Nocturnal feeding migrations of Nerita plicata,N. undata and N. textilis (Prosobranchia: Neritacea) on the rocky shores at Mkomani,Mombasa, Kenya

The vertical zonation of the three common rocky shore neritids at Mkomani, Mombasa, Kenya, Nerita plicata Linnaeus, N. undata Linnaeus, and N. textilis Dillwyn, as a function of feeding migrations and of size, was studied from 28 February to 24 March 1983. These snails perform feeding migrations at night starting at around mid-ebb tide and return to their resting positions with the flood tide. They remain in their resting positions throughout the day until the next nocturnal ebb tide. The direction of migration is sizerelated, with the larger snails of each species moving in the opposite vertical direction to the smaller ones, so that the populations as a whole exhibit no statistically significant net vertical displacement. The larger individuals of two of the species, N. plicata and N. undata, invariably move downwards to their feeding levels, while the smaller individuals move upwards; the larger individuals of N. textilis display a different pattern of migration, moving downwards on and around spring-tide days and upwards on and around neap tide days, while the smaller individuals move in the opposite directions. N. textilis rest above their feeding level around spring tides, and below that level around neap tides. It is demonstrated how these nocturnal migratory feeding rhythms are integrated into the spring-neap and seasonal cycles of the snails' daytime resting positions. The adaptive significance of these migrations is also discussed.     

Trail following in the chiton Acanthopleura gemmata: operational and ecological problems

The Indo-Pacific, intertidal chiton Acanthopleura gemmata (Blainville) is a central place forager which occupies definite, actively dug and defended scars, and migrates during nocturnal low tides up to 3 m away to feed on algal grounds. After each feeding excursion, most chitons home precisely to their scars. Photographic tracking, using lightemitting diodes (LED) glued to the chitons, and experiments were conducted throughout 45 d (August to September 1984) along the Nimu Peninsula (Somalia) in order to elucidate the importance of trail following in the homing and feeding of this species. LED tracking of the chitons throughout each activity phase and trail interruption experiments showed that following the personal outgoing trail is a basic homing mechanism in this species. Translocation of the chitons on a conspecific trail and homing performance analysis revealed that the trail-associated information involved in the homing is polymorphic in the population, thus minimizing the probability of following a conspecific trail despite the frequent trail crossings due to high population density. The LED tracking throughout successive activity phases showed a high coincidence between the paths traced by the same chiton night after night. In the few cases when the outgoing and return paths of the first night markedly diverge, the home-feeding ground route of the successive night overlaps the return trail of the previous night. A less pronounced inter-individual trail following also occurs during foodward excursions. These findings suggest that the trails released by A. gemmata are complex, including both quasi-individual and species-specific information. Moreover, this chiton may utilize its trail following capacity not only to home, but also for the retrieval of the feeding grounds night after night.     

Function of the shell spine in the predaceous rocky intertidal snail Acanthina spirata (Prosobranchia: Muricacea)

Feeding observations revealed that Acanthina spirata (Blainville, 1832) uses the shell spine in a ramming and prying motion, resulting in the fracturing and/or forcing open of opercular plates in the barnacles Balanus glandula (Darwin, 1854) and Chthamalus fissus (Darwin, 1854). Attacking barnacles by using the shell spine may represent a second widespread feeding mechanism (in addition to drilling) in muricacean gastropods. Specialization of the shell spine for attacking barnacles is advantageous over drilling within the context of an optimal foraging theory. Rockey intertidal field and laboratory experiments at Palos Verdes and Los Angeles, California, respectively, in 1981–1982 revealed, that snails with a shell spine have lower handling times and feed at a greater rate than spineless snails which drill their prey. Spine penetration of prey as opposed to drilling takes considerably less time than a tidal cycle. This is expected to increase the success rate for completion of feeding, since during tidal flux snails risk being dislodged. Plasticity in attack mechanisms allows A. spirata with broken spines to penetrate prey by drilling while undergoing spine repair. Differences exist in spine-feeding by A. spirata on B. glandula and C. fissus. The attack process takes more than twice as long when snails attack B. glandula versus C. fissus. However, there is no difference in the mean number of spine thrusts required to penetrate the opercular plates of the two species. When feeding on the larger barnacle B. glandula, A. spirata ingests significantly greater dry weight per unit handling time than with C. fissus.     

Effects of Polinices duplicatus (Gastropoda: Naticidae) on infaunal community structure at Barnstable Harbor,Massachusetts, USA

Effects of the predatory naticid snail Polinices duplicatus (Say) on species composition, diversity and density, of the infauna of intertidal sand-flats at Barnstable Harbor, Massachusetts, USA, were studied using field experiments. Responses of molluscs (prey of P. duplicatus) and nonmolluscs (nonprey) were considered separately to distinguish between the effects of feeding and sediment disturbance during foraging. The fauna of 0.25 m² predator-exclusion cages, coarse-mesh cages, sieved areas, and controls was followed for 1 yr. Species associations within cages from which predators were excluded were denser, more diverse, and richer in molluscs than those in other treatments. Larger areas (3×3 m) of natural bottom were fenced and maintained as snail exclosures and enclosures during two feeding seasons. For both molluscs and nonmolluscs, diversity (H'), number of species, evenness (SD), and density all decreased with increasing snail density. Intense predation pressure on molluscs and selective feeding on thin-shelled bivalves, which were rare, removed individuals and species of molluscs from the community. Comparison of samples taken inside and outside trails made by snails showed that disturbance of the surface sediment layers by snails decreased the abundance of spionid polychaetes and total nonmolluscs. The sipunculan Phascolopsis gouldi and the bivalve Gemma gemma dominated the community in all experimental treatments and were little affected by the activity of Polinices duplicatus. Predation and disturbance by snails lowered community diversity by removing individuals of the less abundant species, and generally maintained population densities below the level where strong competition would occur.     

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     

Trail following,speed and fractal dimension of movement in a marine prosobranch,Littorina littorea,during a mating and a non-mating season

We quantitatively studied movement behaviour in the gastropod Littorina littorea in laboratory experiments during periods of their non-mating season (November 1992) and their mating season (April–May 1993). Snails were collected in 1992 and 1993 from one boulder shore on the north west coast of Sweden. In a comparison between the two seasons (one non-mating and one mating) we measured trail complexity of unsexed snails using fractal dimesion, the degree of mucus trail following (coincidence index of marker and tracker trails) and average movement speed of marker and tracker snails. We found no differences in fractal dimension and coincidence index of trails between the two seasons. Tracker snails moved, however, significantly faster than marker snails during both seasons. This could not be explained by trackers, when locomoting, using the mucus trail deposited by the marker to increase their speed, since there was no correlation between coincidence index and tracker speed. During the mating season we also conducted trail complexity, trail following and speed experiments comparing the behaviour of males and females. There was no difference between males and females in the fractal dimension of movement, nor was there any difference between the mean speed of male and female snails, although male marker snails tended to move faster than female marker snails. Males tracking other males, females tracking other females and females tracking males followed trails about equally long distances (i.e., coincidence indices did not differ). In contrast, males following female mucus trails showed a significantly higher degree of trail following than the other sex combinations. This new finding may suggest that females of L. littorea release pheromones in their mucus trails and that males are able to identify them.     

Competition and interaction between Neosarmatium smithi (Crustacea: Grapsidae) and Terebralia palustris (Mollusca: Gastropoda) in a Kenyan mangrove

 In a Kenyan mangrove, we studied the interactions between the gastropod Terebralia palustris and the crab Neosarmatium smithi when foraging on decaying mangrove leaves. Interactions are considerable on account of their wide overlap in zonation (Rhizophora mucronata belt), food items (mangrove leaves) and activity window (diurnal low water). The snails find a leaf by a systematic transecting of the platform and eat the leaf after crawling on to it, usually en masse. The crabs rarely venture beyond 80 cm from their burrows, and once they find a leaf they quickly drag it back into their burrow, probably to reduce the strong intra-specific competition. A crab is able to drag a leaf away even if a number of snails are already feeding on it, by strongly pulling the leaf away or by pushing the snails off the leaf. The success of the “thief ” depends on both the crab's size and the number of snails on the leaf. Received: 25 July 1999 / Accepted: 27 April 2000     

Factors influencing clustering in the intertidal snail Cerithium moniliferum

The snail Cerithium moniliferum Kiener at Heron Island (southern end of the Great Barrier Reef) clusters at outgoing tide and disperses at incoming tide. In containers with a constant water level, but exposed to the normal day and night rhythm, the snails cluster and disperse rhythmically for up to several days. Graphical simulation and spectral analysis, preceded by first difference filtering, showed two components, one corresponding to the tidal, and the other to the diurnal rhythm. Snails kept under continuous light also exhibit a two-component rhythm of clustering and dispersal. In containers with high (40 cm) water level and under continuous light or normal day and night rhythm, the two-component rhythm may—for some unknown reason—be replaced by long fluctuations of dispersal, with clustering occurring in the afternoon only in snails kept under normal alternation of day and night. Snails kept for 24 h in dryness prior to putting them into containers with water, remain clustered (sometimes with an initial dispersal “shock”) for a long time, with subsequent slow fluctuations of clustering and dispersal, probably induced by the light conditions. Factors contributing to clustering and dispersal are: (1) increased rate of dropping from container walls at outgoing tide; (2) lack of locomotory activity at low tide; (3) negative geotaxis at incoming tide; (4) positive geotaxis at outgoing tide; (5) attraction of snails by other snails (positive chemotaxis) throughout the rhythm. Sudden exposure to light increases the rate of dropping from the container walls and may be a contributing factor to stronger clustering during the day. Factors 1–4 show endogenous periodicities.     

Rhythmic vertical migration of the gastropod Cerithidea decollata in a Kenyan mangrove forest

In Mida Creek, Kenya (3°20′S, 40°5′E), at high water, the snail Cerithidea decollata dwells on the trunks of mangrove trees (Avicennia marina), while during low water it migrates to the ground, foraging at various distances from the trunk, where it aggregates again well before the incoming tide. Snails from the upper shore level are 150–200 m distant from those living at the lower shore level and they cluster at lower heights on trunks. In any case, sufficient height is usually attained to avoid being submersed. An experiment was designed (February and October 2005), exchanging individuals from different shore levels subject to different tide regimes, in order to test whether snails rely on internal information or on external, direct cues, to adapt their behaviour to local conditions. Results show that C. decollata mostly rely on internal information, presumably based on an internal clock. When individuals from upper and lower shore levels were exchanged, their internal clocks continued to govern when to ascend the home trunk and how high to climb for five to six successive tides, after which the behaviour was reset to the new local conditions.     

Distribution,movement and diet of the snailSearlesia dira in the intertidal community of San Juan Island,Puget Sound,Washington

Searlesia dira Reeve is a locally abundant, carnivorous gastropod that occurs from Alaska to central California. I studied populations of this snail at a variety of sites on San Juan Island in order to provide information on their distribution and abundance and to analyze their community role, i.e., their potential impact on their prey populations and their dietary overlap with co-occurring invertebrate predators. I established permanent transects, both random and in crevices, at three areas of differing exposure to wave action. On these transects I recorded the number and size ofS. dira, activity, distance to nearest neighbor, other organisms present, and tidal height on four phases of the mixed semi-diurnal tidal cycle. The results are (1) an increase in density with increase in substrate relief, (2) an occurrence of highest densities in moderately exposed and exposed sites rather than in protected ones, (3) an increase in mean size with increase in depth along the intertidal gradient, (4) an increase in both total activity and feeding success on the high water immediately following an extreme minus low tide occurring during the daytime in the summer, (5) a relative restriction of feeding success to that high tide, and (6) a similarity of the diet to the relative abundance of the main observed prey for most areas. The main exceptions are chitons and one lower intertidal limpet, all of which are taken in excess of their apparent abundance and especially in the middle intertidal. The upward extension of some lower intertidal prey populations, such as chitons, may be limited by the increased susceptibility of individuals at the upper margin of the population to predators such asS. dira; susceptibility appears to be increased after periods with a higher probability of increased physiological stress, such as extreme low tides on summer days.     

Habitats and tidal accessibility of the marine foods of dabbling ducks and brant in Boundary Bay,British Columbia

The role of birds in intertidal food webs is mediated by effects of tides on the habitat, distribution, and accessibility of food organisms. We studied effects of elevation, sediment characteristics, vegetation, and tidal cycles on waterfowl feeding and food organisms in Boundary Bay, British Columbia, Canada from September to March 1990–1992. Percent dry mass of esophageal contents was mainly Zostera japonica leaves (84%) for American wigeon (Anas americana Gmelin), and leaves of Z. japonica (57%) and Z. marina (41%) for brant (Branta bernicla L.). Northern pintails (A. acuta L.) and mallards (A. platyrhynchos L.) ate mostly the leaves, seeds, and rhizomes of Z. japonica (48 and 72%, respectively) and gastropods (12 and 28%). Green-winged teal (A. crecca L.) ate mainly amphipods (68%) and saltmarsh seeds (19%). The well-sorted fine sand (median 0.125 mm) to medium sand (0.25 mm), and the low organic carbon content (1.31±0.12% dry mass), did not appear to vary sufficiently along 2.1- and 2.6-km transects to affect macroinvertebrate distributions. Three habitat zones were recognized: (1) upper intertidal (sandflats); (2) mid-intertidal (largely monotypic Z. japonica); and (3) lower intertidal (mixed Z. japonica and Z. marina). These zones, although related to elevation, had greater influence on amphipod distributions than did elevation alone or detrital biomass. Biomass and numbers of most invertebrate foods of waterfowl were greatest in the eelgrass zones, with the upper elevational limit of Z. japonica occurring at about Mean Water Level (MWL). Accordingly, wigeon, pintails, and mallards fed mostly below MWL. Because of seasonal tidal cycles, daily accessibility of areas below MWL but within reach of surfacefeeding waterfowl decreased from a mean of 10.8 h in October to 9.6 h in December. In October only 10 d and <12 h available feeding time, whereas in December and January every day had <12 h available feeding time. Seasonal effects of tides on accessibility might limit sufficient feeding by some waterfowl species in intertidal areas, increasing their reliance on alternative feeding sites.     

Effect of epibiosis on the fitness of the sandy shore snail<Emphasis Type="Italic"> Batillaria zonalis</Emphasis> in Hong Kong

Batillaria zonalis is a common snail on Hong Kong sandy shores, occupying the low-mid shore. Shells of B. zonalis (basibiont) are often fouled with the rock oyster Saccostrea cucullata and the barnacle Balanus reticulatus (epibiont). This epibiotic interaction has a negative impact on the fitness of B. zonalis in terms of impairing fecundity, mobility and metabolism of fouled individuals. During the reproductive season, fouled snails had lower Gonadosomatic Index and fewer egg-bearing individuals than non-fouled snails. Fouled snails crawled in a more tortuous pathway and with lower speed when compared with non-fouled snails. Reduction in oxygen consumption and metabolic rate in fouled snails further suggested the restriction in mobility of B. zonalis by the epibionts. Glycogen content in the foot muscle tissues and hepatopancreas of fouled and non-fouled snails was, however, similar, revealing epibiosis does not influence the energy storage of B. zonalis. Mortality of fouled and non-fouled snails was similar under laboratory conditions, suggesting epibiosis does not influence the survivorship of B. zonalis in Hong Kong. Impairment in the fitness of fouled B. zonalis indicates that epibiosis can influence the population dynamics of B. zonalis in Hong Kong.Communicated by T. Ikeda, Hakodate     

Effects of an invasive ant on land snails in the Ogasawara Islands

We investigated how Pheidole megacephala has affected endemic achatinellid snails because these snails are excellent indicators of the impact of ants and they have high conservation value in Ogasawara. In 2015 we surveyed the Minamizaki area of Hahajima Island of Ogasawara, designated a core zone of the World Heritage Site, for P. megacephala. In Minamizaki, we determined the distribution and density of achatinellid snails in 2015 and compared these data with their distribution and density in 2005. Land cover in the survey area was entirely forest. We also tested whether P. megacephala preyed on achatinellid snails in the laboratory. P. megacephala was present in the forested areas of Minamizaki. Achatinellid snails were absent in 19 of 39 sites where P. megacephala was present, whereas in other areas densities of the snails ranged from 2 to 228 individuals/site. In the laboratory, P. megacephala carried 6 of 7 achatinellid snails and a broken shell was found. Snail distribution and density comparisons and results of the feeding experiments suggest that the presence of P. megacephala has contributed to the decline of achatinellid snails in forests in the survey area. Yet, P. megacephala is not on the official list of invasive non‐native species. Stakeholders using the list of invasive species to develop conservation programs should recognize that invasiveness of non‐native species differs depending on the ecosystem and that official lists may not be complete.     

Variability in abundances of fishes associated with seagrass habitats in relation to diets of predatory fishes

The spatial, diel and tidal variability in the abundance of piscivorous fishes and their teleost prey, and the dietary composition of predatory fishes were investigated in beds of Heterozostera tasmanica within Port Phillip Bay, Australia, from September 1997 to February 1998. Predatory and prey fish assemblages were sampled from beds of H. tasmanica at three locations during each combination of diel (day and night) and tidal (high and low) cycles. Pelagic and benthic crustaceans represented >60% by abundance of the diets of all predatory fishes. Seven species, 54% of all predatory fishes, were piscivorous. These piscivores consumed individuals from seven families, 36.8% of the fish families being associated with seagrass. Western Australian salmon, Arripis truttacea (Arripidae) (n = 174) and yank flathead, Platycephalus speculator (Platycephalidae) (n = 46) were the most abundant piscivores. A. truttacea consumed larval/post-larval atherinids, gobiids and sillaginids. P. speculator consumed late-juvenile/adult atherinids, clinids and gobiids. While the abundances of piscivores varied between locations (P < 0.001) and diel periods (P = 0.028), the relative differences in piscivore abundance between sites and diel periods were not consistent between tides. The abundances of A. truttacea varied in a complex way amongst sites, diel period and tidal cycle, as shown by a three-way interaction between these factors (P = 0.026). Only during diurnal periods at St. Leonards was the abundance of A. truttacea significantly higher during high than low tides (P < 0.001). During the other diel periods at each site, the abundance of A. truttacea did not vary. P. speculator was significantly more abundant nocturnally (P = 0.017). The abundance of small (prey) fishes varied significantly amongst sites (P < 0.001). During the day, the abundance of small fishes did not vary between high and low tides (P = 0.185), but their nocturnal abundance was greater during low tide (P < 0.001). Atherinids (n = 1732) and sillaginids (n = 1623) were the most abundant families of small fishes. Atherinids were significantly more abundant nocturnally (P = 0.005) and during low tides (P = 0.029), and varied significantly amongst sites (P < 0.001). Sillaginids varied significantly only amongst sites (P < 0.001). Seagrass beds provide a foraging habitat for a diverse assemblage of predatory fishes, many of which are piscivorous. Anti-predator behaviour and amongst-location variability in abundances of piscivorous fishes may explain some of the diel and tidal, and broad-scale spatial patterns in small-fish abundances. Received: 23 July 1999 / Accepted: 18 January 2000     

Grazing rates of organic matter and living fungal biomass of decaying Spartina alterniflora by three species of salt-marsh invertebrates

 The pathway for the flow of salt-marsh grass production into marsh food-webs is still not well defined. We compared the abilities of three marsh macroinvertebrates [salt marsh periwinkles, Littoraria irrorata (Say) (=Littorina irrorata), salt-marsh coffee-bean snails, Melampus bidentatus (Say); and a talitrid amphipod, Uhlorchestia spartinophila Bounsfield and Heard] to access standing-dead leaves of smooth cordgrass (Spartina alterniflora Loisel). The invertebrates were incubated with naturally-decaying leaves, and the rates of removal of organic matter and living fungal biomass (ergosterol) were measured. The impact of invertebrate activity upon fungal growth rates was measured as rates of fungal-membrane synthesis (incorporation of radioacetate into ergosterol). The removal rates of organic leaf biomass per mg individual biomass were highest for amphipods (700 μg mg⁻¹ d⁻¹) and lowest for periwinkles (90 μg mg⁻¹ d⁻¹), but the relatively large biomass of the snails made their removal rates per individual greater than those of amphipods. Net removal of ergosterol by all three invertebrates was >50% for yellow-brown (early-decay) leaf blades. For fully-brown (advanced-decay) blades, >50% removal of ergosterol was found only for periwinkles; exposure to coffee-bean snails and amphipods resulted in a net ergosterol reduction of ≤20%. The lower net reduction of living fungal biomass by coffee-bean snails and amphipods may have been due to fungal-growth stimulation (2.3-fold stimulation in coffee-bean snails and 1.5-fold stimulation in amphipods). Grazing by periwinkles did not stimulate fungal growth, possibly because of its high intensity. Grazing by these three salt-marsh shredders may affect marsh-grass shoot-decay in different ways. Periwinkles may abbreviate the period of fungal production, and incorporate the decaying material relatively quickly into snail biomass and fecal-pellet rain to the sediments. Coffee-bean snails and amphipods may enhance and prolong fungal production, along with the formation of fecal-pellet rain. All three invertebrates fed preferentially on leaf blades rather than leaf sheaths, and feeding rates of gastropods were higher during the night than during the day. Received: 25 November 1998 / Accepted: 4 November 1999     

Evidence of in situ diel vertical migration of a red-tide microplankton species in Ría de Vigo (NW Spain)

The migration capacity of red-tide species in the natural environment was studied at a station in the Ría de Vigo (Rías Bajas, NW Spain) over a 24 h period in September 1991. The Ría de Vigo, where red tides are frequent, normally shows a positive estuarine circulation and is subjected to seasonal upwelling and downwelling phenomena. A marked diel pattern was observed for five species that are capable of causing red tides (Ceratium furca, Scrippsiella trochoidea, Dinophysis acuminata, Mesodinium rubrum, and Eutreptiella sp.). Such diel behaviour could be clearly advantageous in a stratified environment where light and nutrients are often in two separate layers. Active movement enables species such as dinoflagellates and some ciliates to exploit high levels of irradiance at the surface during the day and to take up nutrients in deeper layers at night. Patchy distribution of phytoplankton, shear-induced horizontal dispersion and density variations were considered, but none of them accounted for the vertical changes observed. Vertical migration is thought to be one of the mechanisms that could promote blooms in nutrient-depleted surface layers.     

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.     

Gezeitenrhythmische Nahrungsaufnahme und Kotballenablage einer terrestrischen Milbe (Oribatei: Ameronothridae) im marinen Felslitoral

Feeding, locomotory and defecatory activity of the algivorous orbatid mite Ameronothrus marinus have been measured under defined conditions in the laboratory. Of terrestrial origin, these mites inhabit the intertidal zone of rocky shores along the North and Baltic Seas. Under daylight cycles found at the sampling site and in the absence of tides, feeding and defecation follow an endogenous rhythm with a spontaneous period of 12.3 h; in the tideless littoral of the Baltic Sea these activities are synchronized with diurnal cycles. Feeding and defecation follow a definite sequential pattern, repeated on the average every 4 h, i.e., 3 times between two high tides. The phase of this tidal feeding-defecation rhythm is determined by the onset of the high tide, and different patterns of this rhythm can be observed according to the tidal zones. Inhabitants of the lower eulittoral exposed to longer periods of submersion start feeding earlier, take up more food at one time before the beginning of high tide, quickly defecate part of the food undigested after feeding, and later produce other faecal pellets which contain the real remains of digestion. Inhabitants of the upper eulittoral, exposed to shorter periods of submersion, start feeding later and extend their feeding activity over the whole period of low tide; the difference between faecal pellets with digested and undigested contents is indistinct. The occurrence of these different types of faecal pellets in inhabitants of the intertidal zone is interpreted as a compensatory physiological adaptation resulting from increasing periods of submersion.     

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.     

Effects of temperature on activity,food consumption rates,and gut passage times of seaweed-eating<Emphasis Type="Italic"> Tegula</Emphasis> species (Trochidae) from California

Previous feeding studies on herbivorous marine snails rarely have focused on temperature effects on food intake. If temperature affects food intake, ectothermic snails may experience difficulty obtaining sufficient nutritional resources, limiting their ability to sustain populations at suboptimal temperatures. We hypothesized that the feeding responses of Tegula species would correspond with temperatures characteristic of their geographic distributions. We determined activity, consumption rates, and gut passage times at 11°C, 15°C, 19°C, and 23°C for three Tegula species with distinct thermal distributions: T. brunnea (cold water), T. aureotincta (warm water), and individuals from warm- and cold-water populations of T. funebralis, a broadly distributed species. Activity and consumption rates of T. aureotincta increased with increasing temperature, but were highest for T. brunnea at 19°C, a temperature rarely achieved in habitats occupied by this species, and lowest at 11°C. Warm-water T. funebralis showed significantly lower activity and consumption rates at 11°C, whereas cold-water T. funebralis consumed food fastest at 15°C and were most active at 23°C. Temperature affected gut passage time only in T. aureotincta. These data suggest that temperature might influence the northern limit of T. aureotincta by affecting activity and food consumption rates. T. brunneas activity and ability to consume food were not hindered by warmer temperatures despite the present day restriction of this species to colder waters. Also, widely separated (>300 km) T. funebralis populations may be adapted to regional conditions based on the different temperature responses of northern and southern snails.Communicated by P.W. Sammarco, Chauvin