Dynamic zonation of Nerita plicata,N. undata and N. textilis (Prosobranchia: Neritacea) populations on a rocky shore in Kenya

Vertical zonation of Nerita plicata Linnaeus, N. undata Linnaeus, and N. textilis Dillwyn, inhabiting the rocky cliffs of Nyali, Kenya, was studied through several spring-neap tidal cycles from November to December (northeastern monsoon) 1981 and May to July (southeastern monsoon) 1982. For the first time, a monsoon-related cycle of migration in Nerita plicata and N. undata, superimposed on the shorter term spring-neap cycle is demonstrated. Both species rest predominantly in the upper eulittoral zone and the littoral fringe. N. plicata and N. undata rest at significantly higher levels during the southeastern monsoon (SEM) than during the northeastern monsoon (NEM) (P<0.001). On and around neap-tide days N. plicata, N. undata, and N. textilis occupy significantly distinct mean resting positions (P<0.001) in both seasons; N. undata being the highest in the SEM, followed by N. plicata, with N. textilis always in the lowest level. In the NEM, N. plicata and N. undata sometimes change order. On and around springtide days in both seasons, the upper two species, N. plicata and N. undata, rest close together with no significant difference in their mean resting positions, but with both significantly higher than N. textilis (P<0.001). Both the spring-neap and the monsoon migrations of N. plicata and N. undata are interpreted as either direct or indirect responses to wave action.     

Notes on the Nerita (Archaeogastropoda) populations of Aldabra Atoll,Indian Ocean

Nerita undata, N. plicata, N. polita, N. albicilla and N. textilis are common intertidal gastropods on Aldabra Atoll. Each species prefers a different level on the shore and/or different degrees of exposure to wave action. Patterns of zonation, population size-frequency structure and biomass are given. N. undata exhibits 2 distinct ecophenotypes. Normal types occur on expsed to moderately sheltered shores but are replaced by type B in extremely sheltered conditions. Differences in shell morphology and population structure are also noted. Foraging by all species was confined to hours of darkness, generally at low tide. Activity of N. polita was induced slightly on overcast days. N. undata and N. textilis are cryptically coloured and often visible during the day. N. plicata is conspicuous, but is protected by a strong shell. N. polita and N. albicilla are often brightly and variably coloured, but both are concealed during daylight. No Nerita were found infected with trematodes, suggesting that the degree of infection was extremely low. Habitats of the Aldabran Nerita are compared with 3 species on Barbados, West Indies.     

Field observations on the rhythmic behaviour of Nerita textilis (Gastropoda: Prosobranchia)

Field observations on Nerita textilis Dillwyn inhabiting rocky cliffs at Sar Uanle (Southern Somalia) revealed that during the resting phase some of the snails are scattered on the upper cliff and some are clustered at lower levels. During feeding they all migrate downward, but to different extents. Maximum activity of both groups occurs during low water (LW) between midday and midnight. During spring tides (ST), the scattered snails reside higher up and make smaller feeding excursions than during neap tides (NT). During ST, some clustered snails join the belt of scattered individuals, while the trend reverses during NT. A diurnal and a tidal component appear in their periodicity.Publication of the Centro di Studio per la Faunistica ed Ecologia Tropicali del C.N.R. Researches on the coast of Somalia. The shore and the dune of Sar Uanle, No. 14.     

Leaf consumption by <Emphasis Type="Italic">Sesarma plicata</Emphasis> in a mangrove forest at Jiulongjiang Estuary,China

The feeding ecology of Sesarma plicata (Grapsidae: Sesarminae), the most abundant crab species in a mangrove forest dominated by Kandelia candel at Jiulongjiang Estuary, China, was investigated through field and laboratory experiments. Feeding preference and consumption rates were determined on mature, senescent and decomposed leaves of Kandelia candel, Bruguiera gymnorrhiza and Aegiceras corniculatum. In the laboratory, S. plicata preferred leaves of K. candel over those of B. gymnorrhiza and A. corniculatum, and consumed significantly more decomposed leaves than mature and senescent ones, irrespective of crab size. Field experiments with limited power failed to reveal detectable species preferences despite more consumption of K. candel, but decomposed leaves of each species were again preferred. Leaf characteristics associated with preference changed with plant species and leaf state. Low tannins and high water content characterized the preference for decomposed state of leaves. Species preference was significantly and negatively related to crude fibers and C:N ratios for mature leaves, and crude fiber for senescent leaves, but significantly and positively related to water content for decomposed leaves. Leaf consumption rates averaged for all leaf categories from laboratory no-choice feeding experiments were 0.101, 0.055 and 0.017 gDW ind⁻¹ d⁻¹ for large, medium and small crabs, respectively. In this forest, mean density of S. plicata was 20.5 ind m⁻² as assessed by a manual catching method. Leaf litter removal rate during neap tides by sesarmid crabs was about 1.33 gDW m⁻² d⁻¹ in April 2006. The leaves removed by crabs were grazed on the sediment surface or taken into crab burrows, shredded and stored before being eaten.     

Daytime surface swarming of Euphausia pacifica (Crustacea: Euphausiacea) in the Sanriku coastal waters of northeastern Japan

Biological characteristics of daytime surface swarming of Euphausia pacifica were investigated. Swarms in the Sanriku coastal waters consisted almost exclusively of mature individuals and seemed to be related to breeding behavior. Although maturation of ovarian eggs occurred both at the beginning and the end of the swarming period, females with attached spermatophore were most frequent at the end of the period. Since stomachs of swarming individuals were nearly empty throughout this period, swarming behavior was probably not related to feeding. Swarming individuals did not seem to be transported to the surface by vertical water movement caused by tidal streams, because the daily catch per boat did not differ between spring and neap tides.     

Spatial and temporal strategies in the surface activity of some sandy beach arthropods living along the French Atlantic Coast

During a field study on the eco-ethology of sandy beach arthropods conducted along the French Atlantic Coast it emerged thatTylos europaeus, Phaleria cadaverina andLabidura riparia were among the most abundant species that lived in the eulittoral. Tetradirectional pitfall traps, placed from the base of the dune to the tidal limits, were used. The traps intercepted surface-active arthropods, and the number of traps varied according to the levels of the tides. Data were recorded hourly during two periods corresponding to a spring tide and the following neap tide. At the saine time the most important environmental parameters were registered each hour. The results indicate that all three species show prevalently nocturnal surface activities. ForT. europaeus andPh. cadaverina an influence of the synodic and tidal phases was found on the zonation of the surface activity and on the extent of movement. The latter was due to the greater or smaller distances that the isopods and tenebrionids had to move from the diurnal burrowing sites to the foraging zones. ForL. riparia no variations in the total mean zonations were found between the two synodic phases, and only at neap tide were seaward, followed by landward movements found. Multiple regression analysis showed that the environmental parameters registered in this period were never limiting, as threshold values in this case probably were not reached.     

Diurnal and lunar patterns of larval recruitment of Brachyura into a mangrove estuary system in Ranong Province, Thailand

The present study followed the temporal recruitment pattern of brachyuran larvae in a mangrove tidal creek on the Andaman Sea coast of Ranong Province, Thailand, based on the assumption that the processes governing recruitment are important for the overall population dynamics of mangrove brachyuran crabs. Plankton net samples were taken on five occasions: on two new moon spring tides, one waxing moon neap tide, one full moon spring tide and one waning moon neap tide during October and November 1997. In addition collectors for larval crab megalopae were employed every 3 days through one dry season and one wet season (March–October 1998). Both the plankton net samples and collector samples revealed four major brachyuran groups in three families: Ocypodidae, Grapsidae and Portunidae. The grapsid group was further separated into two morphotypes which were identified as Metaplax and sesarmid species. Identified group mean numbers per cubic metre were ocypodids 3.0, sesarmids 0.8 and Metaplax 0.5, while portunid megalopae were very scarce (≪0.1 m⁻³). Further analysis of plankton net samples showed that when considering the parameters date, depth, current direction and the diel cycle, Metaplax and ocypodids distribute differently in the tidal and lunar cycle. Metaplax recruitment dominates on flood tides and on bottom layers, followed by middle and surface layers. Conversely, ocypodid abundance varied significantly with date only. Notably recruitment was not dependent on the diel cycle for either group. The collector samples of megalopae showed that recruitment of ocypodids, Metaplax and sesarmids occurred on full and new moon spring tides, while portunid megalopae preferred to settle on full moon spring tides. Since tidal currents were related to the lunar cycle megalopa groups are also cross-correlated with tidal amplitude, except for the portunid group. It is concluded that megalopae recruit in a similar manner to what has been found in other regions of the world, except that the abundance of ocypodids and Metaplax is not influenced by the diel cycle. Received: 14 February 2000 / Accepted: 24 November 2000     

Movement patterns of the limpet Cellana grata (Gould) observed over a continuous period through a changing tidal regime

Time-lapse videography of limpets mounted with light-emitting diodes was used to monitor the movements of a population of the non-homing Cellana grata on a vertical gully wall in Hong Kong. Animals were monitored for >7 days to examine spatial and temporal variation in their behaviour as the tides transited from a semi-diurnal to an almost diurnal pattern. Movement was synchronised with the tides, irrespective of the day–night cycles. Limpets rested low on the shore and were stimulated to move by the rising tide. Individuals moved up shore with the flooding tide, maintaining themselves in the awash zone, and then down shore on the ebbing tide, until they reached a resting height when the tide then fell beneath them, exhibiting ‘zonal shuttling’. A tight coupling of limpet position to tide height persisted through the changing tidal pattern, and almost all animals displayed the same organisation of activity over all tides. Initiation of activity and maximum height reached were probably controlled by the tides, but the cessation of activity may have been controlled by an internal clock. The pattern observed is consistent with the threat of attack from aquatic predators coupled with the need to minimise physical stresses while exposed. It is also consistent with the avoidance of grazing lower on the shore where interspecific competition may be more intense. Limpets showed two peaks of activity per tide, corresponding to when the rate of change of tide height was the greatest, except when tides became much reduced during the transit to a diurnal pattern. Movement was triphasic: an initial rapid phase, followed by a slower phase in the high shore and then a rapid phase before the limpets stopped moving. This structure is common in limpets and in this case is likely to be a consequence of animals maintaining themselves within the awash zone. Tide height appears to determine foraging activity, but with modifications in the behaviour in response to factors operating at more local temporal and spatial scales.     

Foraging and homing behaviour in the high-shore, crevice-dwelling limpet Helcion pectunculus (Prosobranchia: Patellidae)

The foraging activity of the high-shore, crevice-dwelling limpet Helcion pectunculus (Gmelin, 1791) (Prosobranchia: Patellidae) from an exposed shore on the eastern Cape coast of South Africa was monitored. Activity was compared during spring and neap tides and between spring, summer, autumn and winter. Rhythms of activity in this limpet varied depending upon microhabitat; individuals inhabiting both east- and west-facing rock surfaces were active during nocturnal low tides, but limpets on west-facing rock surfaces were also active during daytime low tides, whilst in the shade. Individuals travelled further during foraging excursions in winter (xˉ=85.53 cm) than in either spring (xˉ=55.7 cm) or summer (xˉ=48.8 cm) and also during spring low tides (xˉ=89.8 cm) compared with neaps (xˉ=40.9 cm). H. pectunculus exhibited rigid homing to a fixed scar within a crevice, and feeding excursions were found to consist of three distinct phases: a relatively rapid outward phase, a much slower foraging phase and a rapid homeward phase. Whether or not these limpets graze throughout an excursion is not known. Foraging was always highly directional, with a mean vector which took limpets onto an area of the rock face with the highest microalgal biomass. The fact that H. pectunculus took advantage of optimal feeding areas and memorized their location to enable return visits suggests a learning component in herbivorous gastropod foraging behaviour. Received: 29 December 1997 / Accepted: 8 May 1998     

Chlorophyll blooms and the spring/neap tidal cycle: observations at two stations on the coast of Connemara,Ireland

Water temperature, chlorophyll a and phaeophytin were recorded every 2 to 4 d, in order to describe short-term variability in phytoplankton blooms. Data were collected from May to September 1984 and March to September 1985 at shore stations on the coast of Connemara, western Ireland. Comparisons with offshore data suggest that the shore data is representative of coastal water. Chlorophyll a was highest at neap tides in spring, and at spring tides in early summer. In late summer, transient chlorophyll blooms occurred at neap tides, possibly due to a localized accumulation of flagellates in stable conditions.     

Diel vertical migration and feeding of copepods at an oceanic site near South Georgia

Twelve Longhurst Hardy Plankton Recorder (LHPR) profiles were taken over a 16 h period in January 1990, in order to study feeding of four copepod species at an Antarctic oceanic site near South Georgia. Vertical distributions of their life stages, as well as those of dominant competitors and predators, are described in relation to the feeding cycles of Calanoides acutus CV, Calanus simillimus CV, Calanus propinquus CV and Rhincalanus gigas CIII, CV and CVI. Comparisons with vertical ring-net catches, which were used for concomitant gutevacuation experiments, demonstrated the suitability of the LHPR for these fine-scale studies. Planktonic predators, with the exception of the diel migrant Themisto gaudichaudii, resided deeper than the herbivores. During the day and around midnight, when feeding rates were low, species and stages reached their maximum vertical separation. At these times, new generation copepodites of the four species lived progressively deeper and the overwintered generation (i.e., R. gigas Stages CIV, CV, CVI) were progressively shallower. During the afternoon or evening (depending on species), all stages older than CII, as well as Euphausia frigida and T. gaudichaudii, migrated upwards, to amass in the surface mixed layer. Feeding was restricted to darkness, although R. gigas commenced several hours before dusk. In detail their migration and feeding differed widely, with combinations of unimodal and apparent bimodal cycles. As a whole, the results suggest that (1) feeding could occur during sinking as well as during upward migrations, (2) upward migrations were not always associated with feeding increases, and (3) individuals appeared to descend after filling their guts.     

Filter-feeding ethology of benthic invertebrates (ascidians). IV. Pumping rate,filtration rate,filtration efficiency

A combination of a direct and an indirect method has been used for the first time in the study of filter-feeding in benthic invertebrates. Experiments over 12 h periods under constant temperature, pH, light and feeding conditions have been carried out with 3 species of ascidians: Ciona intestinalis (Linné, 1767); Phallusia mammillata (Cuvier, 1815) and Styela plicata (Lesueur, 1823). In C. intestinalis and P. mammillata, no pumping rhythm has been found; the water flow was constant for 12 h, except for some brief accidental interruptions. In S. plicata, the particular rhythm of spontaneous periodical contractions recorded during the experiment were related to gamete emission. The speed of water flow, which was fairly constant during recording, varied considerably during the 12 h period; these variations were generally related to modifications in the diameter of the cloacal siphon. The average speeds, in cm sec^-1, lay between 5.7 and 9.5 for C. intestinalis, 5.2 and 19.4 for P. mammillata, and between 5.3 and 10.7 for S. plicata. Variations in the pumping-rate were small, oscillating around a mean value. In ml h^-1 g^-1 dry weight of organs these were 5,829 to 5,982 (mean=5,906) for C. intestinalis, 6,142 to 6,592 (mean=6,312) for P. mammillata, 1,0508 to 1,1505 (mean=1,0708) for S. plicata. Filtration was continuous without any particular rhythm; filtration rates in ml h^-1 g^-1 dry weight of organs varied between 4,244 and 4,418 (mean=4,331) for C. intestinalis, 4,620 and 4,960 (mean=4,779) for P. mammillata, 8,482 and 9,078 (mean=8,760) for S. plicata. The curves representing pumping and filtration were clearly parallel, indicating that filtration efficiency did not vary greatly during the course of an experiment; the rates obtained were 65 to 87% (mean=74%) for C. intestinalis, 66 to 88% (mean 76%) for P. mammillata, and 73 to 90% (mean=80%) for S. plicata. The slightly higher mean value in S. plicata is probably related to the higher complexity of the branchial apparatus of this species.     

Annual reproductive cycle of the chiton Acanthopleura japonica

Annual reproductive cycle of the chiton Acanthopleura japonica (Lischke) was studied by observing both gonad index and histological features from October 1980 to October 1981. No significant changes in each factor were observed from October to April. Increased values of gonad index and spermatogenesis or vitellogenesis in gondas were observed from May to September. In the same time, gonad index decreased and sperms or oocytes in gonalds disappeared in the days around spring tide, and they were recovered around neap tide. Suggested spawning around spring tide from May to September is also supported by the occurrence of released eggs in the days around the new and full moons.     

Rhythms in larval release of the sublittoral crab Neopanope sayi and the supralittoral crab Sesarma cinereum (Decapoda: Brachyura)

Ovigerous females of the subtidal xanthid crab Neopanope sayi (Smith) and the high intertidal grapsid crab Sesarma cinereum (Bosc) were collected during the summers of 1986 and 1987 in the Beaufort, North Carolina (USA), area and brought into the laboratory, where rhythms in larval release were monitored. When crabs with late-stage embryos were put under a 14 h light:10 h dark cycle in an otherwise constant-environment room, an apparent tidal rhythm in release of larvae was observed for both species, with N. sayi releasing near the time of day and night high tides, and S. cinereum releasing around the time of night high tides. The time of sunset relative to high tide was a complicating factor, since larval release for both species was often concentrated around sunset when evening high tides fell several hours before sunset. When a group of N. sayi and S. cinereum were brought into the laboratory and placed under constant lowlevel light for 5 d, the release rhythm of the population persisted, thus implying that the rhythm is endogenous. Larval release near the time of high tide and often at night is common among brachyurans living in tidal areas, regardless of specific adult habitat, suggesting a common functional advantage. Possibilities include transport of larvae from areas where predation and the likelihood of stranding and exposure to low-salinity waters are high, as well as a reduced probability of predation on adult females. Results of the present study suggest that the importance of release after darkness may increase with increasing tidal height of the adult.     

Rhythmicity of larval release in three species of intertidal brachyuran crabs (Crustacea: Brachyura) from Inhaca Island (Mozambique)

 A study of rhythmicity of larval release in three species of intertidal brachyuran crabs, based on laboratory and field experiments, was undertaken at Inhaca Island, southern Mozambique, using Leptodius exaratus and Macrophthalmus grandidieri from December 1994 to January 1995, and Arcotheres palaensis from April to July 1995. L. exaratus and M. grandidieri showed a semi-lunar cycle in larval release. The release of larvae for L. exaratus, a species having conspicuous larvae, occurred in the first half of the night, after the post-crepuscular high tide, which suggests maximisation of protection of larvae from visual predation. The larval release activity matched the late spring and early neap tides. Results from the field were similar to those from the laboratory. M. grandidieri, having inconspicuous larvae, did not show a pattern related to the light–dark cycle and hatched during spring tides (around full and new moons) to maximise larval dispersion. A. palaensis, living inside the host mussel which inhabits the lowest section of the intertidal zone, did not show a relation with moon phase, tidal or light–dark cycles. Received: 16 February 1999 / Accepted: 8 December 1999     

Flood tide transport of blue crab, Callinectes sapidus, postlarvae: behavioral responses to salinity and turbulence

Blue crab (Callinectes sapidus) postlarvae (megalopae) use flood tide transport to move upstream in estuaries during nocturnal flood tides. The megalopae have a endogenous diel rhythm in activity that is inconsistent with this tidally timed behavior. Thus, it is hypothesized that this behavior is regulated by behavioral changes in response to exogenous cues associated with tidal currents. In a laboratory flow tank, blue crab megalopae were exposed to simultaneous changes in salinity and turbulence to simulate tides in an estuary. On simulated flood tides, megalopae ascended upon exposure to a salinity increase, remained swimming during times of high turbulence, and descended at times of low turbulence. Turbulence stimulated swimming for several hours, approximating the duration of tidal currents in estuaries. Swimming was inhibited by decreasing salinity on simulated ebb tides. These results support a model for regulation of flood tide transport by blue crab megalopae as follows: (1) blue crab megalopae are stimulated to swim into the water column by increasing salinity associated with flood tide; (2) megalopae remain swimming during flood tide in response to high levels of turbulence; (3) megalopae descend at the end of flood tide, when current speed and turbulence decline to low levels; and (4) megalopae are inhibited from swimming on ebb tides by the associated salinity decrease. This is the first model for regulation of flood tide transport in a species lacking a tidal rhythm in activity.     

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.     

Mélange vertical et capacité photosynthétique du phytoplancton estuarien (estuaire du Saint-Laurent)

Time series of chlorophyll a, photosynthetic capacity and many physical parameters were sampled hourly for 167 h in August, 1975, at an anchor station located in the Middle Estuary of the St. Lawrence River, Canada. Sampling was carried out during the transition from neap tides to spring tides. The long-and short-term variations in chlorophyll a are coupled with the advection of water masses which depends on tidal currents. Vertical mixing also influences the chlorophyll a concentration of the cells, since it modifies the physiological state of the phytoplankton. Furthermore, circadian periodicities were observed in the photosynthetic capacity, suggesting that the phytoplankton of this area have a homogeneous light history due to strong vertical mixing. Under these conditions, the photosynthetic capacity is adapted to the mean light intensity in the mixed layer. The semimonthly (M _f) variations of the mean light intensity in the mixed layer depend on the M _f variations in the vertical mixing, whereas in the short-term, the variations in mean light intensity in the mixed layer are circadian.

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Mélange vertical et capacité photosynthétique du phytoplancton estuarien (estuaire du Saint-Laurent)

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Contribution au programme du Groupe interuniversitaire de recherches océanographiques du Québec (GIROQ)     

Abundance of estuarine crab larvae is associated with tidal hydrologic variables

Abundances of brachyuran megalopae and juveniles were measured throughout consecutive tidal cycles during six 2 to 3 d sampling periods in summer 1992, and associated with rates of change of tidal hydrologic variables in the Newport River Estuary. Current speeds and rates of pressure change fitted sinusoidal (tidal) models well; however, rates of salinity and temperature change did not. Analysis of plankton samples taken during spring and neap tides showed peak abundances during nighttime rising tides for all taxonomic groups: Callinectes sapidus, Uca spp., Xanthidae, and Pinnixa spp. megalopae, and Pinnotheres spp. juveniles. Megalopal and juvenile abundances from time-intensive sampling were related to rates of changes in the hydrologic variables using stepwise logistic regression. No hydrologic variable accounted well for the presence of Uca spp. megalopae. Megalopal presence was best predicted by current speed for Pinnixa spp. megalopae, and rates of changes in pressure for xanthid megalopae and Pinnotheres spp. juveniles, and salinity for C. sapidus megalopae. These variables might act as cues causing megalopae to ascend into the water column at a particular point in the flooding tide, and subsequently descend to or near the bottom prior to ebb flow. In this way, larvae which develop on the continental shelf or lower estuary undergo transport up the estuary by behaviorally altering their swimming activity and depth concurrent with tidal changes.     

Monitoring spatio-temporal variation in molluscan grazing pressure in seasonal, tropical rock pools

In Hong Kong, low-shore rock pools support a patchy mosaic of different successional stages of crustose coralline algae. Echinoid and molluscan grazers are abundant in these low-shore rock pools and spatial and temporal variation in grazing pressure is thought to play an important role in structuring these assemblages. Direct records of graze marks using a modified wax-disc method were used to monitor spatial (different pools) and temporal (periods, 3 or 4 days each; spring vs. neap tides; summer vs. winter) variations in grazing pressure. Radula marks of Chlorostoma argyrostoma, Lunella coronata and Nerita albicilla were common in all pools. Grazing pressure was highly variable with tidal state and season but also within these two factors. Lunella and Nerita exerted highest grazing pressure at neap tides during summer, whilst Chlorostoma exerted highest grazing pressure in winter. Although overall patterns of temporal variation were similar and consistent in all pools, there was between pool variation in grazing pressure and individual species showed different patterns. Within the seasonal and tidal variations, large period-to-period differences were found for all species, illustrating small-scale temporal variation. Fine-scale spatial and temporal patterns of variation in grazing pressure of different species, therefore, contribute to the highly dynamic nature of these low-shore rock pools.