The intertidal movements and distribution of young fish on a sandy beach with special reference to the plaice (Pleuronectes platessa L.)

The intertidal movements of young fish were studied on a sandy beach over several tidal cycles during the day and the night. Young plaice (Pleuronectes platessa L.), by virtue of their abundance, received the most attention, and it was found that they migrate up and down the beach with each tide in company with several other less abundant species. The dab (Limanda limanda (L.)), does not make this migration.The vertical distribution of the commoner fish species was charted and each was found to occupy a specific zone in relation to the water, but with large overlaps in many cases. A close relationship between the length of plaice and dabs and the depth in which they were caught was evident during the summer months, but this relationship broke down in the winter due mainly to an offshore movement of larger fish.The food intake of plaice was studied throughout the day and night; feeding occurred mostly during the day. No effect of the tide on feeding could be detected.The possible mechanisms controlling the tide-related movements and the size-depth relationship are discussed and a general hypothesis is put forward in which it is proposed that the fish are able to select and maintain their position at a particular depth throughout a tidal cycle thereby being guided up and down the beach. Attributes of depth which the fish might use as a reference point in their movements are discussed.     

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.     

Effets des marées sur la variation circadienne de la capacité photosynthétique du phytoplancton de l'estuaire du saint-laurent

The photosynthetic capacity of phytoplankton was measured at an anchor station in the Middle Estuary of the St Lawrence, over a period of 120 h. Hourly incubations were made in constant light conditions, under three different colour filters of 3, 4 and 23 W · m^?2. Differences in the oscillations of the photosynthetic capacities were observed in relation to the tidal regime. During spring tide, variations in the photosynthetic capacity are circadian, whereas a tidal pattern was also observed during the neap tide period. Differences in the patterns of photosynthetic capacity were probably due to changes in the gradients of physical factors and to the physiological state of phytoplankton relative to these gradients. The circadian rhythms appeared to be endogenous since they were apparent at non-saturating light intensities and since the photosynthetic potential (U_max) changed during the day.     

Movements of intertidal gastropods

Movements of marked individuals of Nerita atramentosa Reeve, Austrocochlea constricta (Lamarck), Bembicium nanum (Lamarck), and Cellana tramoserica (Sowerby) were recorded at various sites in New South Wales in 1974 and 1975. A model of random movements during feeding at high tide was proposed, i.e., that animals wandered in random directions and the distances moved were exponentially distributed. Appropriate statistical tests for departures from such a model are given. The types of departure tested were non-random directions, non-exponentially distributed dis- tances, different distances in different directions, and correlated distances or directions of movement between days. With two exceptions, the movements of the animals over short periods were adequa- tely described by the assumption of random movements. One exception was A. constricta in one locality where movements of the sample were limited by it being in a large rock-pool and the other was the limpet C. tramoserica which displayed homing behaviour, so that the population could be divided into homers and non-homers. Movements of non-homing limpets were random.The mean distance moved by N. atramentosa was not significantly different from that of A. con- stricta. The mean distances moved by these two species were greater than those of B. nanum and the non-homing C. tramoserica, which were not different from each other. Adult and one-year old N. atramentosa did not differ significantly in mean distance moved, but both moved more than juveni- les. These patterns of movement are discussed with respect to hypotheses about feeding in these species.The model of movement was used to predict the effect of different percentage cover in small pools on the movement of A. constricta, which shows a preference for this micro-habitat. Increased availability of small pools reduced the mean distance moved by this species.     

Effects of Natural and Anthropogenic Change on Habitat Use and Movement of Endangered Salt Marsh Harvest Mice

The northern salt marsh harvest mouse (Reithrodontomys raviventris halicoetes) is an endangered species endemic to the San Francisco Bay Estuary. Using a conservation behavior perspective, we examined how salt marsh harvest mice cope with both natural (daily tidal fluctuations) and anthropogenic (modification of tidal regime) changes in natural tidal wetlands and human-created diked wetlands, and investigated the role of behavioral flexibility in utilizing a human-created environment in the Suisun Marsh. We used radio telemetry to determine refuge use at high tide, space use, and movement rates to investigate possible differences in movement behavior in tidal versus diked wetlands. We found that the vast majority of the time salt marsh harvest mice remain in vegetation above the water during high tides. We also found no difference in space used by mice during high tide as compared to before or after high tide in either tidal or diked wetlands. We found no detectable difference in diurnal or nocturnal movement rates in tidal wetlands. However, we did find that diurnal movement rates for mice in diked wetlands were lower than nocturnal movement rates, especially during the new moon. This change in movement behavior in a relatively novel human-created habitat indicates that behavioral flexibility may facilitate the use of human-created environments by salt marsh harvest mice.     

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.     

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.     

Physiological performance of intertidal coralline algae during a simulated tidal cycle

Intertidal macroalgae endure light, desiccation, and temperature variation associated with sub‐merged and emerged conditions on a daily basis. Physiological stresses exist over the course of the entire tidal cycle, and physiological differences in response to these stresses likely contribute to spatial separation of species along the shore. For example, marine species that have a high stress tolerance can live higher on the shore and are able to recover when the tide returns, whereas species with a lower stress tolerance may be relegated to living lower on the shore or in tidepools, where low tide stresses are buffered. In this study, we monitored the physiological responses of the tidepool coralline Calliarthron tuberculosum (Postels and Ruprecht) E.Y. Dawson and the nontidepool coralline Corallina vancouveriensis Yendo during simulated tidal conditions to identify differences in physiology that might underlie differences in habitat. During high tide, Corallina was more photosynthetically active than Calliarthron as light levels increased. During low tide, Corallina continued to out‐perform Calliarthron when submerged in warming tidepools, but photosynthesis abruptly halted for both species when emerged in air. Surprisingly, pigment composition did not differ, suggesting that light harvesting does not account for this difference. Additionally, Corallina was more effective at resisting desiccation by retaining water in its branches. When the tide returned, only Corallina recovered from combined temperature and desiccation stresses associated with emergence. This study broadens our understanding of intertidal algal physiology and provides a new perspective on the physiological and morphological underpinnings of habitat partitioning.     

Effects of Tidal and Diel Cycles on Dugong Habitat Use

ABSTRACT Quantifying the factors influencing behaviors of aquatic mammalian grazers may enhance the generic understanding of grazer ecology. We investigated diel and tidal patterns in movements of the dugong (Dugong dugon) by Global Positioning System—tracking 12 animals in 5 inshore—intertidal and 3 offshore—subtidal habitats along the coast of Queensland, Australia. We examined effects of tide height and time of day on the dugong's distance from 1) the nearest coast, 2) water >3 m deep, 3) actual water depth (bathymetry + tide ht) experienced, and 4) distribution of the directions of movements. Both tidal and diel cycles influenced dugong movement. Tracked dugongs tended to be closer to shore at high tide than at low tide and closer to shore at night than during the day. Onshore movement was more prevalent on incoming tides and in the afternoon and evening. Offshore movement was more prevalent on outgoing tides and from midnight through the morning until midday. Tidal and diel variation in water depths used by the inshore—intertidal dugongs was small, but probably underestimated, hidden by a sampling bias in the telemetry equipment. Onshore movement at high tide allowed dugongs to exploit intertidal seagrass beds. Dugongs are closer to shore in afternoons and evenings than in mornings. This behavior may be related to the avoidance of predators or watercraft. Our findings can be used to predict spatial patterns of dugongs within areas of conservation management significance and to assess, avoid, and mitigate adverse effects of anthropogenic disturbance.     

Settlement patterns of Chthamalus spp. at Praia da Oliveirinha (SW Portugal)

Recruitment of Chthamalus spp. is intense during spring and summer but the number of settled cypris larvae is very low when compared to the high number of Chthamalus spp. metamorphs. In the summer of 1997 at Praia da Oliveirinha (SW Portugal), two hypotheses were tested: 1) the cypris larvae of Chthamalus spp. metamorphose in one tidal cycle; and 2) the settlement of Chthamalus spp. is higher during diurnal high tide compared with nocturnal high tide. The results indicate that cypris larvae of Chthamalus spp. can metamorphose in one tidal cycle. The hypothesis of higher settlement during diurnal high tide could not be tested at the mid and upper levels of C. montagui distribution due to a significative heterogeneity of variances. At the lower level, the results support this hypothesis. In spite of these data having been collected on a single shore and on a few sampling dates, these results suggest that more studies using different spatial and temporal scales are needed to see if this pattern is general.     

VERTICAL MIGRATION OF A MIXED‐SPECIES EUGLENA (EUGLENOPHYTA) ASSEMBLAGE INHABITING THE HIGH‐INTERTIDAL SANDS OF NYE BEACH,OREGON1

Comparatively little is known about the vertical migration of the microphytobenthic community forming visible patches on high‐energy beaches. We collected surface and cored samples to evaluate the timing and extent of downward migration of a multispecies Euglena assemblage inhabiting Nye Beach, Oregon. Euglena density at the surface was highly variable and was not correlated with the time of low tide or instantaneous irradiance measurements; however, triplicate cores collected at low and high tides revealed a tidal rhythm in mean depth. On average, 95% of the assemblage occurred within 1 cm of the surface during low tide, but 54% of the assemblage was collected between 1 and 8 cm below the surface during high tide. A midday shading experiment revealed that short‐term changes in irradiance levels altered the Euglena density at the sediment surface by inducing vertical migration. This response to short‐term fluctuations in light may explain the weak correlation between cell density at the surface and time of day. The high‐intertidal location of these patches prevented the removal of nonmigrating cells by daily high tides, which increased the variability in surface samples and obscured the tidal migration rhythm detected in the core samples. Due in part to the semidiurnal nature of Oregon tides, this study provides in situ confirmation of past mesocosm research indicating that sediment disturbance during daily submersed periods is an important process in maintaining the quasi‐tidal rhythm in the appearance and disappearance of Euglena spp. from the surface of beaches and intertidal sandflats.     

Community Structure of Coastal Fishes in Relation to Heavily Impacted Human Modified Habitats

Diversity and community structure of coastal fishes were compared between reforested mangrove and reclaimed sandy habitats at Pasir Ris, in the eastern part of Singapore. Both habitats supported a total of 91 species, but a single species, Ambassis kopsii dominated in abundance. Mean fish density was significantly higher at the reclaimed sandy shore than the reforested mangrove, but the reforested mangrove habitat yielded higher diversity. Apart from A. kopsii, abundant species in the mangroves included Ambassis interrupta, Thryssa hamiltonii, Acentrogobius sp., Scatophagus argus and Arius sagor. Anodontostoma chacunda, Acentrogobius sp., Leiognathus decorus, Sillago sihama and Stolephorus sp. were abundant in the reclaimed sandy habitat. Both habitats exhibited a higher mean density and average species richness during high tide. Several species from the family Gobiidae dominated the catch during low tide while pelagic fishes were most abundant during high tide. Diel variation showed significantly higher density during the day than at night, a result that contrasts to similar studies elsewhere. However, species richness showed no significant variation between day and night catches. Interaction between combined factors of time of the day and tidal height on fish density and average species richness was significantly different. The results indicated both habitat types to be important fish nurseries.     

Evidence for female-biased juvenile dispersal in corophiid amphipods from a New Zealand estuary

This study examined the distribution and dispersal of corophiid amphipods (Paracorophium spp.) in a temperate New Zealand estuary. Individuals in the sediment (resident population) and those caught in bed-load (local-scale movement) and suspended-load (larger-scale movement) were counted, measured, and sexed, during two tidal cycles at 4-6 week intervals spanning 13 months. Gravid females and juveniles were present in the resident population on all sample dates suggesting near continuous reproduction. Juveniles (< 2.2 mm body length) accounted for 49.8% of individuals in the sediment, compared to 65.4% of those caught in the bed-load and suspended-load traps. For the majority of sample dates, there was a net export of juveniles from the estuary. The total number of juveniles captured in suspended-load traps was significantly greater during the ebb tide relative to the flood tide (n = 541 versus 256), whereas no significant differences were found for adults. Similarly, numbers in the bed-load traps were significantly greater for juveniles during the ebb versus the flood tide (n = 900 versus 484), and also for adult females (n = 180 versus 86). Juvenile sex ratios were female biased in the sediment (9.8:1) and in suspended-load traps (7.6:1) and both were significantly more female-biased than that recorded for juveniles caught in bed-load traps (5.8:1). By contrast, there was a significantly lower proportion of adult females to adult males found in bed-load traps (1.7:1) and in suspended-load traps (0.98:1) compared to the resident population in the sediment (3.6:1). The mean size of juvenile females caught on a flood tide was significantly larger compared to those on the ebb tide, yet juvenile males were not significantly different, suggesting that smaller females were not returning on the flood tide. We suggest that female-biased juvenile dispersal for estuarine Paracorophium spp. lowers the risk of inbreeding and reduces competition. This may have evolved in response to a polygamous breeding system with little or no social organisation where a female produces more female offspring to maximise her inclusive fitness.     

Capacity of a temperate intertidal seagrass species to tolerate changing environmental conditions: Significance of light and tidal exposure

Seagrass meadows perform several important ecological roles in coastal areas. However, with multiple stressors threatening these aquatic plants, their current rate of decline is likely to increase, so understanding their ability to acclimate to a range of environments may be key to their survival. Light availability is critical for seagrass health, with loss of water clarity likely to cause decline. This study assessed the ability of the temperate intertidal species, Zostera muelleri, to compensate for variations in light in its natural environment. The impacts of light during inundation (high tide) and exposure (low tide) were examined along a vertical gradient from the high to low intertidal at two sites with differing light histories. Photo-acclimation was evaluated through morphological and physiological characteristics over two tidal cycles. Results were consistent with seagrasses having optimized their photosynthetic capacity, with physiological acclimations being site specific. Longer-term morphological changes were also noted, suggesting a dissimilar light history between sites for an extensive period. Interestingly, at the site with the greatest light range, but the lowest light penetration at depth, a significant reduction in photosynthetic activity during air exposure was observed. This suggests that Z. muelleri is capable of tolerating light stress through photo-acclimation, allowing for the more efficient harvesting of light at low levels. Overall, this study demonstrates that Z. muelleri has the ability to adjust both physiologically and morphologically to changing environmental conditions, a key aspect to survival and persistence in temperate intertidal zones.     

Entrainment of the activity rhythm of the mole crab Emerita talpoida

The mole crab Emerita talpoida migrates with the tide in the swash zone of sand beaches. A circatidal rhythm in vertical swimming underlies movement, in which mature male crabs show peak swimming activity 1-2 h after the time of high tides at the collection site. In addition, there is a secondary rhythm in activity amplitude, in which crabs are maximally active following low amplitude high tides and minimally active following high amplitude high tides. The present study determined the phase response relationship for entrainment of the circatidal rhythm with mechanical agitation and whether the cycle in activity related to tidal amplitude could be entrained by a cycle in the duration of mechanical agitation at the times of consecutive high tides. After entrainment with mechanical agitation on an orbital shaker, activity of individual crabs was monitored in constant conditions with a video system and quantified as the number of ascents from the sand each 0.5 h. Mechanical agitation at the times of high tide, mid-ebb and low tide reset the timing of the circatidal rhythm according to the timing relationship to high tide. However, mechanical agitation during flood tide had no entrainment effect. In addition, a cycle in duration of mechanical agitation entrained the rhythm in activity amplitude associated with tidal amplitude. Both rhythms and entrainment effectiveness over the tidal cycle may function to reduce the likelihood of stranding above the swash zone.     

Diel Fluctuations in the Abundance and Community Diversity of Coastal Bacterioplankton Assemblages over a Tidal Cycle

The diel change in abundance and community diversity of the bacterioplankton assemblages within the Pacific Ocean at a fixed location in Monterey Bay, California (USA) were examined with several culture-independent (i.e., nucleic acid staining, fluorescence in situ hybridization {FISH}, and 16S ribosomal RNA gene libraries) approaches over a tidal cycle. FISH analyses revealed the quantitative predominance of bacterial members belonging to the Cytophaga-Flavobacterium cluster as well as two Proteobacteria (α- and γ-) subclasses within the bacterioplankton assemblages, especially during high tide (HT) and outgoing tide (OT) than the other tidal events. While the clone libraries showed that majority of the sequences were similar to the 16S rRNA gene sequences of unknown bacteria (32% to 73%), however, the operational taxonomic units from members of the α-Proteobacteria, Bacteroidetes, Firmicutes, and Cyanobacteria were also well represented during the four tidal events examined. Comparatively, sequence diversity was highest in OT, lowest in low tide, and very similar between HT and incoming tide. The results indicate that the dynamics of bacterial occurrence and diversity appeared to be more pronounced during HT and OT, further indicative of the ecological importance of several environmental variables including temperature, light intensity, and nutrient availability that are also concurrently fluctuating during these tidal events in marine systems.     

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

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

Aerial and underwater metabolism of <Emphasis Type="Italic">Patella vulgata</Emphasis> L.: comparison of three intertidal levels

Intertidal molluscs are known to possess specific respiratory organs that permit aerial breathing during emersion. Patella vulgata is a widely distributed intertidal species found from low-water spring tide to high-water neap tidal level. In order to determine metabolic adaptations to habitat, carbon fluxes associated with respiration and calcification of P. vulgata living at high-shore, middle-shore and low-shore levels were compared. Seasonal aerial respiration was measured using an infrared gas analyser; seasonal underwater respiration and calcification were calculated from dissolved inorganic carbon and total alkalinity. P. vulgata showed net CaCO₃ deposition at all seasons, although the high-shore level limpet annual calcification rate was relatively low due to longer air exposure. Both aerial and underwater respiration rates were highly correlated with seasonal temperature variations and followed the vertical shore gradient, with stronger fluxes for low-shore tidal level limpets and lower fluxes for high-shore level limpets that must limit energy expenditure. P. vulgata appears to be well adapted to aerial exposure, with average hourly respiration fluxes stronger in air than in water. This study demonstrates that P. vulgata calcification and respiration are reduced in upper shore levels and are important factors determining the upper distribution limit of the species.     

Kulturverfahren zur Bestimmung der Salz- und Über-flutungsverträglichkeit vonPuccinellia spp. (Gramineae)

In the ecology ofPuccinellia species (P. maritima, P. distans, P. retroflexa andP. limosa of the German flora described to date) two environmental factors seem to be very important: the salinity of the soil or of the tidal sea water and, especially in regard toP. maritima, the periodical inundation during the tidal cycle. The latter species forms an important community (Puccinellietum maritimae) a few decimetres above the middle tide level. The other species seem to be specialized to localities of higher levels. Some 450 natural localities covered withP. maritima on the western coasts of Schleswig-Holstein (W. Germany) have been analysed with respect to the type of plant community, dominance ofP. maritima, frequency of inundations, level above or below middle tide level, and soil factors (for instance: Cl-ion content, electrical conductivity, pH, clay content, etc.). Physiological experiments have been carried out in greenhouses, experimental grounds, and culture-rooms to test the effects onP. maritima (number 57 from the optimal level within the Puccinellietum maritimae of the banks of the River Eider) of soil composition, light intensity and daylength, temperature, inundation, etc.P. maritima grows with long creeping stolons at daylengths of 16 hrs and more; it forms a caespitose tufted plant at daylengths of 12 hrs and less (light intensity: 10,000 lux and more). Optimal growth occurs between 17° and 23° C and under conditions of artificial flooding up to soil surface twice a day (for about 10 minutes) by artificial sea water of 25 ‰ S. Two culture techniques have been developed for testing growth and survival after 8 weeks under greenhouse conditions (20° C or more; 10,000 lux or more; 15 hrs daylength or more, after comparable pre-cultivation over 8 weeks under 12 hrs day and artificial light conditions; sand + 3 % peat). The first technique has been used for testing salt tolerances between 0 and 60 ‰ S (artificial sea water), the second for testing inundation tolerances (0–2.5 hrs inundation twice a day; artificial sea water of 25 ‰ S). Considering different proveniences ofPuccinellia species on German coasts, it became clear thatPuccinellia exhibits broader ecological and morphological than taxonomical variabilities. WithinP. maritima, for example, several ecological types can be separated. This finding indicates the possibility of isolating different ecological types for specific localities in land reclamation areas.