Flooding regimes increase avian predation on wildlife prey in tidal marsh ecosystems

Within isolated and fragmented populations, species interactions such as predation can cause shifts in community structure and demographics in tidal marsh ecosystems. It is critical to incorporate species interactions into our understanding when evaluating the effects of sea‐level rise and storm surges on tidal marshes. In this study, we hypothesize that avian predators will increase their presence and hunting activities during high tides when increased inundation makes their prey more vulnerable. We present evidence that there is a relationship between tidal inundation depth and time of day on the presence, abundance, and behavior of avian predators. We introduce predation pressure as a combined probability of predator presence related to water level. Focal surveys were conducted at four tidal marshes in the San Francisco Bay, California where tidal inundation patterns were monitored across 6 months of the winter. Sixteen avian predator species were observed. During high tide at Tolay Slough marsh, ardeids had a 29‐fold increase in capture attempts and 4 times greater apparent success rate compared with low tide. Significantly fewer raptors and ardeids were found on low tides than on high tides across all sites. There were more raptors in December and January and more ardeids in January than in other months. Ardeids were more prevalent in the morning, while raptors did not exhibit a significant response to time of day. Modeling results showed that raptors had a unimodal response to water level with a peak at 0.5 m over the marsh platform, while ardeids had an increasing response with water level. We found that predation pressure is related to flooding of the marsh surface, and short‐term increases in sea levels from high astronomical tides, sea‐level rise, and storm surges increase vulnerability of tidal marsh wildlife.     

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.     

Rhythmic egg-release in Littorina littorea (Mollusca: Gastropoda)

Batches of female Littorina littorea from four estuarine sites were kept in aquaria throughout their breeding season (December to July), some permanently submerged in circulating sea-water and others in a regime of simulated semi-diurnal tides of constant small amplitude. Females without the tidal experience released few eggs with little relation to the lunar month. Those in the tidal regime released more than four times the total number of eggs and released far more in periods of new and full moon than in periods of half-moon. Over the whole season, release was 1–5–3–25 times faster (at different sites) during full and new moon but the difference was more accentuated at the height of the season in February to early March and in early May. While individual females differed considerably and there were minor variations between the four sites, it is concluded that a two-weekly rhythm relates egg release to the spring-neap tide cycle and that this rhythm requires a tidal cycle of immersion and emersion for its maintenance. The special significance of such a rhythm for a species with pelagic eggs and larvae in an estuary is discussed.     

长江口低盐淡水区潮间带鱼类群落结构季节及半月相变化

基于2006年春（5月）、夏（7月）、秋（9月下旬）和冬季（2月）对长江口低盐淡水区水域进行的潮间带鱼类调查结果,分析了鱼类种类及丰度的半月相及其季节变化. 结果表明：长江口低盐淡水区水域潮间带鱼类共有53种;其中鲤科鱼类最多,虾虎鱼科次之,其他科较少. 2顶网具渔获物的鱼类种类和丰度均以春、夏、秋季较高,冬季最低.各个季节的平均种类数和平均丰度均为小潮期小于大潮期. Shannon-Weaver指数（H′）在一个潮周期内的波动幅度较大,且小潮期H′值略低于大潮期. 对鱼类种类丰度进行聚类分析发现,冬季鱼类聚成一组,春、夏和秋季聚成一组. 从整年的情况来看,潮间带鱼类的丰度变化与水温的相关性最高;与潮高的相关性次之;而与盐度的相关性最小.     

Some features of ocean currents in Gulf of Nicoya, Costa Rica

The spatial and vertical structure of the water currents and its relationship with the tidal cycles were studied using current meters in the Gulf of Nicoya. In the upper gulf, the vertical marine current differences increase as the depth increases. The water column at the station near Chira Island (upper gulf) shows the smaller changes in currents and in temperature. The flow at the station between San Lucas Island and Puntarenas (middle gulf) is the most stratified in this region. Currents with magnitudes over 100 cm/seg were measured during spring tides. Changes in the lags of the surface and bottom tidal flows were measured on the order of 100 minutes. In general, in this upper region the flows are toward the head of the gulf when the tide is flooding and toward the mouth when the tide is ebbing. In the lower gulf the circulation is more complex. Along an axis between Tárcoles and Negritos Islands, changes of velocity vectors are identified between surface and bottom. The current rotates in a different way in the water column in this region and their patterns cannot be explained only by the tidal cycles predicted for Puntarenas. These results demonstrate that the spatial and vertical variation of the marine currents of the Gulf of Nicoya is not only related to the thermohaline structure, but also to the tidal cycles and tide ranges that take place in this estuary.     

The influence of diel cycle,tidal direction and trawl alignment on beam trawl catches in an equatorial estuary

Synopsis Using a 3 m beam trawl, catch variation in the fish and prawn assemblage of the Labu estuary, Papua New Guinea was investigated during July and December 1981 and September 1984. About six times more organisms, three and a half times the biomass and twice as many species were caught per trawl at night than during the day. The greatest number of species, individuals and biomass were caught just after dusk. With the exception ofSecutor ruconius, there was no significant difference in the catches trawled at ebbing and flooding tides. Over twice as many individuals ofS. ruconius were caught during ebbing tides. Alignment of trawl direction in relation to the tidal current did not have a significant effect on the fish catch, but affected the prawn (Metapenaeus demani) catch. It is postulated that most species are not carried to and fro by tidal currents, but maintain their position in the estuary with respect to the substratum. Only prawns appeared to be carried by the currents. This study indicates that the most representative trawl results in terms of species diversity and abundance in shallow estuaries may be obtained at night and, that in estuaries with a small tidal range, tidal direction and velocity have little influence on catch. It also indicates that analyses of demersal nekton assemblages from daytime only estuarine trawl surveys must be approached with caution.Lizard Island Research Station     

Foraging strategy of the mudskipper Periophthalmus sobrinus Eggert in a Kenyan mangrove

The feeding behaviour of Periophthalmus sobrinus was studied in a population living along the channels of the Tana river delta. In November 1993 mudskippers were sampled every 2 tidal h for two consecutive spring and neap tides both during the day and night. The fishes were sexed and their stomach contents analysed. The results indicated that the mudskipper has a carnivorous diet and forages during both day and night although feeding greatly decreases after dark. Quantitative and qualitative differences in diet were found during the day and night of the two synodic and tidal phases. These differences were also demonstrable in the two sexes. The study showed that females of P. sobrinus are more selective than males and that differences between the two sexes may be related to reproductive condition. Furthermore, the mudskippers showed a tidal rhythm in feeding behaviour with peaks of activity around low tide (LT) on spring tides and around high tide (HT) on neap tides. Foraging activity seems to be influenced by the different spatial and temporal distributions of the mudskipper's prey caused by synodic, tidal and diel changes in the environment.     

The turbidity maximum in the Tamar estuary

Many estuaries of medium to high tidal range exhibit an accumulation of fine cohesive material in their upper reaches in the region of the limit of saline intrusion. Much, or all, of this material is suspended each tidal cycle and the entire region undergoes a seasonal variation which appears to depend on fluvial input. Two factors which are throught to influence the formation and maintenance of turbidity maxima are the differing magnitudes of the bed shear stress (τ₀) on flood and ebb tides and the large vertical density gradient which developes on the ebb tide. Crucial to the importance of the first factor is that τ₀ exceeds a critical value, at which erosion occurs, for a greater period on the flood than on the ebb. The effect of the density gradient is that upward propagation of bed generated eddies is inhibited and the sediment is not transported into the upper part of the flow where it will be most effectively transported. It is not clear which, if either, of these mechanisms is dominant. Data consisting of vertical profiles of velocity, salinity and suspended solids were collected at four stations in the Tamar estuary during a high range tidal cycle. One station, at which the depth mean salinity (S _d ) varied from 0.0 to ∼ 12.0‰, was occupied permanently. The other stations were occupied such that data were collected asS _d varied in the range 0.0 to ∼ 4.0‰. In this way each station was occupied for a period of time on the ebb and flood tide. Observations show that during the early ebb, when the flow is relatively deep and slow, stratification persist untilS _d ∼ 0.0‰ and that no significant transport occurs while the flow is saline but that there is a rapid increase in suspended solids concentrations after this time. During the later ebb the shallower faster flow allowed the density gradient to be erode and significant transport was observed atS _d ∼ 5.0‰. On the flood tide the flow in the low salinity region is well-mixed troughout. Computation of the fluxes and total transport per unit breath of estuary show that on the ebb tide the quantity of solid material being transported by the low salinity (0–3‰) region remains nearly constant as this region of the flow is advebted seaward. On the flood tide, however, as the same region is advected landward the quantity of material being transported increases. It is concluded that in the Tamar estuary the early ebb tide stratification contributed to the formation and maintenance of a turbidity maximum which is strongly associated with the low salinity region of the flow. It is also speculated that the differences in the ebb and flood tide transport are caused by differences in the availability of mobile material on the bed at different stages of the tidal cycle.     

Tidal and diurnal variations in larval fish abundance in an estuarine inlet in Ariake Bay,Japan: implication for selective tidal stream transport

We have conducted a preliminary study of tidal and diurnal variations in the distribution of dominant larval and juvenile fishes in the Chikugo River inlet (Ariake Bay, Kyushu, Japan) to determine whether selective tidal stream transport (STST) occurs. Larval and juvenile fish were collected from the mesohaline zone of the Chikugo River inlet during spring 2002. Temperature, salinity, depth, and current velocity were measured. Larval and juvenile abundance were compared among four tidal conditions, flooding tide, high tide, ebbing tide, and low tide, and between day and night. A total of 12 families, 15 species, and 5,577 individuals were collected. Temperature did not vary significantly with tidal conditions whereas salinity, depth, and current velocity varied significantly. Salinity also was correlated significantly and positively with depth. The abundance of most of the fishes was correlated positively and significantly with salinity and depth. Lateolabrax japonicus, Trachidermus fasciatus, Acanthogobius hasta, and other gobiid larvae (Gobiidae spp.) were significantly more abundant during high tide; in contrast, Coilia nasus and Neosalanx reganius were most abundant during low tide. The abundance of most of the fishes was higher during high tides at night than during the day, indicating the existence of STST, which may be strategically associated with ascending progress to upstream nursery areas.     

Tidally driven distribution of phytoplankton blooms in a shallow, macrotidal estuary

The influence of the ebb tide on the abundance and distributionof bloom-forming species, as well as the mechanisms for theselection of those species which remain, were investigated inthe shallow, tidal-flushed Urdaibai estuary, north Spain. Phytoplanktonwas collected monthly from May to September 1998 during differenttidal conditions (neap-spring) at five stations along the salinitygradient of the estuary. During the neap tides of May, Leptocylindrusdanicus dominated in the lower estuary, together with Thalassiosiraguillardii and Peridinium foliaceum in the upper segment; T.guillardiiexperienced the broadest displacement along the estuary. Duringthe June and July cruises, coinciding with mid-tidal amplitudesand high temperatures, Peridinium quinquecorne densely aggregatedin the upper estuary at the slack high tide, whereas Chaetocerossalsugineum bloomed at the intermediate stations. Cyclotellaatomus and Protoperidinium achromaticum reached high concentrationsin the upper zone only during the late stages of the ebb. Duringthe spring tides of September, Prorocentrum minimum, Heterocapsapygmaea and Heterocapsa rotundata appeared in the lower marineestuary, being washed out at low tide. By contrast, the diatomsSkeletonema costatum, Thalassiosira guillardii and Cyclotellaatomus were the most abundant species in the upper reaches,peaking during the ebb. Physical trapping and high water residencetimes served to retain blooming species in the upper estuary.The intense growth of the estuarine diatoms may compensate forthe advective seaward losses of cells during the ebb, thus allowingthe development of stable populations in the estuary. Only Peridiniumquinquecorne seems to combine an endogenous tidal rhythm witha photic response to remain in the tidal area of the estuary.     

Tidal amplitude and fish abundance in the mouth region of a small estuary

Using an acoustic underwater camera (Dual Frequency IDentification SONar, DIDSON), the abundance and direction of movement of fishes > 80 mm total length (L_T) in the mouth of a small South African estuary during spring and neap tidal cycles were observed. While the sizes of fishes recorded were consistent across both tide cycles, the number of fishes passing the camera was significantly greater during the smaller neap tides. Schooling behaviour was more pronounced for fishes that were travelling into the estuary compared to fishes swimming towards the ocean.     

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

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

Meiofaunal emergence from intertidal sediment measured in the field: significant contribution to nocturnal planktonic biomass in shallow waters

Field studies on the occurrence of meiobenthos in the water column above intertidal sandflats have been performed near the Island of Sylt in the northern Wadden Sea. Swimming meiobenthos was strongly dominated by harpacticoid copepods. Many of them have a semiplanktonic life-style. They rest in superficial sediment layers at low tide and swin in the water column at high tide. Swimming activity correlated negatively with light. The abundance in the water column was one order of magnitude higher during the night. Strong currents caused by storm tides significantly decreased meiobenthic abundance in the water column. Light and flow being constant, no significant changes of meiobenthic abundance per unit area occurred over a tidal cycle. Since holoplankton and meroplankton abundances correlated positively with the height of the water column, semiplanktonic meiobenthos may dominate the mesozooplankton in shallow waters. On an average, emergence of meiobenthos increased the mesozooplanktonic biomass by about 2% during diurnal high tides over the entire tidal cycle, and by about 50% during nocturnal high tides. Because of seasonal cycles of the dominant harpacticoids, this high contribution to planktonic biomass may be a summer phenomenon.     

Movement of adult Atlantic salmon in the Usk estuary, Wales

A total of 56 salmon was tagged in the Usk estuary using combined acoustic and radio tags. Those fish migrating within the estuary oscillated with the tide over c . 10 km, being towards the seaward end at low water and moving upstream on the flood tide. Fish migrating through the estuary moved upstream on the flood tide and stemmed displacement downstream during the ebb. These findings, together with information on the hydrodynamics of the estuary, indicate that the fish utilize tidal currents to migrate passively in their preferred direction.     

我国近海赤潮多发区域及其生态学特征

根据有关的研究资料,分析我国近海赤潮多发区分布、主要赤潮生物种类、环境状况及赤潮发生生态学特点.结果表明,我国辽东湾、渤海湾、莱州湾、大连湾、长江口、舟山海域、杭州湾、厦门湾、柘林湾、大鹏湾、珠江口等为赤潮多发区.引发赤潮的因素较多,它与气象、水动力、营养盐及生物环境的变化密切相关,人类活动(如海水养殖、陆源污水排放等)影响加剧近海富营养化是引发赤潮的重要因素;但富营养化并非发生赤潮的唯一条件,低营养海区也可能发生赤潮.目前对诱发赤潮的关键因子及赤潮发生机理,应加强定点长期监测和开展深入研究.     

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.     

Temporal Occurrence and Spatial Distribution of Red Tide Events in China's Coastal Waters

Between the years 1933–2001, 460 red tide events were found in China's coastal water. The scope of red tide occurrence has extended over all the coastal provinces of China, and the three major areas with high red tide frequency are the Bohai sea, the sea area near the Shengsi Archipelago and Huaniaoshan Island on the outer side of the Changjiang River estuary and the Hangzhou Bay, and the coastal waters on the east side of the Zhujiang River estuary from Hong Kong to East Guangdong. In the past 20 years, the frequency of red tides has been tending upwards. The years of 1988–1990 and 1998–2001 saw the most serious red tides along China's coastal waters, with the latter period as the peak. The average area of a single red tide, i.e., the scale of red tides, is expanding every year, and in 2001 rose to about 500 km². Every year, China's red tides occur from south to north, with those in the South China Sea occurring from March to May, those in the East China Sea from April to August and those in the Bohai and Huanghai Seas from May to September.     

Changes in bacterial populations during red tides caused by Mesodinium rubrum and Gymnodinium catenatum in North West Coast of Spain

Heterotrophic bacterial communities associated with four red tides caused by Mesodinium rubrum and Gymnodinium catenatum in two Galician Rias (North West Spain) were examined. Three of these were produced by the Mesodinium rubrum and the causative organism of a toxic bloom was Gymnodinium catenatum . In early stages of all the blooms, the diversity decreased but the total marine bacterial counts increased by one or two logs. Vibrio numbers were also incremented by two logs in two blooms of M. rubrum , while in the other bloom of this organism and in the red tide caused by G. catenatum a decrease in number of these bacteria was observed. A total of 116 bacterial strains were identified at the genus level and grouped into 12 phena. During the decomposition processes of two blooms of M. rubrum a zooplanktonic-type bacterial succession was observed ( Vibrio , pseudomonads and Moraxella-Acinetobacter ). On the other hand, during decomposition of the other red tide of M. rubrum and the bloom of G. catenatum , a typical phytoplanktonic-type succession occurred, as Pseudomonas and Moraxella groups became dominant for all the process. These results support the conflicting taxonomical position of M. rubrum . After the blooms, the changes in the community point towards the restablishment of the normal bacterial flora of the estuary (increase in diversity and decreases of bacterial numbers). Only the Vibrio strains, isolated from the non-toxic first and second red tides, displayed cytotoxic activities. A relationship among bacterial cytotoxicity and toxic effects of blooms cannot therefore be established.     

The response of salt marsh vegetation to tidal reduction caused by the Oosterschelde storm-surge barrier

In 1986 a sluice gate barrier was completed in the mouth of the Oosterschelde estuary. The barrier has been partially or completely closed during 1986 and the first months of 1987. Consequently the high tides were reduced to such a level that the salt marshes were scarcely flooded. Since April 1987 the barrier has been closed on average during two out of 706 high tides a year. Although the barrier allows tidal exchange the tidal flow has been restricted as a result of the reduced width of the mouth of the estuary. This restriction of the tidal flow caused a 26 cm decrease of mean high water in the eastern part of the estuary. As a result the inundation frequencies of the salt marshes decreased.The response of salt-marsh vegetation to this tidal reduction was analyzed using annual records (1982 till 1990) of species composition in 57 permanent plots in two marshes at the southern shore of the estuary. Analysis of the response of individual species to marsh elevation in the pre and the post-barrier situation revealed that most species moved down the marsh elevation gradient. The first axis of an ordination (DCA-1) was significantly negatively related to inundation frequency. Between 1984 and 1990 all plots were displaced towards a significantly higher ordination score, indicating a trend towards a species composition from higher up the marsh. The position of most plots along DCA-1 remained stable until 1985 and started to increase in 1986 or 1987. The vegetation in plots dominated either by Halimione portulacoides or by Spartina anglica, started to change in 1985. This premature change was attributed to frost damage in January 1985. The initially high rate of change along DCA-1 decreased in 1989 and 1990. This would suggest that the vegetation re-equilibrated with the newly established tidal conditions. Further analysis revealed no significant difference in the relation between inundation frequency and sample score along DCA-1 between 1984 and 1990. This corroborates the view that species composition had re-equilibrated with the tidal conditions. Along DCA-2 the samples were displaced towards a significantly higher score as well. The change was attributed to an increase of the perennial Halimione and the annual Suaeda maritima. The annual Atriplex hastata displayed an increase from 1986 till 1988, but strongly declined in 1990. The transient response of this annual was described by DCA-3.During the pre-barrier phase several attempts had been made to predict the vegetation response to tidal reduction. We developed a multivariate model (CCA) with inundation frequency as a constraining factor to describe the relation between inundation frequency and species composition in the pre-barrier phase in 1984. Next we used this model to predict the score of the samples along CCA-1 in the post-barrier phase from the inundation frequencies in 1990. The actual ordination score in 1990 was calculated from the observed species composition. The predicted ordination scores did not differ significantly from the observed ones. In retrospect it is concluded that the response of the vegetation to the reduction of the tides, as far as the fraction of species composition which is related to inundation frequency is concerned, could have been predicted by our model.

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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.