitgeber receptors, and coupled endocrine components are unknown. Here, a lunar‐rhythmic change of shielding pigment transparency in the larval ocelli of the intertidal midge Clunio marinus is demonstrated for the first time as a possible access to the neurobiology of lunar timing mechanisms. We studied third instar larvae (Vigo strain) throughout the lunar cycle by light‐ and electron-microscopy as well as by x‐ray fluorescence analysis for the identification of the pigment. Moonlight detection is a prerequisite for photic synchronization of the lunar clock. The larval ocelli of Clunio putatively may function as moonlight receptors and are also controlled by the circalunar clock itself, hence being primary candidates for tracing input and output pathways of the lunar pacemaker. Additionally, the demonstration of a reversible optical change of shielding pigment transparency in Clunio is a novel finding, not reported so far in any other animal species, and reveals a mechanism to enhance photosensitivity under the condition of very dim light. It represents a remarkable change of a sense organ from an imaging device to a radiometer. Its restriction to the developmental stage susceptible to lunar timing elucidates a unique sensory strategy evolved at the level of sensory input. It also raises basic questions about the biochemistry of optically active pigments, like melanin, and their intracellular control.     

TIDAL RHYTHMS: THE CLOCK CONTROL OF THE RHYTHMIC PHYSIOLOGY OF MARINE ORGANISMS

1. A great number of vital processes are rhythmic and the rhythms quite often persist in constant conditions. The best-known rhythms are circadian; much less is known about circalunadian rhythms, and this review was prepared in an attempt to rectify this deficiency. All through the article comparisons are drawn between circalunadian and circacian rhythms. 2. Activity rhythms. (a) The activity patterns of 28 intertidal animals are discussed. All describe a periodicity with a basic component of 24.8 hours, and this approximate period persists in the laboratory in constant light and temperature and in the absence of the tides. The duration of persistence ranges from a few cycles to months, and is a function of the species studied, the conditions imposed, and individual tenacity. (b) In those few cases where relatively long-term observations have been made, there is a trend for the period of the rhythm to become circatidal, or better, circalunadian. (c) The ‘desired’ phase relationship between rhythm and tidal cycle is species-specific. Geographical translocation experiments have shown that the phase is set by the local tides. (d) In some cases the amplitude of the persistent rhythm mimics the semidiurnal inequality of the tides. (e) In about a third of the species discussed, a circadian component has been found combined with the tidal component. Many of the other studies were of such short duration that a low-amplitude circadian component would have gone unnoticed. (f) The tidal rhythm is innate. However, the rhythm is (i) sometimes lacking in organisms living in non-tidal habitats, or (ii) fades after a spell of incarceration in constant conditions. Various treatments — some aperiodic — can induce the expression of the missing tidal rhythm. (g) In the green crab, removal of the eyestalks destroys the activity rhythm. 3. Vertical migration rhythms. (a) A rather surprisingly large number of intertidal animals have been found to undergo migration rhythms between the upper layers of the substratum and its surface. The movements are synchronized with the tides in nature, but most species have either been shown to be diurnal in constant conditions, or in cases where adequate testing has not been done, suspected of being so. (b) In only one species has confirming work shown that the fundamental frequency is truly tidal. This finding is especially important as it shows that tidal rhythms need only the single-cell level of organization for expression. Even at this level there appears to be a dictatorial override by a circadian clock. 4. Colour change. Low-amplitude tidal rhythms in colour change — superimposed on a more dominant circadian change — have been reported to be intrinsic in four species and inducible in a fifth. 5. Oxygen consumption. Tidal rhythms in oxygen consumption have been described for seven invertebrates and one alga; six of the species have superimposed solar-day rhythmic components also. 6. Translocation. A total of five geographical translocation experiments, in which the organisms were maintained in constant conditions throughout, have been tried. Unequivocally in one case, and possibly in a second, the test organisms rephased spontaneously to the times commensurate with local tidal conditions. In two other cases, the pretranslocation phase was retained. The fifth experiment has not been reproducible. 7. Determination of phase. (a) The tidal cycle on the home shoreline sets the phase of the inhabitant's rhythms. Even the location of a crab's burrow on the beach incline can play a determining role. (b) Paradoxically, the periodic wetting by inundation is not an important entraining factor for most intertidal organisms. Instead, the effective portions of the tidal cycle include one or more of the following. (i) Mechanical agitation, especially for animals living in an uprush zone where they are periodically subjected to the pounding surf, (ii) Temperature cycles, though they have not yet been systematically investigated, have very pronounced entraining roles in crabs. (iii) Pressure is probably not a generally important entraining agent for most intertidal organisms, but it is so for the green crab. (c) Light-dark cycles in general, whether daily or tidal in length, have no effect on the entrainment or phase setting of many tidal rhythms. There are two exceptions: (i) a 24-hour light-dark cycle is known to keep a tidal locomotor rhythm (one that becomes circalunadian in constant conditions) at a strict tidal frequency. (ii) In rhythms with both daily and tidal components, when the former is shifted by light stimuli, the latter is affected in a nearly identical manner. 8. Temperature. (a) The role of temperature on tidal rhythms is compared with its role on circadian rhythms. (b) The effects of different constant temperatures have so far been studied on only four tidal rhythms. All studies indicate a lack of any permanent change in period, which is not so with most circadian rhythms; the latter having temperature coefficients around 1.1. In two of the studies the rhythms under test temperatures were followed for less than a day, and a third study cannot be repeated. (c) Short exposure to very cold temperature pulses produced a response that may be interpreted as a temporary stoppage of the clock. Exposure to relatively less-cold pulses appear simply to reset the hands of the clock. The same responses have been demonstrated with circadian rhythms. (d) In the case of green crabs, which had become arrhythmic during prolongued captivity in the laboratory, a tidal rhythm could be reinitiated by a single short cold treatment. The cold pulse also set the phase of the rhythm. (e) A few superficial studies employing temperature steps or pulses have produced results which suggest that a phase-change sensitivity rhythm — just like that found associated with circadian rhythms — may underlie tidal rhythms. Certainly a determined search for this rhythm should be made in the near future. 9. Clock control of rhythms. (a) An argument is constructed claiming that tidal rhythms have a basic period of about 24–8 hours rather than the more expected tidal interval of 12.4 hours. In constant conditions, a circalunadian period is usually displayed. (b) After speculating that a frequency-transforming coupler may function between the clock and the overt rhythm, reasons are given that lead to the further speculation that both circadian and circalunadian rhythms could be generated by a single clock, via specific coupling mechanisms. (c) Two current hypotheses concerning the nature of the clockworks are reviewed and discussed. (d) Suggestions are made for future investigations.     

Lunar-related reproductive behaviour in the badger (Meles meles)

Cyclicity in behaviours, including reproduction, in relation to the lunar cycle is widely documented in some phyla, but weak or unknown in Class Mammalia. In this paper we present long-term video surveillance data of wild Eurasian badgers Meles meles, which reveal a strong correlation between reproductive behaviour and the lunar cycle. Squat marking and raised-leg urination, which increase in frequency at times of reproductive activity, showed maxima around day 22 of the lunar cycle (i.e. new moon). These findings were supported by observations of matings, together with published records, which showed significantly higher occurrence in the lunar dark phase (last quarter to first quarter). We propose that the lunar cycle has the potential to act as a regulator of the reproductive cycle in the badger.     

Number of reproductive cycles of Varroa jacobsoni in honey-bee (Apis mellifera) colonies

Very little data exists concerning the number of reproductive cycles performed by individual Varroa mites. To understand the population dynamics of the Varroa mite it is necessary to know the number of fertile female offspring each Varroa female produces during her lifetime. The lifetime reproduction capacity of the mite consists of the mean number of fertile female offspring produced during each reproductive cycle multiplied by the mean number of cell passages. This paper describes an experimental design to estimate the number of reproductive cycles where mites are transferred to new mite-free colonies for reproduction in sealed brood cells. The data presented suggests that the mean number of reproductive cycles performed by the individual female mite is larger than previously accepted. Under optimal conditions, the mean number of reproductive cycles by Varroa females is probably greater than 1.5 but less than 2. Furthermore, the results show that the reproductive success of Varroa females going into cells to reproduce is not influenced by previous brood cycles.     

Patterns of larval release in the Florida stone crab, Menippe mercenaria

Larval release patterns in brachyuran crabs are often synchronized with environmental cycles. While previous studies have focused extensively on supratidal and intertidal taxa, there have been relatively few investigations of subtidal species. This study examined patterns of larval release by the Florida stone crab, Menippe mercenaria, from three different tidal regimes. Ovigerous stone crabs were collected from Sebastian Inlet on the east coast of Florida, Tampa Bay on the west coast of Florida, and the Florida Keys. Patterns of larval release were monitored in the laboratory in relation to local tidal and diel cycles. Results showed a significant diel pattern in initiation of hatching by crabs from each of three study areas. Larval release consistently occurred during the diurnal phase despite the maintenance of females in constant laboratory conditions for up to 96 h prior to hatching. This implies that release may be controlled by a circadian clock. Patterns of release by stone crabs in relation to tidal cycle were more variable. Larval release by females from populations near Tampa Bay and Sebastian Inlet were not synchronized with the tides, whereas females collected from the Florida Keys exhibited a pattern that was strongly related to tidal cycle. These results may be explained by differences in tidal amplitude at the three sampling locations.     

Circannual rhythm of food intake in spotted munia and its phase relationship with fattening and reproductive cycles

Summary The tropical spotted munia,Lonchura punctulata, exhibits a pronounced seasonality in reproduction, body weight and food intake. The body weight cycle parallels the reproductive cycle, whereas the food intake cycle is almost in antiphase to it. There is evidence that constant dietary restriction (50% of the maximum intake) causes dissociation of the body weight and reproductive cycles. Because of these relationships we performed experiments on the food intake cycle and its phase relationships with the reproductive and body weight cycles in birds held under constant light. The results indicate persistence of all three cycles. The phase relationships of these three parameters were almost normal, which may indicate that the endogenous oscillators for the cycles of reproduction, body weight and food intake are linked.     

Field Chronobiology of a Molluscan Bivalve: How the Moon and Sun Cycles Interact to Drive Oyster Activity Rhythms

The present study reports new insights into the complexity of environmental drivers in aquatic animals. The focus of this study was to determine the main forces that drive mollusc bivalve behavior in situ. To answer this question, the authors continuously studied the valve movements of permanently immersed oysters, Crassostrea gigas, during a 1-year-long in situ study. Valve behavior was monitored with a specially build valvometer, which allows continuously recording of up to 16 bivalves at high frequency (10?Hz). The results highlight a strong relationship between the rhythms of valve behavior and the complex association of the sun-earth-moon orbital positions. Permanently immersed C. gigas follows a robust and strong behavior primarily driven by the tidal cycle. The intensity of this tidal driving force is modulated by the neap-spring tides (i.e., synodic moon cycle), which themselves depend of the earth–moon distance (i.e., anomalistic moon cycle). Light is a significant driver of the oysters' biological rhythm, although its power is limited by the tides, which remain the predominant driver. More globally, depending where in the world the bivalves reside, the results suggest their biological rhythms should vary according to the relative importance of the solar cycle and different lunar cycles associated with tide generation. These results highlight the high plasticity of these oysters to adapt to their changing environment. (Author correspondence: d.tran@epoc.u-bordeaux1.fr)