Topographically generated fronts, very nearshore oceanography and the distribution and settlement of mussel larvae and barnacle cyprids

Flow patterns adjacent to shore may prevent or aid shorewardmigration of benthic invertebrate larvae. We hypothesized thata front at the mouth of Sunset Bay, Oregon, prevents shorewarddispersal of larvae, significantly altering settlement of mussellarvae and barnacle cyprids. Settlement was measured at threesets of moorings (three moorings per site) distributed acrossthe front at Sunset Bay. From 6 July to 4 September 2000, sampleswere collected roughly every other day. Concurrently, we madevertical zooplankton tows adjacent to each mooring site andcollected physical oceanographic data. During upwelling-favorablewinds, the front was always present at the bay mouth, separatingsignificantly cooler, saltier and denser offshore water fromthat within the bay. During downwelling winds, the front brokedown and we found no significant difference in the surface physicaloceanographic parameters across the bay mouth. During upwelling,the concentration of mussel larvae was higher seaward of thefront than landward, but there was no significant differencein concentration during downwelling, suggesting that the frontmay act as a barrier to the shoreward dispersal of mussels.Mussel settlement was too low and sporadic to allow statisticalanalysis. There was no difference in cyprid concentrations acrossthe bay mouth whether the front was present or not. Cyprid settlementwas, however, nearly an order of magnitude lower at mooringsseaward of the front than at those landward. A significant cross-correlationwas found between settlement at the offshore mooring and tidalrange (r = 0.464, lag = 0 days) and between settlement at themid and inner moorings and downwelling winds (r = 0.532 midbay, r = 0.532 inner bay, lag = 0 days). Seaward of the front,settlement varied with tidal range, while landward of the front,most settlement occurred as brief pulses during downwellingwinds, periods when the front was not present. We found largedifferences in the distribution of cyprids, and mussel larvaeand cyprid settlement relative to the front; larval distributionsand settlement varied with upwelling versus downwelling windsand was due to differences in the very nearshore (i.e. within100–1000 m of shore) coastal oceanography.     

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.     

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.     

象山港熊本牡蛎幼体分布、附着及生长的空间格局

牡蛎幼体迁移与附着影响着牡蛎的空间分布、资源补充、种群维持及生态服务功能。本研究通过野外调查,监测了1个完整潮汐周期(9个潮时采样)内象山港2个试验点(采苗场和产卵场)表层、底层水体中熊本牡蛎幼体的时空动态,并开展野外附着试验,检验了熊本牡蛎在2个试验点和3个潮区(上层T、中层M和底层B)附着及生长的空间格局。结果表明: 在1个完整潮汐周期内,2个试验点表层水体中牡蛎幼体丰度均有显著的时间变化,采苗场表层水体中熊本牡蛎幼体丰度最大值(20.8±5.6 ind·L^-1)出现于高平潮时,显著高于其他8个潮时;而产卵场相反,高平潮时牡蛎幼体丰度最低(0.1±0.1 ind·L^-1);2个试验点底层水体中牡蛎幼体丰度在不同潮时之间均无显著差异。采苗场附着牡蛎总丰度、成活率和壳高在不同潮区间差异显著,牡蛎总丰度大小顺序为B>T=M,成活率和壳高的大小顺序为T>M>B,活体牡蛎丰度在潮区之间无显著差异。产卵场附着牡蛎总丰度、成活率、活体牡蛎丰度和壳高在不同潮区之间均无显著差异。表明高平潮时是熊本牡蛎附着的主要时段,产卵场和采苗场具有相似的牡蛎稚贝附着密度。     

Prezygotic barriers to hybridization in marine broadcast spawners: reproductive timing and mating system variation

Sympatric assemblages of congeners with incomplete reproductive barriers offer the opportunity to study the roles that ecological and non-ecological factors play in reproductive isolation. While interspecific asynchrony in gamete release and gametic incompatibility are known prezygotic barriers to hybridization, the role of mating system variation has been emphasized in plants. Reproductive isolation between the sibling brown algal species Fucus spiralis, Fucus guiryi (selfing hermaphrodite) and Fucus vesiculosus (dioecious) was studied because they form hybrids in parapatry in the rocky intertidal zone, maintain species integrity over a broad geographic range, and have contrasting mating systems. We compared reproductive synchrony (spawning overlap) between the three species at several temporal scales (yearly/seasonal, semilunar/tidal, and hourly during single tides). Interspecific patterns of egg release were coincident at seasonal (single peak in spring to early summer) to semilunar timescales. Synthesis of available data indicated that spawning is controlled by semidiurnal tidal and daily light-dark cues, and not directly by semilunar cycles. Importantly, interspecific shifts in timing detected at the hourly scale during single tides were consistent with a partial ecological prezygotic hybridization barrier. The species displayed patterns of gamete release consistent with a power law distribution, indicating a high degree of reproductive synchrony, while the hypothesis of weaker selective constraints for synchrony in selfing versus outcrossing species was supported by observed spawning in hermaphrodites over a broader range of tidal phase than in outcrossers. Synchronous gamete release is critical to the success of external fertilization, while high-energy intertidal environments may offer only limited windows of reproductive opportunity. Within these windows, however, subtle variations in reproductive timing have evolved with the potential to form ecological barriers to hybridization.     

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.     

Developmental shift in the selective tidal-stream transport behavior of larvae of the fiddler crab Uca minax (LeConte)

Following hatching, larvae of the fiddler crab Uca minax (La Conte) are exported from the adult habitat in estuaries to coastal and shelf waters where they undergo development prior to re-entering estuaries as postlarvae (megalopae). Studies of the spatial distribution of both newly hatched zoeae (Stage I) and megalopae indicate they undergo rhythmic vertical migrations associated with the tides for dispersal and unidirectional transport (selective tidal-stream transport) both within estuaries and between estuaries and the nearshore coastal ocean. We tested the hypothesis that U. minax zoeae possess a circatidal rhythm in vertical migration that facilitates offshore transport in ebb tidal flows, while postlarvae (megalopae) return to estuaries using a similar flood-phased endogenous rhythm. We also determined if the expression of the rhythm was influenced by the salinity conditions zoeae and megalopae experience as they transition between low-salinity regions of estuaries and high-salinity coastal waters. Stage I zoeae were collected by holding ovigerous female crabs in the lab until hatching. Megalopae were collected from the plankton and identified to species using molecular techniques (PCR-RFLP). Under constant laboratory conditions, both zoeae and megalopae exhibited endogenous circatidal rhythms in swimming that matched the principal harmonic constituent of the local tides (12.39 ± 0.07 h; X¯ ± SE). Upward swimming in Stage I zoeae occurred 2.5-4 h after high tide near the time of expected maximum ebb currents in the field. Rhythmic swimming of megalopae occurred slightly earlier in the tide (2.5 ± 0.09 h after high tide; X¯ ± SE) but was not entirely synchronized with flood currents, as expected. Salinity conditions had no apparent effect on the expression or pattern of the rhythms. Results indicate that this circatidal rhythm forms the behavioral basis of selective tidal-stream transport (STST) in early stage U. minax zoeae, but does not undergo a sufficient phase shift to account for vertical distribution patterns exhibited by megalopae in the field.     

Recolonization pattern of the polychaete Lanice conchilega on an intertidal sand flat following the severe winter of 1995/96

The recolonization process of Lanice conchilega was studied on an intertidal sand flat in the northern Wadden Sea after an intertidal population had been wiped out during the severe winter of 1995/96. After very low recruitment in the first two years, the population fully recovered in the third year after its destruction. By then, the distribution pattern closely resembled the pattern before the severe winter with >95% of all Lanice occurring below –0.5 m mean tidal level and dominance of the density categories 6–50 individuals (ind) m^–2 and 51–200 ind m^–2. Lanice larvae have been observed to attach preferentially on the tubes of adults. However, the high recruitment in the third year was also seen in areas devoid of adult Lanice, indicating that settlement facilitation by adults was of little importance for the recolonization process. Instead, we suggest that the observed recruitment pattern was a consequence of a large-scale decline in the metapopulation in the tidal basin and the onshore coastal subtidal which resulted in insufficient larval supply onto the tidal flats and low recruitment in the first two years following the severe winter. Received in revised form: 7 May 2001 Electronic Publication     

Empirical evidence for large‐scale human impact on intertidal aggregations,larval supply and recruitment of Pyura praeputialis around the Bay of Antofagasta,Chile

In northern Chile, Pyura praeputialis is an invasive species inhabiting rocky intertidal and subtidal habitats restricted exclusively to the Bay of Antofagasta where it forms extensive aggregations. The negative impact of Pyura gathering on mid‐intertidal abundances of this species has recently been reported at the south‐eastern end of this bay. In the present study we have increased sampling sites to cover the entire bay toward the north‐western end and the northern section, where a coastal marine reserve for the scallop fishery partially restricts shellfish gathering. Therefore, the sampling sites were chosen to represent different levels of shellfish gathering access along the northern shore of the bay. Long‐term monitoring (1999–2014) of changes in tunicate cover and the abundances of larvae and recruits at seven sites are reported. The opening of a remodelled artificial and recreational beach in 2012, on the central‐eastern shore of the bay, has increased accessibility to rocky intertidal platforms that started to be massively visited by Pyura gatherers from the summer of 2013. This allowed for the implementation of an intensive short‐term monitoring program of changes in tunicate cover and the abundances of their larvae and recruits. When gathering access was present the reduction in intertidal cover was generalized to the entire bay and followed by reductions in larvae and recruits. However, these reductions were not found in sites with more restricted gathering access. We conclude that continuous extraction by Pyura gatherers followed by reductions of conspecific larvae and recruits are the main drivers behind the reduced abundance of P. praeputialis in the entire bay of Antofagasta. Thus, if gathering is not stopped important ecosystem services provided by this tunicate in the bay may be threatened. Similar consequences may be expected if other massive and irreversible reductions in other species of the Pyura complex, that inhabit other coasts in the southern hemisphere, occur. The controversy concerning the impacts of invasive species and whether they cause negative, positive or neutral impacts to original ecosystems and fisheries is discussed.     

Seasonal variation in dry weight and elemental composition of the early developmental stages of Petrolisthes laevigatus (Guérin, 1835) (Decapoda: Porcellanidae) in the Seno de Reloncaví, southern Chile

In the Seno de Reloncaví, southern Chile, seasonal changes in dry weight (DW) and elemental composition (CHN) were studied in embryo (initial embryonic stage), newly hatched zoeae, and newly settled megalopae of a porcelain crab, Petrolisthes laevigatus. Samples were taken throughout the seasons of egg laying (March-December), hatching (August-February), and settlement (October–February). Values of DW and CHN per embryo or larva, respectively, were consistently minimum in the middle of each season and maximum near its beginning and end. Patterns of seasonal variation in early embryonic biomass may thus be carried over to larvae at hatching and, possibly, to the settlement stage. Such carry-over effects may be selectively advantageous, as zoeae released at the beginning or near the end of the hatching season face conditions of poor planktonic food availability in combination with low winter temperatures or decreasing temperatures at the end of summer (enforcing long development duration). Hence, an enhanced female energy allocation into egg production may subsequently translate to enhanced yolk reserves remaining at hatching, allowing for a larval development under unfavourable winter conditions. In summer, by contrast, plankton productivity and temperatures are generally high, allowing for fast larval growth and development. This coincides with minimal biomass and energy contents both at hatching and settlement. In conclusion, our data suggest that seasonal patterns in the biomass of early developmental stages of P. laevigatus may reflect phenotypic variability as an adaptive response to predictable variations in environmental conditions, allowing this species to reproduce in temperate regions with marked seasonality in water temperature and plankton productivity.     

Sediment dispersion: part 2, characterisation by size of sand fraction and percent mud

In part 2 of this contribution, examples are drawn from the River Mersey and Liverpool Bay illustrating the use of simple statistical parameters to describe dispersion of sands and muddy sediments. The River Mersey and Liverpool Bay, eastern Irish Sea, were sites of intensive studies on the dispersal of dumped harbour mud and sewage sludge during the mid 1960's–70's. The combined effects of strong tidal scour, wave action and shoreward near-bed residual drift result in shoreward transport of large volumes of sand in the bay. Large amounts of mud (silt/clay mixtures) oscillate in the river estuary, and naturally derived and dumped muds also move shoreward in the bay. Unpublished historic geochemical data have been combined with reprocessed particle size data and both have been used to reassess sedimentological techniques for defining transport and dispersal pathways. River and bay muds have similar size compositions, but river muds have excess Cd > V > U > As = Zn relative to bay muds. The lower relative concentrations of heavy metals in the bay are thought to reflect desorption and degradation of organic matter from the river. Trends in sediment distribution data based on the means of the sand size fraction, alone, provide sensitivities comparable to those of higher order moment measures and are usually easier to interpret than full size spectrum analyses.     

Seasonal density dynamics and distribution of brachyuran crab larvae (Decapoda: Brachyura) in Amursky and Ussuriysky bays, Sea of Japan

The species composition, period of occurrence, density, and distribution of brachyuran crab larvae (Decapoda: Brachyura) in the Amursky and Ussuriysky bays (Sea of Japan) were studied in May–October 2007 and 2008. The larvae of this group were not numerous, with peak densities occurring in July at maximum water temperatures. The average brachyuran larvae density was up to 126.9 ind./m³ in Ussuriysky Bay and 68 ind./m³ in Amursky Bay; the contribution of Brachyura to all decapod larvae reached 80–90%. In Amursky Bay, the highest density of brachyuran larvae (up to 584 ind./m³) was observed along the western coast over depths to 20 m; in Ussuriysky Bay, the larval density was highest (up to 1817 ind./m³) in the shallowwater northern part and progressively decreased with depth. Larvae of 18 brachyuran species were found in plankton samples. In Amursky Bay, intertidal crab larvae of the genus Hemigrapsus dominated, Eriocheir japonica larvae were fairly numerous. Larvae of the pea crabs Pinnixa rathbuni and Tritodynamia rathbunae and zoeae of the bigtooth rock crab Glebocarcinus amphioetus were predominant in Ussuriysky Bay.     

Morphogenesis of sense organs and behavioural changes in larvae of the brown-marbled grouper Epinephelus fuscoguttatus (Forsskål)

The morphogenesis of sense organs and related behavioural changes in the hatchery-reared brown-marbled grouper Epinephelus fuscoguttatus larvae were examined to gain better understanding of its early life history because ecological field observations for grouper species is difficult. The newly hatched larvae (2.1 mm total length) had developing eyes and otic vesicles, a pair of free neuromast on the head and ciliated olfactory epithelium. At 3 days post hatching (dph), the eyes became fully pigmented with pure-cone retinae, the semicircular canals formed in the inner ear, and the larvae (2.8 mm) were able to swim horizontally, preying on rotifers. Retinal rods and the intra-oral taste buds at pharyngeal appeared next. The olfactory lamellae and the head lateral line system then formed, and the inner ears developed completely in the larvae during the metamorphosis period (15–40 dph; 5.1–18.1 mm). At settlement (50 dph; 32.8 mm), the fish possessed taste buds in the mouth entrance region, and the lateral line system developed completely. The sensory development correlates well with the known aspects of its life history at sea whereby the larvae can feed early and avoid predators during the passive drift, are able to swim shoreward to search nursery ground along the metamorphosis stage and survive in seagrass beds at settlement.     

Habitat differentiation in the early life stages of simultaneously mass-spawning corals

The settlement process of coral larvae following simultaneous mass-spawning remains poorly understood, particularly in terms of population and community parameters. Here, the larval settlement patterns of Acropora corals, which are the most diverse genera of scleractinian corals at the species (haplotype) level, were investigated within a single subtropical reef. Across a 4-year period (2007–2010), the mitochondrial and nuclear molecular markers of 1,073 larval settlers were analyzed. Of the 11 dominant haplotypes of recruited populations, nine exhibited non-random patterns of settlement distribution. This result suggests that the actual habitat segregation starts during the early swimming larval stages of their life history, rather than by natural selection after random settlement. In addition, the presence of a depth-related settlement pattern supports that species-specific vertical zonation of coral larvae may play a role in the establishment of habitat segregation. Moreover, in some species that showed a preference toward the shoreward area of the bay, the settlement pattern was consistent with that of the adult distribution. This result indicates that the gametes were not mixed between fore and back reefs in the period from fertilization to settlement during the mass-spawning event, even within a single small reef. Another compatible hypothesis of this pattern is that the larvae are able to recognize various types of environmental information, facilitating the selection of optimal micro-habitats. Overall, Acropora coral larvae that are produced from a simultaneous mass-spawning event may have adapted to complex reef topography by means of multi-step habitat selection at settlement, corresponding to different spatial scales.     

The surf zone: a semi-permeable barrier to onshore recruitment of invertebrate larvae?

The supply of larvae to the shore is important for population replenishment and intertidal community dynamics but its variability at most scales is not well understood. We tested the relationship between nearshore mussel larval abundance and intertidal settlement rates over several years at multiple spatiotemporal scales in Oregon and New Zealand. Abundance of competent larvae nearshore and intertidal recruitment rates were simultaneously quantified using collectors mounted at different depths on moorings 50-1100 m from shore, and at adjacent rocky intertidal sites. Total mussel larval abundance and oceanographic conditions were also measured in some locations. At all scales, abundance of nearshore mussel larvae was unrelated to intertidal recruitment rates. In the intertidal, patterns of mussel recruitment were persistent in space, with sites of consistently high or low recruitment. In contrast, nearshore competent larval abundance showed generally similar abundances among sites except for a high, and spatially-inconsistent, variability in Oregon during 1998 only. On moorings, recruitment tended to be greater on midwater collectors than shallower or deeper. Finally, on moorings larval abundance in traps and recruitment on collectors was unrelated. These results suggest that (1) among sites, the size of the nearshore larval pool is relatively uniform while onshore recruitment varies and is unrelated to larval abundance, (2) temporal variability in nearshore larval availability is not strongly expressed onshore, (3) vertical stratification of competent larvae nearshore is strong and may influence transport and recruitment, and (4) within-coast variability in onshore recruitment is strongly driven by processes occurring locally within the surf zone that need to be studied to understand coastal recruitment dynamics.     

Spatial variation in the environmental control of crab larval settlement in a micro-tidal austral estuary

Settlement of benthic marine invertebrates is determined by the interaction between physical factors and biological processes, in which the tide, wind, and predation can play key roles, especially for species that recruit within estuaries. This complexity promotes high variability in recruitment and limited predictability of the size of annual cohorts. This study describes the settlement patterns of megalopae of the commercially important crab Cancer edwardsii at three locations (one in the center and two at the mouth of the estuary) within the Valdivia River estuary (~39.9°S), over three consecutive years (2006–2008). At each location, 12 passive benthic collectors with a natural substratum were deployed for 48?h at 7-day intervals, over a lunar cycle. Half of the collectors were covered with mesh to exclude predators. The main findings were as follows: (1) circulation changes due to upwelling relaxation or onshore winds controlled crab settlement at sites within the mouth of the estuary, (2) at the internal estuarine site, settlement was dominated by tidal effects, and (3) the effect of predation on settlement was negligible at all scales. The results show that the predominant physical factor controlling the return of competent crab larvae to estuarine environments varies spatially within the estuary. The lack of tidal influence on settlement at the mouth of the estuary can be explained by the overwhelming influence of the intense upwelling fronts and the micro-tidal regime in the study area.     

Cross-shelf dispersion of Dover sole (Solea solea) eggs and larvae in Biscay Bay and recruitment to inshore nurseries

We studied the cross-shelf dispersion of Dover sole (Solea solea)eggs and larvae from the offshore spawning grounds to the coastalnurseries of Biscay Bay in 1987. Eggs and larvae were retainedover the spawning grounds from late February to early April,consistent with the limited movement of satellite-tracked Lagrangiandrifters at different depths in the water column. The distributionof the larvae coincided with maximum zooplankton abundance.In mid April, both sole larvae and drifters moved northwardin response to wind forcing but the advection rate of the larvaewas about one-third that of the drifters. No significant onshoreadvection of the pelagic stages was observed. No evidence wasfound for a behavioural selection of tidal currents by the pelagiclarvae that could lead to onshore transport Unless that behaviourdeveloped after settlement to the bottom, cross-shelf diffusionof the pelagic stages would represent the main avenue of transportto the coastal nursery grounds. This dispersion strategy wouldimply that the vast majority of sole larvae fail to recruitto the bays and estuaries and arc lost to the population.     

Tidal movements of the macrofauna on an exposed sandy beach in South Africa

All epifaunal and shallow-burrowing, intertidal macrofauna on an exposed sandy beach appear to undergo tidal migrations which vary in extent but are all upshore on the incoming tide and downshore on the outgoing tide during day and night. In the crustaceans this is probably endogenous rhythmic behaviour and they show increased activity at night. In the molluscs this movement is not synchronised by an intrinsic mechanism but results from behavioural responses to changing physical conditions. The large wedge clam, Donax serra, shows no tidal migrations but does show a semilunar rhythm of movement from just above mean tide level (MTL) during springs to the low tide level during neap tides.     

Unravelling the Ecological Significance of Endogenous Rhythms in Intertidal Crabs

Organisms living along the shore are exposed to complex sets of environmental oscillations. In addition to solar (24.0 h) and lunar (24.8 h) cycles, local tides may reoccur on a 12.4 h schedule. Beyond daily routines, biweekly, monthly and annual rhythms may each have a significant impact on an animal's activity. For some time, it has been established firmly that intertidal crabs possess several internal biological clocks with distinctly different periods and properties. However, the versatility of these clocks has not been obvious. Crabs living in the littoral zone must adjust their internal chrono-meters to be synchronous with the specific temporal structure of the immediate habitat. Fine adjustments in their clocks will depend upon on a particular tide province and the location of their niche in the intertidal zone. Over a wide geographic range, the location of an intertidal habitat for one species may be in as many as four tidal provinces. Based on wave form and harmonic components, tide provinces are characterized as either a) semidiurnal, b) mixed, mainly semidiurnal, c) mixed mainly diurnal, or d) diurnal. Likewise, the primary frequency associated with an intertidal niche in each tide province may be augmented by diel (24 h) and semilunar (14 day) periods. In addition, supralittoral habitats may be influenced by monthly (28 day) and seasonal rhythms. Since some species live in several tidal provinces and different positions in the littoral zone, locomotor and larval release rhythms of intertidal crabs must naturally be adjusted to the timetable of the local habitat. Flexibility in ambulatory and egg hatching rhythms of crabs are discussed from this environmental perspective. The nature and location of the underlying circadian and tidal oscillators tracking these environmental rhythms are reviewed.     

The role of niche measures in explaining the abundance–distribution relationship in tropical lotic chironomids

In situ measurements of both community metabolism (primary production and respiration) and PAM fluorometry were conducted during emersion on intertidal sediments in the Mont Saint-Michel Bay, in areas where oysters and mussels were cultivated. Results highlighted a low benthic metabolism compared to other intertidal areas previously investigated with the same methods. Comparisons between gross community primary production and relative electron transport rates confirmed this statement. More specifically, primary productivity remained very low all over the year, whereas the associated microalgal biomass was estimated to be high. We suggest that the microphytobenthic community studied was characterized by a self-limitation of its primary productivity by its own biomass, as previously shown in Marennes-Oléron Bay for example. The almost permanent high biomass would represent a limiting factor for micromigration processes within the first millimetres of the sediment. This could be explained by very low resuspension processes occurring in the western part of the bay, enhanced by the occurrence of numerous aquaculture structures that could decrease tidal currents in the benthic boundary layer. Handling editor: N. Desroy