Cardiac responses to abiotic factors in two tropical limpets, occurring at different levels of the shore

Laboratory experiments tested whether the cardiac activity of two species of limpets (Cellana spp.), co-occurring on the rocky shores of Hong Kong but having different zonal distributions, was differently affected by variations in physical environmental characteristics. Experiments reproduced exposure of limpets to dehydration, high water temperature and reduced salinity, all conditions that can occasionally be faced by animals during low tides. Cardiographic traces were obtained using a non-invasive method based on infrared light sensors glued onto the limpet shell in a position directly over the heart. The signals were filtered and amplified, viewed on portable oscilloscopes and automatically transferred to computer. When tested under acclimation temperatures, the lower shore C. toreuma had faster heart rates than the upper shore C. grata, both in air and when submerged in seawater. Regular alternation of heartbeating and acardia were observed in C. grata exposed to both dry and saturated air, whilst C. toreuma showed transient acardia only during exposure to dry air. Both species showed a similar increase in heart rate with increasing temperature (Q₁₀~1.8 in the range 25-34°C). When submerged in freshwater, C. grata regularly alternated acardia with beating at a rate slightly lower than in seawater, whilst C. toreuma showed strong bradycardia with reduction of beat amplitude, followed by steady acardia. Recovery of heartbeating after return to initial salinity was faster in C. grata, whilst C. toreuma, initially bradycardic, showed a consistent return to tachycardia. Overall, this study shows that the two congeneric species exhibit different cardiac responses to variations in environmental characteristics, which may be related to their differing vertical distribution.     

Ecological impacts on the limpet Lottia gigantea populations: human pressure over a broad scale on island and mainland intertidal zones

Here long-term monitoring data taken at 33 sites in southern and central California coast and islands were used to evaluate the size structure of the large intertidal limpet, Lottia gigantea in restricted-access and in easily accessible intertidal zones that encompass a wide range of ecological variables. Using multi-dimensional analysis of population size structures, we found that sites on islands and strictly protected mainland sites have significantly larger median limpet sizes and a greater range of limpet sizes than unprotected mainland sites, while no pattern occurs in latitudinal or regional comparison of sites. Although intertidal predators such as oystercatchers were not the primary focus of the monitoring efforts, extensive natural history notes taken during sampling visits support the argument that predation was not a primary cause for the size structure differences. Finally, substratum differences were determined not to have biased the observation of larger limpets in protected sites. In regard to human interactions with limpets, we conclude that the degree of enforcement against poaching is the better predictor of limpet size structure than proximity to population centers or visitation to intertidal sites.     

Factors affecting variability of foraging excursions in a population of the limpet Patella vulgata (Mollusca,Gastropoda)

Paths moved by the limpet Patella vulgata L. were monitored on a sheltered vertical rocky shore in North Wales using time-lapse photography throughout nocturnal low tides in April and November 1992, roughly corresponding to periods of minimum and maximum gonad ripeness, respectively. Various motion parameters, including total duration, total length, and maximum distance reached from home were computed from 124 complete foraging routes obtained from 18 higher-zoned and 18 lower-zoned limpets. P. vulgata typically performed a single loop per night (average total length, ca. 70 cm), moving from the individual home scar to graze the surrounding algal grounds (average maximum distance, ca. 25 cm), then following its own trail back home. On the average, limpets moved from the home scar during three of four available nocturnal low tides and exploited about half the emersion time. Evidence for size-related variability in behaviour was found, as a positive correlation was assessed between both total duration and legth of the excursions, and shell length. In addition, zonation was proved to affect the limpets' foraging behaviour, since low-shore limpets moved faster and covered greater distances than high-shore ones. Moreover, consistent seasonal variation in foraging behaviour emerged, since in April the excursions were longer and longer-lasting than in November, but limpets exploited a larger fraction of potential activity phases in November than in April. The within-population variability in the temporal and spatial characteristics of the foraging excursions is discussed in relation to the available data on zonal and seasonal variation of food resources and in relation to physiological changes due to reproductive cycle.     

Sex change in a giant endemic limpet,Patella kermadecensis,from the Kermadec Islands

Sex-ratios of the limpetPatella kermadecensis Pilsbry, collected at three times of the year during 1984 and 1985 at Raoul Island in the Kermadecs group, varied with the size of the limpets. Small limpets were predominantly male, while amongst larger ones about half were female. The presence of hermaphroditism inP. kermadecensis was confirmed by microscopic examination of gonad material from over 50 limpets, which revealed a high percentage of individuals with both male and female gametes in their gonads. It is postulated that protandric sex change occurs in this large limpet, and that this phenomenon is related to a change in habitat from the backs of large shells (limpets less than approximately 50 mm) to bedrock (larger limpets). The pattern of sex change inP. kermadecensis is compared with that in other species of limpet, and it is suggested that a reproductive system consisting of gonochoristic males and sequential hermaphrodites may be particularly advantageous for a large, slow-growing species that has a very restricted geographical range.     

How hot for how long? The potential role of heat intensity and duration in moderating the beneficial effects of an ecosystem engineer on rocky shores

Studies of the importance of ecosystem engineers have focused on their benefit to biodiversity through ameliorating environmental stress, without understanding the exact benefits gained by associated organisms. On monsoonal tropical shores, species experience strong seasonality in environmental conditions from almost temperate winters to tropical summers when mass mortalities can occur during daytime emersion. The limpet, Cellana grata, associates with an ecosystem engineer, the barnacle Tetraclita japonica, in summer, but not in winter. To understand the benefits of this association, physiological responses (heart rates and osmotic responses) of the limpet either amongst Tetraclita or on open rock surfaces were investigated under three environmental conditions in a laboratory mesocosm: awash (non-stressed), low (30 °C) and high thermal stress (40 °C) of varying durations (3 or 6 h). In general, at 30 °C, limpets showed similar physiological responses under all conditions for both exposure durations. After 6 h at 40 °C, however, all limpets on open rock surfaces died, whereas those associated with barnacles survived. The surviving limpets experienced similar levels of stress as those exposed to 40 °C on open rock surfaces for half the time (3 h), showing that both the level and duration of stress were important. Limpets, therefore, gain benefits from engineering species when conditions are extreme or stress is prolonged. Under low temperatures or for short durations (e.g. winter), associating with barnacles does not provide physiological benefits. Understanding how, and to what extent, associates benefit from ecosystem engineers highlights how close the margin between survival and mortality can be when conditions are extreme.     

Intra-specific variability in the temporal organisation of foraging activity in the limpet<Emphasis Type="Italic"> Patella vulgata</Emphasis>

Patella vulgata is one of the most common limpet species on rocky shores throughout the north-eastern Atlantic. Foraging activity of this species has been described during various different diel and tidal phases and several factors have been suggested that might influence the timing of its activity. Among these, inclination of rock surfaces seems to have a clear role, with limpets on horizontal rocks being active during daylight at high water, while limpets on vertical rocks were active during night-time low water. However, by comparing the results of previous investigations it is difficult to separate clearly the effect of any single factor from the effects of their interactions. In the present paper we investigated the simultaneous influence of height on the shore, slope of rock surface and date of sampling (August–September and March–April) on the behaviour of a population of P. vulgata on the Isle of Man (UK). The study was carried out using modern telemetric methods, allowing the activity patterns of limpets to be recorded for long periods of time. The results showed that limpets concentrated their activity during two distinct temporal windows—daytime tide-in and night-time tide-out—while no significant activity was observed during daytime tide-out or night-time tide-in. The observed behaviour appeared to be driven by complex interactions among the three factors examined, and the influence of rock slope alone was not so strong as to override the effect of other factors completely. In addition, a key finding of this study was the conspicuous variability in the individual behaviour of P. vulgata. The greatest level of variability was observed in limpets on vertical surfaces on the upper shore, which showed an extreme switch in behavioural patterns, according to date of sampling (daylight/tide-in foraging during the spring and night-time/tide-out foraging during late summer). In addition, individuals apparently sharing the same set of conditions (inclination, height on the shore and sampling date) differed in their behaviour, and a given behavioural pattern was not always shared by all individuals from a given group. Finally, variability was also observed at the intra-individual level since the same individual could be active during different tidal and diel phases. In conclusion, although the existence of endogenous rhythms has previously been demonstrated in this species, our data clearly show that P. vulgata was able to modulate its behavioural patterns in relation to a variety of exogenous determinants.Communicated by R. Cattaneo-Vietti, Genova     

Parasitic castration in slipper limpets infested by the symbiotic crab Calyptraeotheres garthi

Two ill-explored hypotheses might explain host castration by parasitic pea crabs. The ‘energy drain’ hypothesis states that castration is caused by host-derived nutrient consumption of parasites that ultimately diminishes host-energy intake. The ‘steric interference’ hypothesis states that castration occurs when parasites physically inhibit host reproduction. This study evaluated whether Calyptraeotheres garthi, a pea crab from the southwestern Atlantic, is a parasitic castrator and explored whether the two hypotheses above explain castration in the limpet Crepidula cachimilla. None of three studied limpet species brooded embryos during the reproductive season when infested by mature female pea crabs. Also, limpets of C. cachimilla infested by C. garthi did not reproduce during a 90-day experimental period while crab-free limpets did spawn embryos during this period. Limpets resumed reproduction soon after pea crabs were experimentally removed from their brooding chamber. Thus, C. garthi does castrate limpets, and castration is reversible. Pea crabs ‘steal’ food from limpets, and infested limpets did not modify their feeding behavior to counteract nutrient loss. Thus, infested limpets are expected to ingest less food which provides partial support for the ‘energy drain’ hypothesis. However, the limpet’s body condition increased or was not affected by pea crabs during the breeding season which argues against the same hypothesis. Furthermore, that limpets promptly recovered reproductive activity once pea crabs were experimentally removed, that castration was not induced by the smallest pea crabs in the population (that fill only partially the brooding chamber), and that parasitized limpets did exhibit fully mature ovaries, support the ‘steric interference’ hypothesis explaining parasitic castration.     

Analyses of limpet defense and predator offense in the field

Between November 1982 and August 1984 diving observations of two intertidal limpets' defensive behaviours in response to two predatory species were made near the Seto Marine Biological Laboratory of Kyoto University, in Wakayama Prefecture, Japan, for over 800 h. The focus of the observations was a simple isolated rock, 0.9 m² in area with an inclination of 75°. The pulmonate homing limpet Siphonaria sirius responded by fleeing from its home in response to the whelk Thais clavigera and by calmping tightly to its home scar in response to the starfish Coscinasterias acutispina. Both behaviours were successful for avoiding predation. After a whelk moved far away, limpets returned to their homes. The patellid limpet Cellana toreuma showed mantle folding behaviour or fled for an average distance of 8 cm vertically upwards in response to the whelk and for an average distance of 21 cm vertically upwards in response to the starfish. The amount of time predators spent foraging increased in the low intertidal zone relative to the mid and high zones, and most predator attacks were observed during the limpets' resting periods (submersion in late afternoon or nighttime). The upward directional flight displayed by C. toreuma was related to the upward orientation of the limpets' heads when they are in a resting position. Half of the limpets observed fleeing downward away from a starfish, a movement elicited by a downward attack, received a second attack. These limpets were preyed upon at the second encounter 67% of the time. The upward fleeing behaviour of C. toreuma resulted in the shifting of its resting site higher up the study site where fewer attacks occurred and where the mortality rate was at its lowest. It is suggested that the homing behaviour of S. sirius and the upward fleeing behaviour of C. toreuma are mechanisms by means of which interspecific competition between limpets is reduced and coexistence on a small rock surface is maintained.     

Ecological studies on a tropical limpet,Cellana radiata. Structural variations in the shell in relation to distribution

Structural variations in the shell of the tropical limpet Cellana radiata (Born) inhabiting the high and low water zones and the exposed and protected surfaces at Palm Beach, Waltair Coast, India, were investigated. Attempts have also been made to study which environmental factors have a marked influence upon the characters of the shell. The results are discussed and are compared with those on temperate climate limpets.     

Scarred limpets at hydrothermal vents: evidence of predation by deep-sea whelks

The chaotic physical and chemical environment at deep-sea hydrothermal vents has been associated with an ecosystem with few predators, arguably allowing the habitat to provide refuge for vulnerable species. The dominance of endemic limpets with thin, open-coiled shells at north Pacific vents may support this view. To test their vulnerability to predation, the incidence of healed repair scars, which are argued to reflect non-lethal encounters with predators, were examined on the shells of over 5,800 vent limpets of Lepetodrilus fucensis McLean (1988) that were collected from 13 to 18 August 1996. Three vent fields on the Juan de Fuca Ridge at ca. 2,200 m depth were sampled, two within 70 m of 47°56.87′N 129°05.91′W, and one at 47°57.85′N 129°05.15′W with the conspicuous potential limpet predators, the zoarcid fish Pachycara gymninium Anderson and Peden (1988), the galatheid crab Munidopsis alvisca Williams (1988), and the buccinid snail Buccinum thermophilum Harasewych and Kantor (2002). Limpets from the predator-rich vent were most often scarred, a significant difference created by the high incidence of scars on small (<4 mm long) limpets in this sample. Collected with the limpets were small (median shell diameter 4.4 mm) buccinids. They, rather than the larger, more conspicuous mobile fishes and crabs are argued to be the shell-damaging predator. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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     

Size-assortative mating and non-reciprocal copulation in a hermaphroditic intertidal limpet: test of the mate availability hypothesis

Mating behaviour and mating patterns are affected to a large extent by body size in both hermaphrodites and gonochorists. Detailed research on mating patterns, mate choice, pre- and post-copulatory sexual selection in marine pulmonates is wanting, thus warranting more attention as a study system in the future. The simultaneous hermaphroditic limpet Siphonaria capensis (Pulmonata: Basommatophora) shows size-dependent fecundity, and acts as a suitable organism to test the effects of body size on mating patterns, mating success, gender expression and reciprocity in hermaphroditic marine gastropods. We mainly used bootstrap resampling techniques to estimate the effects of different factors on mating patterns. In the populations studied, a strong size-assortative mating pattern was observed, where small-scale spatial distribution of potential mating partners (the mate availability hypothesis) could explain 65% of this pattern, while mate choice and mating constraints explaining the rest. No significant difference in mating success between limpets with different body size was found, even though in one population, the sperm recipients were larger than the non-copulating limpets. Interestingly, we found that intromission was non-reciprocal during copulations. This may mean that this species does not conform to the common rule of reciprocity predicted for hermaphrodites, unless there is sex-role alternation between individuals in a mating pair. The mating partners consisted of similar sized, acting males and females, thus without any indication of the body size determining the gender expression. The matings took place in early mornings only at spring tides and the animals were observed laying egg masses only during neap tides. Since siphonariid limpets possess both a spermatheca and a seminal vesicle, which may have either a sperm storing and/or digestive function, post-copulatory sexual selection (e.g., sperm competition) cannot be dismissed.     

Overestimations of food abundance: predator responses to prey aggregation

Understanding and predicting the consequences of trophic interactions for community processes requires knowledge of the role of food availability, which is often wrongly conflated with prey abundance. For prey animals in groups, this is not fully understood. Previous work has shown that oystercatchers more frequently attack solitary rather than aggregated limpets and are more successful in predation attempts on singletons. It has also been demonstrated that an attack on one limpet in a group alerts the entire group, all of which then clamp down and become unavailable. I show that Eurasian Oystercatchers (Haematopus ostralegus L.) attack only one limpet in a group and then move on to attack another individual limpet, and I also demonstrate that the distance they move is greater than the distance at which groups of limpets have been known to detect attacks. Thus in the oystercatcher-limpet predator-prey system on rocky shores, groups of limpets are actually one prey item independently of the number of limpets in the group. This has implications for assessment of food supply for avian predators on rocky shores, with consequences for our understanding of previously documented trophic cascades.     

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.     

Intraspecific density mediates sex-change in the territorial patellacean limpet Lottia gigantea

The effects of intraspecific density and agonistic interactions on sex-change were studied in the territorial limpet Lottia gigantea. In a one-year field experiment (1982–1983) on San Nicolas Island off the southern California coast, USA, male limpets transplanted to large enclosures changed sex more frequently than those transplanted to small enclosures (9 of 13 vs 1 of 10; p=0.013), indicating that intraspecific density can profoundly influence the probability of sex change. Large limpets were more likely to change sex than small ones. Observations of gender-age distributions as well as field behavior suggested that each limpet's territorial status prior to the experiment may have been an important component of this size effect, although other interpretations including an effect of age are possible. Pooling the results with those of two previous studies confirmed that sex-change is enhanced by low density. This enhancement was observed among the largest members of a local population in the first year of each experiment, while among the smaller members the enhancement was delayed until the second or third year. Low density may be a correlate of high mortality, and therefore an adaptive cue for an earlier age of sex change. Dominant territorial status correlates with an individual's size, and therefore egg-producing capacity, relative to its neighbors, and thus may also be a good cue for the initiation of sex-change.     

A multi-factorial experiment on heart rate variations in the intertidal crab Pachygrapsus marmoratus

A multi-factorial experiment was designed to investigate the effect of the following factors on the cardiac activity of the intertidal crab Pachygrapsus marmoratus: respiratory medium (air, water), temperature (four levels, 10, 17.5, 25, 32.5 °C), season (winter, summer) and body size (two levels, carapace ≤21 mm and carapace >21 mm). The results showed that the heart rate of P. marmoratus increased linearly with temperature and decreased when the specimens were exposed to air rather than water. Moreover, the heart rate values in summer were lower than those in winter at the corresponding temperature and body size. The summer heart rate–temperature regression line was laterally (to the right) shifted with respect to the winter line, suggesting a seasonal acclimation. Body size affected heart rate only at the acclimation temperature (17.5 °C), while no significant effect was detected at lower or higher temperatures. During the reproductive season a separate experiment was carried out to assess the effect of sex and reproductive status on heart rate. No significant difference was found among mean values of males, berried females and females without eggs. The results of the present study confirm the high physiological plasticity of this species, suggesting that P. marmoratus is a truly amphibious crab, able to deal with both water- and air-breathing during its activity. Received: 5 January 1999 / Accepted: 7 July 1999     

Food availability and diet composition of three coexisting Mediterranean limpets (Patella spp.)

The gut contents of three intertidal patellid limpets were analysed by collecting foraging specimens on a breakwater on the Tyrrhenian coast (central Italy) between May 1988 and October 1989. The three species coexist there showing a different, but partially overlapping zonation: Patella aspera dominates the infralittoral fringe; the majority of P. caerulea inhabits the lower midlittoral, while P. rustica is most abundant in the upper midlittoral. The algae present on slivers of substratum over which each limpet collected was moving were identified. Moreover, floristic surveys were made along the shore in order to characterize the algal cover of the different zonal belts. The floristic study revealed that the basic elements of algal communities typical of western Mediterranean rocky shores are present in the study area. The algae found on the slivers under the foraging limpets were generally representative of the algal community typical of the same zone. There was a marked difference between the diets of P. rustica and P. aspera due to the fact that the first species forages on a few low lying epilithic and endolithic Cyanophyceae, while P. aspera feeds on a large number of species belonging to all the main algal classes and life forms considered, including frondose epilithics and epiphytics. The diet of P. caerulea resembles that of P. aspera in algal heterogeneity, but is dominated by Cyanophyceae as in P. rustica. A detailed analysis of the differences between gut contents of each limpet species and the relative slivers showed an obvious general correspondence, but revealed also that the diets of the three species do not completely reflect the availability of algae. These findings suggest that the basic diet segregation mechanism between the three populations is their zonal separation. However, the difference in gut contents of heterospecific limpets foraging in the same zone suggests the existence of supplementary morphy-functional or behavioural mechanisms for diet segregation between the three species.     

Mineralization of major lateral teeth in the radula of a deep-sea hydrothermal vent limpet (Gastropoda:Neolepetopsidae)

Radular teeth of a neolepetopsid patellogastropod limpet, Paralepetopsis ferrugivora, from a deep-sea hydrothermal vent site have similar elemental composition (O, Si, P, S, Cl, K and Fe) to radular teeth of the shallow-waters patellogastropod limpets. However, in contrast to shallow waters limpets, the fully mature teeth of P. ferrugivora do not show any crystalline phases. Amorphous silica was found in the cusp of the teeth and amorphous iron oxide in the junction zone and base of the teeth. Ferritin-rich vesicles were observed in cells adjacent to the junction zone of the early mature teeth, suggesting that these vesicles can mediate the delivery of iron to the tooth matrix. The similarity in the elemental composition between the hydrothermal limpet P. ferrugivora and shallow-waters patellogastropod limpets shows that the extreme hydrothermal environment did not alter the elemental composition of the radular teeth in the deep-sea species.Communicated by P. W. Sammarco, Chauvin     

Carbon emission associated with respiration and calcification of nine gastropod species from the intertidal rocky shore of Western Europe

Intertidal rocky shores are characterized by vertical zonation that results from the interplay between environmental conditions, organism physiology, and species interactions. Metabolism of intertidal organisms is highly variable between species and it changes with vertical position along the intertidal gradient. The present study aimed to quantify the carbon metabolism of nine intertidal rocky shore gastropods, in order to clarify their respective roles in carbon production during emersion and immersion. The influences of monthly temperature variation and tidal level were tested for each species. Analyses were performed in the laboratory using the infrared gas analyzer method for measuring aerial respiration rates, and the dissolved inorganic carbon and total alkalinity technique for measuring aquatic respiration rate and calcification. Hourly carbon fluxes were calculated for the mean annual temperature of 13 °C measured in both air and underwater in the study area. Respiration rates were similar for emersion (8–25 μmol CO₂ g AFDW^?1 h^?1) and immersion (10–23 μmol DIC g AFDW^?1 h^?1). For all species, underwater respiration fluxes were more influenced by monthly temperature variation than by air fluxes, probably as an adaptation to the rapid changes occurring during emersion. Calcification was an important factor influencing annual carbon fluxes for all studied species; every species showed different calcification rates according to its size and position on the intertidal zone. Annual carbon emissions were calculated using the mean immersion/emersion time of each species. Intertidal gastropod carbon emission was primarily influenced by body biomass and their vertical position within the intertidal zone.     

Life-history features of the marine intertidal limpet Notoacmea scutum (Gastropoda) in central California

Growth of individually marked specimens of Notoacmea scutum (Rathke) was followed for 15 months (April, 1978 to June, 1979) at a field site in central California, USA. Growth rates were highly seasonal, being highest from late spring through early summer and lowest during winter. Growth rate also varied significantly with intertidal height: limpets lower in the intertidal zone grew faster. Both relationships may have been related to the relative abundance of food or to the amount of time available for foraging. Recently settled limpets, which were present throughout the year, grew to approximately 17 mm in length by the end of their first year; by then, they had also reached the size at which mature gonads were consistently found (16 mm). Limpets were almost 26 mm long by the end of the second year. Substantial mortality occurred throughout the year, but it was particularly severe during winter, when a large portion of the reproductive population was lost. This pronounced winter mortality was probably not caused by invertebrate predators such as seastars. Most limpets at the study site were less than 2 yr old.