Zur Fortpflanzung des Polychaeten Anaitides mucosa

In 1974, spawning polychaetes, Anaitides mucosa (Oersted, 1843), appeared in great numbers (up to 148 individuals/m²) at the surface of an intertidal flat in the German Waddensea from mid-February until 10th April. A. mucosa forms mucous bags which usually contain more than 10,000 green eggs. The eggs measure 113.4±5.4 μm in diameter, including the fructification membrane (18 measurements). Production if these egg bags was observed 5 times in nature and 4 times in the laboratory. Four to 17 males and 1 female form a ball of interlacing bodies. At the same time a mucous mass is secreted by the female — maybe also by the males — which increases to a diameter of as much as 2 cm within about 8 min. Eggs and sperm are shed into this mass of mucus. The worms appeared at the surface of the flat from early sunset until sunrise from about 3 h before to 1 h after low tide. Judging from observations of worms in the laboratory, it seems possible that the rhythm of sexual activity is controlled endogenously.     

Diatoms of the microphytobenthic community in a tropical intertidal sand flat influenced by monsoons: spatial and temporal variations

Seasonal variations in the microphytobenthic diatom community were investigated in an intertidal sand flat of a tropical marine environment influenced by monsoons. Cores of sediments were collected along the beach gradient: low tide, mid tide and high tide zone up to a depth of 15 cm.. Diatom abundance was lowest during the monsoons and highest during the post-monsoons and the early pre-monsoon season throughout the intertidal transect. Diatom diversity was highest at the mid tide, followed by the high and low tide zones. Diatoms were viable up to a depth of 15 cm throughout the intertidal transect. The diatom community included the pennates, the permanent residents of this area, centric genera, which lead an attached mode of life and also some planktonic genera, brought in from ambient waters. Among the pennates, Navicula and Amphora were the dominant genera whereas in the case of centrics, Thalassiosira dominated the community throughout the intertidal transect down to a 15 cm depth. . Grain size fractions, which served as predictors of some diatom genera changed with tidal zones. The effect of winds on the resuspension of the pennate diatoms was evident only at the low tide zone down to a depth of 5 cm . Chlorophyll a concentration proved to be a good predictor of both pennate and centric diatom abundance at the low tide zone down to a depth of 10 cm and at the mid tide zone down to a depth of 5 cm.. However, even though chlorophyll a concentrations failed to reveal any positive correlation with the diatom abundance at both the deeper sediment layers and the high tide zone, the fact that viable cells were present at these areas reveal that the diatoms adopt survival strategies, contributing significantly to the carbon budgets of such unstable habitats.     

Influence of seasonal food abundance and quality on the feeding habits of an opportunistic feeder,the intertidal crab <Emphasis Type="Italic">Pachygrapsus marmoratus</Emphasis>

The abundance of trophic sources on the intertidal zone is discontinuous and their supply can vary both in a predictable or unpredictable way. The Mediterranean semi-terrestrial crab Pachygrapsus marmoratus, is known, as adult, to entirely rely on the intertidal trophic sources, and, consequently, it faces the fluctuations of nutritional sources and quality. To clarify the relationships between the feeding habits of an Italian population of P. marmoratus and the temporal variation of its food sources, we carried out a 2-year sampling protocol. Data on seasonal variation in composition of intertidal food item assemblages, on the average content in N and C of the commonest algae, on seasonal changes in crabs feeding habits were collected and compared using a suite of multivariate and univariate techniques. Results showed that P. marmoratus takes advantage of the recruitment phase of the most common invertebrates, affecting and controlling the abundance even of those species whose adults are out of its reach. It can act both as a herbivore, and as a carnivore that shifts between bivalves, more abundant in spring/summer, and the periwinkle Melaraphe neritoides, during winter. In conclusion, we provide evidences on the food choice of this common rocky shore species, which is very plastic and capable of relying on many trophic sources, possibly influencing the abundance and/or the population structure of a number of intertidal populations.     

No zone is always greener: variation in the performance of<Emphasis Type="Italic"> Fucus gardneri</Emphasis> embryos,juveniles and adults across tidal zone and season

The responses of the different stages of an organisms life-cycle to different environments can have important implications for their persistence in those habitats. The marine intertidal zone is considered among the most stressful of all marine environments, because daily exposure at low tide results in daily fluctuations in a range of important environmental factors. In this study we examined the link between daytime exposure and the performance of embryos, juveniles and adults of the seaweed Fucus gardneri across zones and seasons on San Juan Island, Washington, USA. Specifically we asked: (1) whether three measures of performance for F. gardneri: growth rates, surface area of reproductive tissue and survivorship differed between high- and mid-intertidal zones, and among months throughout the year and, then, (2) whether total daytime exposure within each zone each month was a good predictor of variation in these measures of performance. At our study site, seasonal differences in the timing of low tides result in long periods each day during summer of very harsh conditions (exposure to the air, high temperatures and low humidity) compared to winter when exposure to the air during the day is virtually absent. Contrary to the expectation of lower performance in the high zone, we found no consistent differences between zones over the entire year in any measure of performance for any stage. For nearly all traits, there were significant differences among months and, importantly, significant interactions between zone and month. Thus, for each stage there were some months when thalli in the high zone performed better and some months when thalli in the mid-zone performed better. Regressions between total daytime exposure and performance each month indicated some stage-specific effects. Larger thalli appeared more strongly affected by the amount of exposure per month; there were significant negative relationships between total daytime exposure and (1) growth rates of medium and large adults in the high zone and (2) the surface area of reproductive tissue of medium adults in the high zone and small adults in the mid-zone. The lowest survivorship for all stages occurred in the 2–3 months that followed summer, suggesting a cumulative effect of long daytime exposure in summer on survivorship. Post-settlement survivorship of embryos was very low throughout the year, with only 2 out of 5395 seeded zygotes surviving to a visible size. One-month post-settlement survivorship ranged from 0% to 16.7%, and there were no differences in the survivorship curves between high- and mid-intertidal zones for each cohort. Although it is often assumed that the performance of intertidal species is lower in high-intertidal zones because of exposure to environmental extremes for longer, our results do not support this paradigm. Instead, they indicate a complex and fluctuating pattern of variation in the performance of different stages of the F. gardneri life-cycle throughout the year in different intertidal zones.Communicated by M.S. Johnson, Crawley     

Seasonal abundance,distribution and growth of postlarval and juvenile grass shrimp (Palaemonetes pugio) in a Georgia,USA, salt marsh

Postlarval and juvenile grass shrimp (Palaemonetes pugio Holthuis) ≦15 mm total length (TL) were abundant at low tide in shallow aquatic microhabitats (i.e. puddles and films of residual tidal water) in the intertidal zone of a salt marsh on Sapelo Island, Georgia, USA from 1982 to 1984. The highest concentrations of young P. pugio occurred at 190 to 200 cm above mean low water, ∼10 to 20 cm below mean high water. The intertidal distributions of young grass shrimp expanded and contracted with changes in tidal amplitude. Postlarval grass shrimp (6 to 8 mm TL) continuously recruited into the intertidal marsh population from May until October, but densities varied in a regular pattern with peaks in abundance occurring at ∼2-week intervals, corresponding to spring tide periods in the lunartidal cycle. Although present nearly year-round in the intertidal marsh, juveniles (9 to 15 mm TL) were most abundant from August to October. Apparent growth rates of individuals up to 15 mm TL averaged 0.268±0.026 (mean±95% C.I.) mm d^-1 from May to October and 0.070±0.032 mm d^-1 in November and December. Unlike larger aquatic organisms, which can forage in the emergent marsh only when it is flooded by the tide, juvenile grass shrimp have constant access to intertidal resources. Although potentially important predators in this system, the role of young P. pugio in the trophic organization of salt marsh benthic invertebrate assemblages has yet to be examined. Contribution No. 576 of the University of Georgia Marine Institute     

Life history of the sandy-beach amphipod Neohaustorius schmitzi (Crustacea: Haustoriidae)

Neohaustorius schmitzi Bousfield, the dominant macroscopic invertebrate on sandy beaches in North Carolina, USA, is strictly intertidal, and distributed from mean low tide to high tide, with an average density of 790/m². Location of greatest densities within the intertidal zone is dependent upon season and sex. An aggregated distribution is shown. N. schmitzi has 2 generations/year: a winter generation which lives about 8 months, and a summer generation which lives about 4 months. The reproductive season extends from February through October, with spring and summer peaks. Each female appears to produce only 1 brood of young. The number of young ranges from 2 to 14, depending upon the length and generation of the female.     

Distribution of Juvenile,subadult and adult Scylla serrata (Crustacea: Portunidae) on tidal flats in Australia

In Deception Bay, northern Australia, during 1979–1981, a study was made of the distribution of Scylla serrata (Forskal) in an area having a broad intertidal zone. Juveniles (20 to 99 mm carapace width) were resident in the mangrove zone, remaining there during low tide. The majority of subadult crabs (100 to 149 mm) migrated into the intertidal zone to feed at high tide and retreated to subtidal waters at low tide. Adults (150 mm and larger) were caught mainly subtidally and only small numbers were captured in the intertidal at high tide. Few crabs were captured in the coolest months (May to August). Adults were captured on the flats mainly in the warmest months (January to April), but subadults could be captured over the entire summer (September to March). Juveniles were found in the upper intertidal throughout the year.     

Estimate of removal rate ofNereis virens (Polychaeta: Nereidae) from an intertidal mudflat by gulls (Larus spp.)

The feeding behavior of herring gulls (Larus argentatus), ringed-billed gulls (L. delawarensis) and great blackbacked gulls (L. marinus) on an intertidal mudflat in Maine, USA, was investigated. Remains of fish, mussels, crabs, insects, and the polychaeteNereis virens were recovered from gull feces. Forty-three percent of the fecal samples containedN. virens jaws, setae, or both. A comparison of jaws from fecal material and from worms collected from the natural community demonstrates that gulls preferentially preyed upon larger worms. Feeding was largely confined to 3 h around low tide, and birds fed mostly in the low intertidal and below mean low water where the largest worms were located. Individual birds remained on the flat for a mean of 28.0 min and consumed a mean of 19.2 worms per visit. It was calculated that gulls remove a mean of 808 largeN. virens from the flat per tide from June to October, representing an estimated 0.04% of the standing crop of largeN. virens.     

Amphipods and isopods in the rocky intertidal: dispersal and movements during high tide

Animals in the intertidal, both mobile and sessile, generally exhibit some zonation pattern, in which each species shows a preference for, or is confined to, some height levels. The study of zonation patterns is, however, almost exclusively based on surveys made during low tide, when many animals are relatively inactive. We studied zonation patterns of amphipods and isopods on rocky shores in southwestern Iceland, both by traditional sampling at low tide as well as by sampling during high tide. The distributional patterns seen at high tide differed significantly from that at low tide. One amphipod, Anonyx sarsi, was common around baits at all levels at high tide but absent from the intertidal at low tide. Several other species were either relatively more common or tended to be recorded higher, or in one instance, lower on the shore when the tide was in than at low tide. There was also evidence of some species changing habitats within the intertidal with the tidal cycle. Many species, however, moved little away from their respective zones occupied at low tide, and for some species, including some capable of rapid swimming, very limited mobility was indicated. We conclude that low-tide surveys of the intertidal give an incomplete picture of the community structure, and even key species may be missed in such surveys.Communicated by L. Hagermann, Helsingør     

No evidence for homeoviscous adaptation in intertidal snails: analysis of membrane fluidity during thermal acclimation, thermal acclimatization, and across thermal microhabitats

Many eurythermal organisms alter composition of their membranes to counter perturbing effects of environmental temperature variation on membrane fluidity, a process known as homeoviscous adaptation. Marine intertidal gastropods experience uniquely large thermal excursions that challenge the functional integrity of their membranes on tidal and seasonal timescales. This study measured and compared membrane fluidity in marine intertidal snail species under three scenarios: (1) laboratory thermal acclimation, (2) thermal acclimatization during a hot midday low tide, and (3) thermal acclimatization across the vertical intertidal zone gradient in temperature. For each scenario, we used fluorescence polarization of the membrane probe DPH to measure membrane fluidity in individual samples of gill and mantle tissue. A four-week thermal acclimation of Tegula funebralis to 5, 15, and 25°C did not induce differences in membrane fluidity. Littorina keenae sampled from two thermal microhabitats at the beginning and end of a hot midday low tide exhibited no significant differences in membrane fluidity, either as a function of time of day or as a function of thermal microhabitat, despite changes in body temperature up to 24°C within 8 h. Membrane fluidities of a diverse group of snails collected from high, middle, and low vertical regions of the intertidal zone varied among species but did not correlate with thermal microhabitat. Our data suggest intertidal gastropod snails do not exhibit homeoviscous adaptation of gill and mantle membranes. We discuss possible alternatives for how these organisms counter thermal excursions characteristic of the marine intertidal zone.     

Measurement of chlorophyll fluorescence reveals mechanisms for habitat niche separation of the intertidal seagrasses <Emphasis Type="Italic">Thalassia hemprichii</Emphasis> and <Emphasis Type="Italic">Halodule uninervis</Emphasis>

In Taiwan, Thalassia hemprichii dominates the upper intertidal zone, whereas Halodule uninervis occupies the lower intertidal zone. We tested the hypothesis that T. hemprichii is better adapted to high irradiance and more resistant to air exposure than H. uninervis. The photosynthetic efficiency, damage, and extent of recovery were determined by measuring chlorophyll fluorescence using pulse amplitude modulated fluorometry. Both species growing in tidal pools, in response to high irradiance alone, revealed a small depression in maximal quantum yield of photosystem II (Fv/Fm) at noon. The second experiment compared the effect of air exposure alone and the combined effect of air exposure with high irradiance by interposing a shading screen on both species, growing in the intertidal zone over a diurnal cycle. Values of Fv/Fm of both the shaded and irradiated T. hemprichii remained high at low tide. However, H. uninervis exhibited a marked depression following air exposure and a synergistic depression under the combined effect. The experimental manipulations of exposure time demonstrated that the tolerance of T. hemprichii to the combined effect was longer and the recovery from air exposure following re-submersion was better than those of H. uninervis. Both species were more susceptible to the combined effect in the dry season than in the wet season. Our results suggest that air exposure is more important than high irradiance in constraining the distribution of H. uninervis in the upper intertidal zone. This was confirmed by transplantation experiments in which a rapid decline of H. uninervis was observed after transplantation into the upper intertidal zone. In the lower intertidal zone, measurements of the response of the photosynthetic electron transport rate to irradiance demonstrated that the transplanted T. hemprichii exhibited a sun-type response and H. uninervis a shade-type response.     

The contribution of seagrass beds (<Emphasis Type="Italic">Zostera noltii</Emphasis>) to the function of tidal flats as a juvenile habitat for dominant,mobile epibenthos in the Wadden Sea

Since the substantial loss of subtidal eelgrass (Zostera marina L.) in the 1930s, seagrass beds in the Wadden Sea are limited to the intertidal zone and dominated by Z. noltii Hornem. This study deals with the effect of vegetated tidal flats on quantities of mobile epifauna and proves empirically the function of seagrass canopies as a refuge for marine animals remaining in the intertidal zone at ebb tide. Drop-trap samples were taken in the Sylt-Rømø Bight, a shallow tidal basin in the northern Wadden Sea, on vegetated and unvegetated tidal flats during July and August 2002, and during the entire growth period of Z. noltii from May to September in 2003. The species composition in Z. noltii and bare sand flats showed minor differences since only two isopod species (Idotea baltica and I. chelipes) occurred on Z. noltii flats exclusively. Juvenile shore crabs (Carcinus maenas L.), brown shrimps (Crangon crangon L.) and common gobies (Pomatoschistus microps Krøyer) were also found abundantly on bare sand flats. However, the results showed significantly higher abundances and production of these dominant species on vegetated tidal flats. Additionally, the analyses of faunal size classes indicated higher percentages of small individuals in the seagrass bed during the entire sampling period. Despite drastic diurnal fluctuations of dissolved oxygen at low tide, faunal density in the residual water layer remaining in seagrass canopies at ebb tide was found to be consistently higher than that found in artificially created tide-pool units. Although species composition of mobile epifauna did not basically differ between vegetated and unvegetated tidal flats, Z. noltii beds are considered to contribute quantitatively to the function of tidal flats, as an extended juvenile habitat for some of the most important species of the Wadden Sea food web.Communicated by O. Kinne, Oldendorf/Luhe     

Migration of the sandworm Nereis virens during winter nights

There have been many previous reports of the sandworm Nereis virens Sars swimming in the water column. This behavior usually has been attributed to reproductive processes. Sandworms were found swimming in surface waters at night on ebb tides during many nights of January, February and March in a Maine (USA) estuary. None of the specimens examined contained gametes or possessed other characteristic spawning or pre-spawning modifications. Several age classes were found, with worms measuring 9 to 38 cm in length, weighing 0.5 to 19.8 g, and having 82 to 187 segments. The greatest numbers of worms were observed during near-average tides on evenings in which low tides occurred a few hours after sunset but prior to moonrise. Up to 83 worms per minute were observed swimming seaward through a 20 m transect, while none were observed swimming landward at any stage of the tide. It is concluded that sandworms swimming during winter nights is unrelated to reproduction and that it is an inherent behavior pattern.Contribution No. 102 of the Ira C. Darling Center, Walpole, Maine 04573, USA.     

Factors affecting the feeding rate of the winkle Littorina littorea

The winkle Littorina littorea (L.) feeds mainly when moistened by the tide and is quiescent when exposed to the drying effects of the wind and sun. Upper shore winkles thus experience a reduced feeding time compared with individuals living lower in the intertidal zone, but are able to completely compensate by an increased rate of radular activity when immersed. The rate of radular activity is influenced by the body size of winkles, by experimental temperature and by acclimation temperature, but is mainly modified by the duration of exposure to air which directly controls the rate of radular activity on immersion.     

The breeding cycles of two species of grapsid crabs (Crustacea: Decapoda) from the North Coast of Kyushu,Japan

The incidence of ovigerous females in populations of two grapsid crabs, Hemigrapsus penicillatus (de Haan, 1835) and Sesarma (Parasesarma) pictum (de Haan, 1835) were followed from August 1975 to November 1976. H. penicillatus, which inhabits the lower intertidal region near the mouth of Tatara-Umi Estuary, breeds from March to November. S. pictum, which inhibits crevices and abandoned holes of other species and is abundant at and above the upper intertidal region of the estuary, breeds from May to September. H. penicillatus matures when the female reaches 6 to 7 mm carapace width, whereas S. pictum becomes mature when the carapace width is 12 to 13 mm, although the maximum size attained by females of both species is almost the same. H. penicillatus produces 5 to 6 broods, S. pictum 2 to 3 broods, during a breeding season. The size of a newly laid egg of H. penicillatus is smaller than that of S. pictum. The number of eggs produced by a female H. penicillatus is greater than that of S. pictum during a breeding season. In both species the peak of the breeding season is during summer, at which time the ovarian activity is also apparently accelerated. The major environmental factor which controls the breeding in these crabs appears to be temperature. H. penicillatus is submerged at every high tide, and is relatively inactive from late November to February when the ambient water and air temperatures are rather low. S. pictum is dormant in crevices or understones of the splash zone from November to March. The length of the breeding season of these crabs appears to be inversely proportional to the period of their winter dormancy.     

Diel eclosion rhythm of a sublittoral population of the marine insect Pontomyia pacifica

Within the scope of studies on adaptation of insects to intertidal and sublittoral environments, correlations between the reproduction period of the short-lived chironomid Pontomyia pacifica Tokunaga and tidal conditions were examined at the only known Japanese location of this species in 1980. The larval habitat is situated, except for deeper tide pools in the lower midlittoral zone, mainly within the inner sublittoral zone, whose upper area is covered by Hypnea choroides, the dominating algae during the summer. The aerial adults emerge on the surface of the open sea and swarm there independent of the tidal situation above the submersed sublittoral habitat. Eclosion always started during dusk and all reproductive activities were ended within 2.5 h after sunset. The adaptation of P. pacifica (sublittoral habitat, diel eclosion after sunset with mass concentration of the adults on the water surface, no semilunar or lunar timing of the reproduction period and sinking egg masses) corresponds with those of the convergent marine chironomid Clunio balticus from Europe. On the basis of a few laboratory observations with P. pacifica and detailed experiments with C. balticus, it is supposed that the diel eclosion of P. pacifica is also controlled by an endogenous, circadian timing mechanism and the 24-h light-dark cycle as an environmental time cue.Communicated by O. Kinne, Oldendorf/Luhe     

Regulation of intertidal food webs by avian predators on New England rocky shores

Although there is a large body of research on food webs in rocky intertidal communities, most of the emphasis has been on the marine benthic components. Effects of avian predation on highly mobile predators such as crabs, remains practically unstudied in rocky shore ecosystems. The crab, Cancer borealis, is an important component of the diet of gulls (Larus marinus, L. argentatus) at the Isles of Shoals, Maine, USA. C. borealis prey include the predatory gastropod Nucella lapillus L., the herbivore Littorina littorea, and mussels Mytilus edulis L. We hypothesized that gulls reduce abundance of C. borealis in the low intertidal and shallow subtidal, thereby allowing C. borealis prey to persist in high numbers. A study of crab tidal migration showed that C. borealis density nearly doubled at high tide compared to low tide; thus, crabs from a large subtidal source population migrate into the intertidal zone during high tides and either emigrate or are removed by gulls during low tides. Results from a small-scale (1 m2) predator caging experiment in the low intertidal zone indicated that enclosed crabs significantly reduced L. littorea abundance when protected from gull predation. In a much larger-scale gull exclusion experiment, densities of C. borealis increased significantly during low and high tides in exclosures relative to the controls. C. borealis density was inversely correlated with changes in the abundance of two mesopredators Carcinus maenas and Nucella lapillus, and with the space-occupier M. edulis. There was a similar negative correlation between abundance of C. borealis and the change in abundance of the herbivore L. littorea, but the trend was not significant. Mortality of tethered L. littorea was associated with C. borealis density across sites. However, preferred algae did not change in response to L. littorea density during the experiment. Thus, we found suggestive, but not conclusive, evidence for a three-level cascade involving gulls, crabs, and L. littorea. Our studies strongly suggest that gulls, as apex predators, generate three-level trophic cascades in rocky intertidal food webs by preventing the highly mobile subtidal predator, C. borealis, from establishing substantial populations in the low-mid intertidal zone thereby indirectly enhancing densities of two key mesopredators (N. lapillus, Carcinus) and blue mussels (M. edulis).     

How a subsocial intertidal beetle,Bledius spectabilis,prevents flooding and anoxia in its burrow

Summary All stages of Bledius spectabilis Kratz (Staphylinidae) dig wine-bottle shaped burrows in the intertidal Salicornia zone. The adult female lays her eggs around her burrow and, by remaining with them, prevents both flooding by the tide and anoxia. Flooding is prevented by an ever-ready burrow which exploits a surface tension effect and by blocking the burrow once the tide comes in. The burrow has a narrow neck (about 2 mm in diameter) leading to a living chamber 5 mm in diameter. Tides over artificial tubes in agar showed that a critical minimum neck diameter of 2–3 mm prevented sudden flooding, giving time to block the neck with mud. Blocking took about four minutes. Ensuring the burrow is reopened at each low tide is a vital role of the brooding female in anaerobic or impermeable soils. Field models of burrows became anoxic in 4 days, much less than the 4 week long egg stage. A lab model system with anaerobic agar and calculations of oxygen uptake and diffusion supported this conclusion. Mortality of orphaned larvae may be much lower, however, in burrows within large Bledius aggregations because of mitigating good drainage and soil aeration: larval mortality from physical causes in these burrows did not increase over 3 weeks but without the mother, 14% of the eggs were attacked by mould, and two burrows were taken over by a predatory carabid, Dichierotrichus gustavi, and all larvae eaten. The surface tension effects of small air-filled openings may be used by small air-breathing animals in many intertidal habitats. Like Bledius, other intertidal animals, including large ones, may block their burrows at high tide to keep them full of air. Maternal care, in particular the combination of behaviours which protect the brood from the tide and anoxia, enables this airbreathing insect to colonise the inhospitable habitat of the intertidal saltmarsh.     

The “piece-by-piece predation” of<Emphasis Type="Italic"> Eteone longa</Emphasis> on<Emphasis Type="Italic"> Scolelepis squamata</Emphasis> (Polychaetes )—traces on the sediment documenting chase,defence and mutilation

On many sea shores of the Niedersachsen coast, the polychaete Scolelepis squamata is the dominant animal species living in the sediment of exposed beaches.The population of the predatory species Eteone longa, with a main distribution in more sheltered intertidal and subtidal habitats, has a certain overlap with S. squamata. In these restricted areas, S. squamata suffers from a permanent pursuit by E. longa during low tide. Field and laboratory studies have revealed that this predator-prey relationship follows a distinct behavioural pattern and is an exploitation of surviving animals: the predator does not ingest the whole prey individual but only feeds on regenerable parts of the body. Both species draw characteristic tracks on the sediment surface that illustrate the phases of the chase, the attempts at defence and the mutilation of the prey species.Communicated by O. Kinne, Oldendorf/Luhe     

Plasticity in the temporal organization of behaviour in the limpet <Emphasis Type="Italic">Cellana grata</Emphasis>

The behaviour of intertidal consumers is often tightly constrained to tidal movements, although activity patterns can vary within these constraints. Spatio-temporal variability in behaviour of a limpet, Cellana grata, was analysed over different tidal conditions (spring and neap tides) and during different times of the year (one summer and one winter) at sites in Hong Kong. Activity was generally dictated by tidal movements, being concentrated when animals were awash. Plasticity in behaviour was observed, with some limpets anticipating activity during the summer period and delaying activity during winter time. Limpets were active for a time equal, or slightly less, than time awash. As the time awash exceeded ~14–16 h, however, activity duration decreased. Within this general pattern, tidal variation as well as variation among times of the year was noted, with the lowest dependence on time awash being recorded during winter neap tides. Limpets showed a slight preference for being active during nighttime, which was particularly evident when animals were emersed during the summer period. Although the basic activity in C. grata is constrained to a specific temporal window, this limpet is able to modulate its foraging strategies and resting height, according to local, daily changes in environmental conditions.