Hatching rhythms of Uca thayeri Rathbun: timing in semidiurnal and mixed tidal regimes

We compared the timing of larval release by Uca thayeri exposed to different tidal regimes. Crabs on Florida's East Coast experience semidiurnal tides, whereas crabs on the Florida's West Coast experience mixed tides.

In both populations, hatching occurred shortly after high tide. On the East Coast, most crabs released their larvae between dusk and midnight, a few days before the maximum amplitude spring tides. On the West Coast, most crabs released their larvae during the afternoon tropic tides of greater amplitude. West Coast crabs may release during the day because ebbing tides at night are too weak for effective transport. Thus, at each location, hatching occurs when phase relationships between the ebbing tides, the light–dark cycle, and tidal amplitude are most favorable. Further study is required to determine whether females on each Coast show fixed responses to each tidal regime, or whether they can alter their hatching rhythms upon exposure to different tides.     

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.     

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.     

Larval life history strategies of sub-tropical southern African estuarine brachyuran crabs and implications for tidal inlet management

Soon after hatching the larvae of many estuarine crabs migrate from estuaries to adjacent coastal waters soon, where larval development is completed before the post larval stages recruit to estuaries to settle. This study investigated the larval flux of several brachyuran crabs resident within a subtropical estuary on the east coast of South Africa. Plankton sampleswere collected over two intensive sampling periods during the spring of1997 (19 days) and the summer of 1998 (26 days). Larvae were releasedmaximally when high tide within the estuary was crepuscular, whilerecruitment occurred during nocturnal flood tides, with peak abundanceson the maximum amplitude nocturnal flood tides. The data are discussedin relation to freshwater inflow problems facing South African estuaries andthe active management of the tidal inlets of affected systems.     

The vertical and transverse distribution of larval herring in the River Blackwater estuary, Essex

The vertical and transverse distributions of spring-spawned larval herring were studied in the River Blackwater estuary, Essex. Sampling was undertaken during May when larval densities within the estuary were at a maximum for the year. At this time, larval lengths were in the range 7–16 mm. No clear diel pattern in the vertical distribution was found, although larval distribution at night was more even.
The tides were found to influence the vertical distribution. Surface waters (< 1.5 m in depth) were avoided during both flood and ebb tides, but used during slack water. Larval densities at ebb were higher near the sea bed than during the flood tide. It was concluded that this behaviour will tend to transport larvae into and retain them within the estuary.     

Patterns of temporal occurrence of brachyuran crab larvae at Saco mangrove creek, Inhaca Island (South Mozambique): implications for flux and recruitment

Temporal patterns of larval occurrence of brachyuran taxa weredescribed from Saco mangrove creek, Mozambique, based on planktonsamples. Brachyuran larvae were collected hourly in four 24h cycles during neap and spring tidal periods at a fixed station,in November 1997 and February 1998. Results indicate a semilunarcycle of larval release activity for most species. Newly-hatchedstages of sesarmids showed a peak occurrence in post-crepuscularebbing tides, Macrophthalmus spp. and Uca spp. showed highestdensities during ebbing tides at night. Dotilla fenestrata andPinnotheridae, in spite of showing a semilunar pattern, didnot display significant differences between day and night. Mostmangrove taxa showed thus a clear larval exportation behaviourduring ebb tides, with little return during the following floods,with varying degrees of dependence on the diel period. Megalopaewere restricted to the night period, and highest values wereobtained during spring tidal periods. However, maximum valuesof megalopae were obtained both during flood and ebb periods,contrary to most previous studies. This is interpreted as abi-directional transport of non-competent megalopae into andfrom the mangrove area.     

Distribution and patterns of emergence of suprabenthic and pelagic crustaceans in a shallow temperate estuary (Ria de Aveiro,Portugal)

Ecology and dynamics of the zooplankton in the Ria de Aveiro, Portugal, were researched to understand the correlation between the patterns of abundance of suprabenthic and zooplanktonic crustaceans and its tidal, day/night and lunar cycles. This study involved the collection of a series of samples during nine 25 h sampling cycles, separated weekly, at a fixed station located near the mouth of the lagoon. In each 25 h cycle, the samples were collected every 2 h with a 500 μm net, at three depths, at the surface, mid-water and over the bottom. The overall effect of the phases of tide and day and depth on the vertical position of the organisms was further investigated through a three-way analysis of variance (ANOVA), in which the phase of the tide (ebb, flood), the phase of the day (day, night) and depth (surface, mid-water, bottom) constituted the three factors in the analysis. The distribution of the species in the water column exhibited distinct tidal, day/night, and lunar patterns, reflecting the ecological particularities of the different species in relation to their seasonal occurrence and behavioural relationships. This distribution showed that for the dominant species, there was a significant interaction between behavioural strategies and cyclic changes of environmental conditions.     

Role of the tidal cycle in the gonadal development and spawning of the tropical wrasse Halichoeres trimaculatus

Many wrasses on coral reefs exhibit daily spawning that peaks around daytime high tides. In this study, we examined tidal-related ovarian development in the threespot wrasse, Halichoeres trimaculatus, a species common on coral reefs in the Indo-Pacific Ocean. When the fish were collected in the morning at different tidal phases, the gonadosomatic index (GSI) and ovarian histology changed; concomitant with increases in GSI towards high tide, a clutch of the most advanced oocytes developed from vitellogenic to maturation stages. Ovulated eggs and post-ovulated follicles (POF) existed in most ovaries around high tide, but only POF remained around ebb tide, suggesting that spawning occurred during or after high tide. We noticed that tidal-related spawning was considerable in the morning and that most ovaries collected on the afternoon high tide exhibited post-spawning features. This suggests that certain labrid species possess plasticity with regard to their spawning time and utilize potent environmental cues to ensure their reproductive success. When pieces of ovary were incubated with precursor steroids, high conversion of testosterone to 17beta-estradiol occurred during high and ebb tides, while that of 17alpha-hydroxyprogesterone to 17alpha,20beta-dihydroxy-4-pregnen-3-one and 17alpha,20beta,21-trihydroxy-4-pregnen-3-one was observed during low and flood tides. Incubation of pieces of ovary with human chorionic gonadotropin resulted in similar fluctuations in the steroid hormones with tidal phase. Production of these steroid hormones correlated with oocyte development in the ovaries and was probably regulated by gonadotropin. These results demonstrate that the daily cycle is fundamental for oocyte development, and that the tidal cycle is superimposed on this process.     

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.     

Reproductive cycles in tropical intertidal gastropods are timed around tidal amplitude cycles

Reproduction in iteroparous marine organisms is often timed with abiotic cycles and may follow lunar, tidal amplitude, or daily cycles. Among intertidal marine invertebrates, decapods are well known to time larval release to coincide with large amplitude nighttime tides, which minimizes the risk of predation. Such bimonthly cycles have been reported for few other intertidal invertebrates. We documented the reproduction of 6 gastropod species from Panama to determine whether they demonstrate reproductive cycles, whether these cycles follow a 2‐week cycle, and whether cycles are timed so that larval release occurs during large amplitude tides. Two of the species (Crepidula cf. marginalis and Nerita scabricosta) showed nonuniform reproduction, but without clear peaks in timing relative to tidal or lunar cycles. The other 4 species show clear peaks in reproduction occurring every 2 weeks. In 3 of these species (Cerithideopsis carlifornica var. valida, Littoraria variegata, and Natica chemnitzi), hatching occurred within 4 days of the maximum amplitude tides. Siphonaria palmata exhibit strong cycles, but reproduction occurred during the neap tides. Strong differences in the intensity of reproduction of Cerithideopsis carlifornica, and in particular, Littoraria variegata, between the larger and smaller spring tides of a lunar month indicate that these species time reproduction with the tidal amplitude cycle rather than the lunar cycle. For those species that reproduce during both the wet and dry seasons, we found that reproductive timing did not differ between seasons despite strong differences in temperature and precipitation. Overall, we found that most (4/6) species have strong reproductive cycles synchronized with the tidal amplitude cycle and that seasonal differences in abiotic factors do not alter these cycles.     

High larval abundances in the Ria Formosa (Southern Portugal)-methodological or local effect?

Zooplankton abundance at a station in the Ria Formosa (SouthernPortugal) was monitored every week or second week over 18 monthsat incoming tide, high tide, ebb tide and low tide. The watersamples were filtered by 32µ mesh and examined under aninverted microscope. The proportion of planktonic larvae washigh all year round. They formed 50–70% of the metazoicplankton organisms during winter and 80 to >90% during summer.An extrapolation implied that this corresponds to 40–80%of the biomass in summer and <20% in winter. On average,roughly three-quarters of the larvae were nauplius larvae; theremaining quarter were meroplanktonic stages, especially ofgastropods, bivalves and polychaetes. Larval abundances variedindependently of tide (high tide-low tide), but showed a significantlynegative correlation with the tidal elevation (spring tide-neaptide) for total larval number and for most larval groups separately.It is outlined that next to a geographical component, theserelatively high larval figures are the result of the mesh sizeused. When treated as a component of the mesozooplankton, larvalnumber and biomass will always be greatly underestimated.     

Evidence for tidally-induced vertical migration of some gelatinous zooplankton in the Wadden Sea area near Sylt

Gelatinous zooplankton is more abundant in the Wadden Sea area of North Sylt than in the adjacent North Sea. The hypothesis is tested that medusae maintain their position in the North Sylt Wadden Sea by ascending to the surface at flood tides and descending to the bottom during ebb tides, thus avoiding the strong ebb currents which could carry them out of this food-rich area. Surface sampling at a main tidal channel revealed great differences between high tide and low tide abundances of five species of medusae in the surface layer.Rathkea octopunctata, Sarsia tubulosa, Eucheilota macultata andPleurobrachia pileus were all more abundant around high tide than during low tide. Bongo net sampling in different depth layers revealed thatPleurobrachia pileus, Bougainvillia ramosa andEucheilota maculata showed a preference for the surface layers around high tide, whereas most of the individuals were found in the deepest layer around low tide. The results suggest tidally-induced vertical migration of medusae in tidal channels. This may assist maintenance of the populations in the Wadden Sea area near Sylt.     

Larval hatching in the horseshoe crab, Limulus polyphemus: facilitation by environmental cues

Female horseshoe crabs, Limulus polyphemus (Linnaeus), lay their eggs in nests on sandy beaches near the high water line. Embryos develop within the sand, hatch into trilobite larvae, and enter the water column when the nest is inundated. Given the diversity of tidal and shoreline inundation patterns that populations of L. polyphemus experience throughout their range (semidiurnal and diurnal tides, microtidal, and nontidal), hatching may also be facilitated by environmental triggers that serve to synchronize hatching and larval emergence with periods of high water. The objective of this study was to determine if larval hatching in L. polyphemus is triggered or facilitated by environmental cues. Stage 21 embryos were subjected to one of seven different treatments that simulated conditions experienced during inundation: (1) hydration, (2) agitation, (3) hydration and agitation, (4) hydration and agitation with sand, (5) osmotic shock, (6) terrestrial hypoxia, and (7) aquatic hypoxia. Hatching rates increased significantly under all simulated tidal conditions compared to controls and were highest (96%) for eggs simultaneously exposed to both hydration and agitation with sand. Measurements of the osmolarity of the perivitelline fluid of developing eggs collected from the field indicated that it is hyperosmotic to the ambient seawater and porewater. Thus, when inundated, eggs also experience a hypoosmotic shock, which would likely facilitate hatching by causing the eggs to swell, rupturing the egg membrane and thereby increasing the likelihood that larvae would hatch and enter the water column during periods of high water.     

Microhabitat and rhythmic behavior of tiger beetle Callytron yuasai okinawense larvae in a mangrove forest in Japan

Mangrove forests are regularly flooded by tides at intervals of approximately 12.4 h (tidal rhythm). Larvae of the tiger beetle Callytron yuasai okinawense in a mangrove forest made shallow burrows in mounds up to 1 m in height constructed by the mud lobster Thalassina anomala. No larval burrows were observed on the forest floor, which was very muddy even during low tide. Some larvae plugged the burrow openings before they were submerged at high tide. The mean interval between consecutive burrow plugging events was 12.37 h, which is similar to the period of tidal cycles. Nine out of 30 larvae plugged the burrow openings even when the burrows did not become submerged. Plugging behavior may be governed by an endogenous biological clock, or may be a response to exogenous information about tidal level (e.g. moisture seeping through the ground).     

Diel and lunar spawning periodicity of the hawkfish Paracirrhites forsteri (Cirrhitidae) on the reefs of Kuchierabu-jima Island, southern Japan

We observed spawning behavior of the hawkfish Paracirrhites forsteri on reefs of southern Japan. Spawning generally occurred after sunset, prior to the full and new moon with semi-lunar spawning peak periodicity. No egg predation was observed in spawning after sunset, and high tide often occurred at dusk in the lunar phases. Thus, diel timing and lunar synchronicity may increase larval survival. We found that the mating activity tended to start earlier in the day during early mating season than during mid-to-late mating season. The advantages of the earlier start of mating activity were also examined in relation to adult biology contexts.     

Migratory behaviour of post-smolt Atlantic salmon during initial stages of seaward migration

The movements of 24 hatchery-reared Atlantic salmon Salmo salar smolts, with miniature acoustic transmitters (pingers) implanted surgically, were determined after release in the coastal waters of Passamaquoddy Bay (mean tide range 6 m), New Brunswick, Canada, to describe the first stages of seaward migration. Automated pinger detection at fixed sites, and pinger location and tracking by boat were used. Post-smolts left the release area rapidly, and the majority moved to open waters of the bay within several tidal cycles. Initially, post-smolts moved with a seaward orientation on ebb tides and held positions on flood tides. Their movements into open waters were diurnal, and the timing corresponded with the state of the tide during which they moved through a narrow channel. Post-smolts moved preferentially through this passageway with the aid of the tidal stream. Successful movements out through the channel occurred during ebb tides and any movements back in were during flood tides. Ground speed of fish moving through the channel was 4·2 body lengths s⁻¹ and faster than the tidal stream velocities in the channel. The relative velocity of fish swimming through the channel was 2 body lengths s⁻¹. Post-smolt movement was indicative of active, directed swimming with a reliance on ebb-tide transport for migration through a coastal area with strong tidal currents. Some post-smolts moved seaward directly with no apparent period of acclimation for the transfer to the marine environment, whereas others delayed their departure. These differences in behaviour were probably related to asynchrony in smolting when fish were released.     

Ontogeny of an endogenous rhythm in Eurydice pulchra

Field catches of Eurydice pulchra (Leach) from an Isle of Man beach indicated that adult animals were active in the water column solely during spring tide periods, whereas juvenile animals swam abundantly at high tide irrespective of lunar phase. Under constant conditions in the laboratory, animals kept in actographs with sand showed a circa-tidal rhythm of spontaneous emergence and swimming activity. In adults, the activity was greatest during “expected” night-time high-waters and was expressed solely after collection during spring tide periods. The rhythm was independent of sex, season or reproductive status. Juveniles showed no such diurnal modulation of the rhythm and the pattern was expressed throughout the semi-lunar cycle. These ontogenetic differences in behaviour are discussed in the light of previous work on the rhythm, and in terms of the differing habits of the mature and immature animals.     

The distribution and abundance of planktonic penaeid larvae in Tudor creek,Mombasa, Kenya

The temporal distribution and abundance of penaeid prawn larvae in surface waters of Tudor creek were studied. Only late mysis and early post mysis stages of development were encountered in the creek waters with a descending gradient from the mouth station (station 1) to the innermost station 5. Larval encounter incidence (% incidence) was highest in plankton tows taken during night spring tides (59%) followed by night neap tides (45.4%), day spring tides (34.9%) and day neap tides (21.4%). This encounter incidence was more skewed towards the seaward stations (1 and 2) during the neap tides than during spring tides. Likewise, larval density (mean catch/standard 5 min. tow) decreased less steeply up the creek during spring tides than it did during neap tides. Higher catch rates were in general obtained in night samples than in day samples at each station and in each neap/spring phase. The diel cycle had a greater effect on both incidence and abundance of larvae than did the tidal cycle.Intensified larval incursion into the creek was observed between March and June.     

Hatching rhythms and dispersion of decapod crustacean larvae in a brackish coastal lagoon in Argentina

Mar Chiquita, a brackish coastal lagoon in central Argentina, is inhabited by dense populations of two intertidal grapsid crab species,Cyrtograpsus angulatus andChasmagnathus granulata. During a preliminary one-year study and a subsequent intensive sampling programme (November–December 1992), the physical properties and the occurrence of decapod crustacean larvae in the surface water of the lagoon were investigated. The lagoon is characterized by highly variable physical conditions, with oligohaline waters frequently predominating over extended periods. The adjacent coastal waters show a complex pattern of semidiurnal tides that often do not influence the lagoon, due to the existence of a sandbar across its entrance. Besides frequently occurring larvae (exclusively freshly hatched zoeae and a few megalopae) of the two dominating crab species, those of three other brachyurans (Plathyxanthus crenulatus, Uca uruguayensis, Pinnixa patagonica) and of one anomuran (the porcellanidPachycheles haigae) were also found occasionally. Caridean shrimp (Palaemonetes argentinus) larvae occurred in a moderate number of samples, with a maximum density of 800·m⁻³. The highest larval abundance was recorded inC. angulatus, with almost 8000°m⁻³. Significantly moreC. angulatus andC. granulata zoeae occurred at night than during daylight conditions, and more larvae (statistically significant only in the former species) during ebb (outflowing) than during flood (inflowing) tides. In consequence, most crab zoeae were observed during nocturnal ebb, the least with diurnal flood tides. Our data suggest that crab larvae do not develop in the lagoon, where the adult populations live, but exhibit an export strategy, probably based upon exogenously coordinated egg hatching rhythms. Zoeal development must take place in coastal marine waters, from where the megalopa eventually returns for settlement and metamorphosis in the lagoon. Significantly higher larval frequency ofC. granulata in low salinities (≤12‰) and at a particular sampling site may be related to local distribution patterns of the reproducing adult population. Unlike crab larvae, those of shrimp (P. argentinus) are retained inside the lagoon, where they develop from hatching through metamorphosis. They significantly prefer low salinity and occur at the lagoon surface more often at night. These patterns cannot be explained by larval release rhythms like those in brachyuran crabs, but may reflect diel vertical migrations to the bottom. It is concluded that osmotic stress as well as predation pressure exerted by visually directed predators (small species or life-cycle stages of estuarine fishes) may be the principal selection factors for the evolution of hatching and migration rhythms in decapod larvae, and that these are characteristics of export or retention mechanisms, respectively.     

The mechanism of retention of pelagic tomcod,Microgadus tomcod,larvae and juveniles in the well-mixed part of the St. Lawrence Estuary

Synopsis We tested the hypothesis that the mechanism of retention of tomcod, Microgadus tomcod, larvae and juveniles in the well-mixed part of the St. Lawrence Estuary is similar to that of sympatric smelt, Osmerus mordax, larvae who actively migrate to the surface during flood tides and to the bottom during ebb tides so as to minimize net downstream displacement. The vertical distribution of tomcod larvae and juveniles was documented during two 98-h sampling series at 2 anchor stations in June and July, 1986. An hourly index of the center of mass of fish in the water column calculated to take into account daytime net avoidance in surface waters suggested that tomcod remained deep in the water column and that their accumulation at the head of the estuary was the result of passive upstream transport by net residual circulation rather than active tidal migrations. For both series, depth of fish was inversely related to density of the water suggesting that the buoyancy of fish influenced their vertical distribution. Tomcod larvae and juveniles were advected by tidal currents. In June, larger larvae were found at low slack water indicating that they were located upstream of smaller larvae. In July, larger juveniles were located downstream of smaller juveniles, the difference in mean length between low and high slack water attaining 20 mm. Ontogenetic buoyancy changes may be responsible for these differences in the vertical distribution of tomcod. Comparisons of the early life-history stages of tomcod and smelt retained in the same area under the same hydrodynamical conditions indicate that more than one mechanism permits retention in a well-mixed estuary and that the observed species-specific patterns of vertical distribution are not simply interpretable as adaptations to retention.