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International audience; Sclerochronological data from archaeological shell middens provides crucial information about past environments and subsistence practices of coastal human populations. These studies rely on a strong knowledge of species-specific growth patterns, as it is the main driver of the quantity and quality of information recorded in biogenic carbonates. The limpet Patella vulgata is highly represented in eastern Atlantic shell middens. In this study, the micro-increment formation of its shell was investigated through in situ marking experiments, using calcein fluorochrome (150 mg.L−1) coupled with cohort study and local SST recording. Calcein-stained growth structures confirmed the efficacy of the fluorochrome, and our results shows a disturbed tidal increment formation, with on average more than one increment per day, but less than one increment per tide. Growth is not mainly influenced by environmental factors, but could be forced by competition among limpets and endogenous factor (age and sex), resulting in a highly heterogenous population in terms of growth rates. Growth interruption could not be detected solely by micro-increment observation, causing a major issue for cross-calibration with recorded temperatures, sclerochronological estimation of growth and isotopic profiles. For palaeo-environmental application, we recommend to consider that there is no general population pattern, even at a micro-habitat scale.

Limpets and barnacles are important components of intertidal assemblages worldwide. This study examines the effects of barnacles on the foraging behaviour of the limpetPatella vulgata, which is the main algal grazer in the North-west Atlantic. The behaviour of limpets on a vertical seawall on the Isle of Man (UK) was investigated using autonomous radio-telemetry, comparing their activity patterns on plots characterized by dense barnacle cover and plots from which the barnacles had been removed. Limpet behaviour was investigated at mid-shore level, but two different elevations were considered. This experiment revealed a significant effect of barnacle cover on the activity ofP. vulgata. Limpets on smooth surfaces spent a greater proportion of total time active than did limpets on barnacles. Movement activity was also greater in areas that were lower down in the tidal range. In general, limpets were either predominantly active during diurnal high or nocturnal low tides and always avoided nocturnal high tides. Individuals on barnacles at the higher elevation concentrated their activity during nocturnal low water. All the other groups of limpets (smooth surfaces on the upper level and all individuals on the lower shore) had more excursions centred around daylight hours with an equal distribution of activity between periods of low and high water. Inter-individual variability was, however, pronounced.

In space limited intertidal habitats, non-specific substratum generalists can be observed to increase their survival and settlement opportunities by growing upon other living organisms. To date, few studies have assessed the influence of multiple environmental factors in determining patterns of epibiont diversity and community structure upon gastropod shells, across spatial scales. This study investigated the significance of surface salinity range, wave exposure and shell metrics in determining the composition and diversity of epibionts found upon shells of Patella vulgata (Common Limpet), throughout the Plymouth Sound and lower reaches of the Tamar Estuary, United Kingdom (UK). A total of 292 shells were sampled, identifying 26 epibiont species amongst 9 phyla. Species richness and diversity (H’) were greatest upon shells at exposed marine sites (where salinity variation was lower), decreasing as surface salinity range increased towards the Lynher River. Multivariate analysis revealed epibiont community composition to differ significantly between sites, being influenced mainly by surface salinity range and, to a lesser extent, wave exposure. Species contributing to community dissimilarities were identified to reflect the dominant species between sites, with observed community shifts attributed mainly to a species physiological tolerance to environmental conditions. This study is the first to describe epibiont community variation upon shells of P. vulgata along an estuarine-exposure gradient in the UK. Although salinity variation and wave exposure explained most of the observed community composition patterns, further investigation including exploration of additional environmental factors, is required to fully explain the distribution and diversity patterns of epibionts upon shells of P. vulgata.

Using a combination of geostatistical methods and generalized additive models (GAMs), we have analyzed the combined effect of spatial, environmental and biological factors on the density of Patella vulgata, Patella depressa and Patella ulyssiponensis, three limpet species coexisting in rocky intertidal areas of the Galician coast (NW Iberian Peninsula). The best multivariate GAM fit to the data explained, on average, 54% of the observed variation in density of these species, revealing the influence of spatial (latitude and longitude) and environmental factors (temperature). The influence of biological factors such as reproductive status and density of the species, as well as those of coexisting species, was also detected. The results suggest the existence of spatial and temporal segregation behaviours to decrease intra- and interspecific competition. They confirm P. vulgata as a migratory species and reveal P. depressa also as a migrant limpet, while P. ulyssiponensis belongs to the non-migratory group. Geostatistical analysis showed that P. depressa is the dominant species in areas more exposed to wave action, whilst P. vulgata dominates sheltered area. In addition, the three limpet species are spatially structured in patches, whose density and size vary according to the type of coast (Atlantic/Cantabrian) and to the degree of wave exposure. These results reveal the importance of carrying out assessments at the local level and in those areas particularly vulnerable to exploitation, such as sheltered areas, where densities and recruitment are lower. This information can be useful for decision-making aimed at the management of this resource, as well as for the efficient distribution of effort with respect to geographical fishing areas.

Zooarchaeological faunal remains are commonly examined to investigate harvesting behavior. We determined limpet (Patella vulgata) shell size and shape, and estimated shell age from several middens ...

Long-term cyclical changes in rocky shore community structure were documented over two decades at sheltered shores in Lough Hyne Marine Reserve, County Cork, Ireland. Three peaks of abundance were recorded for the limpet, Patella vulgata (1990-5, 2002-6 and 2010-14) with oscillations that varied in amplitude and frequency among sites. The cover of the fucoid Fucus vesiculosus varied inversely with limpet abundance and showed the strongest negative correlation with a lag time of 0–2 years. The species complex Fucus spiralis/guiryi showed a weaker correlation with a lag time of 1–2 years. Two other fucoid species showed no such negative correlations despite their close proximity to limpets within the lough's compressed tidal range. There was no relationship between overall barnacle cover (dominated by Austrominius modestus ) and the limpet-fucoid cycles, suggesting that the shelter provided by A. modestus for algae to escape from limpet grazing pressure may not be necessary for these cycles to occur on wave-sheltered shores.

Parasites are ubiquitous throughout nature and can have dramatic effects on their hosts. Although much is known about the pathology of parasites, the environmental factors influencing the distribution and abundance of parasites are poorly understood. Investigations into these factors could help predict the effect of parasites on the functioning of ecosystems. The limpet Patella vulgata is a key grazer on European rocky shores and is the first intermediate host for the trematode parasite, Echinostephilla patellae. This study investigated the spatial distribution of E. patellae in relation to P. vulgata at fourteen rocky shores around Ireland. Despite P. vulgata recruiting into rock pools, adults are more abundant on emergent rock and experience greater mortality in pools compared to emergent rock. It has been suggested that greater parasite prevalence in pools may be a factor driving this disjunct distribution pattern. Prevalence of infection was compared among rock and pool habitats. Size and sex of the host were also investigated in order to determine whether host phenotype influenced levels of infection prevalence. Results varied considerably among study sites, probably due to the heterogeneity of environmental conditions. No clear pattern emerged in relation to infection prevalence between habitats but this may be owing to the low numbers of limpets collected from pools. There was a significant positive relationship between infection prevalence and increasing host size. Individuals for which sex was indeterminate were more likely to be infected than distinct males or females, suggesting that infection may be causing castration and subsequent removal of these individuals from the breeding population. We discuss the importance of considering host-parasite dynamics in a period of rapid environmental change.

The use of limpets of the genus Patella as indicators of environmental quality is a priori promising, but few data are available in relation to their sensitivity to toxicants or their ability to accumulate pollutants. The aim of this work was to evaluate the use of Patella spp. as model organisms by (1) assessing the sensitivity of limpet embryos to some common pollutants, (2) validating the use of the limpet embryo-larval bioassay using natural sediment samples, and (3) analyzing the presence of pollutants by using Patella spp. as sentinel species. The obtained EC50 values ranged between 33 and 84 µg/L for Cu, 749–1163 for Cd, 317–1103 for Zn and 813–3411 µg/L for Pb; these sensitivity ranges were similar to those of other marine invertebrates used in ecotoxicology. It was not possible to validate the bioassay with natural samples, since the sampled sites did not cause significant toxicity. Patella spp. accumulated metals and organic pollutants in their soft tissues, showing higher concentrations than mussels for some metals (Cd, As and Cu) and lower concentrations for others (Hg, Zn, Cr), as well as for organic pollutants. Furthermore, bioaccumulation in limpets showed a spatial pattern of contamination similar to that shown by mussels and sediments, revealing limpets as a very good substitute/complement to mussels for biomonitoring marine pollution, especially in sites where mussel populations are absent or present at low numbers.

The reproductive cycle and the physiological status, measured through a standardized condition index, of limpets Patella depressa and Patella ulyssiponensis were studied along the Galician coast (NW of the Iberian Peninsula) from July 2010 to June 2011. The observed data were adjusted using linear and nonlinear general models. This allowed analyzing and quantifying the influence of independent factors, such as individual size and degree of wave exposure, on the cycles. Populations of P. depressa showed mature individuals nine months a year with a resting period from June to September. The amplitude of the fertile period increased with limpet size and with wave protection. On the other hand, the condition index decreased with increasing limpet sizes and, for a given size, it was higher in sheltered areas than in exposed ones. P. ulyssiponensis showed mature individuals eleven months a year with a small resting period in May or June. The amplitude of the reproductive cycle increased with limpet size but this variable did not show any significant influence on the condition index. No size/sex relation was observed either for P. depressa or P. ulyssiponensis. The results have contributed some valuable information to help design exploitation plans: Given that the limpet exploitation in these coasts is multispecific and includes a third species Patella vulgata, in case the resource is intended to be managed by temporal closures during the fertile period, it should be established following the fertile period of P. vulgata, which is restricted to 3–4 cold months, while the studied species show mature individuals throughout much of the year. It is recommendable to not harvest the largest individuals, since energetic condition decreases with size; in addition, larger individuals show longer maturity periods and might therefore contribute to recruitment to a greater extent. Size distribution observed in sheltered areas showed a lack of juveniles. This can be pointing to lower recruitment, suggesting caution in order to avoid overexploitation in these areas.

As climate change is expected to impose increasing thermal stress on intertidal organisms, understanding the mechanisms by which body temperatures translate into major biogeographic patterns is of paramount importance. We exposed individuals of the limpet Patella vulgata Linnaeus, 1758, to realistic experimental treatments aimed at disentangling the contribution of water and air temperature for the buildup of thermal stress. Treatments were designed based on temperature data collected at the microhabitat level, from 15 shores along the Atlantic European coast spanning nearly 20° of latitude. Cardiac activity data indicated that thermal stress levels in P. vulgata are directly linked to elevated water temperature, while high air temperature is only stressful if water temperature is also high. In addition, the analysis of the link between population densities and thermal regimes at the studied locations suggests that the occurrence of elevated water temperature may represent a threshold P. vulgata is unable to tolerate. By combining projected temperatures with the temperature threshold identified, we show that climate change will likely result in the westward expansion of the historical distribution gap in the Bay of Biscay (southwest France), and northward contraction of the southern range limit in south Portugal. These findings suggest that even a minor relaxing of the upwelling off northwest Iberia could lead to a dramatic increase in thermal stress, with major consequences for the structure and functioning of the intertidal communities along Iberian rocky shores.

The use of three limpet species, Patella vulgata Linnaeus, 1758, Patella depressa Pennant, 1777 and Patella ulyssiponensis Gmelin, 1791 as model organisms in marine ecotoxicology has been evaluated. Initial laboratory experiments were aimed to standardize a biological test with embryos and larvae of Patella spp, establishing the percentage of normal trochophore larvae as endpoint. Before conducting in vitro fertilization, oocytes must be maturated artificially by incubation in an alkaline solution; therefore, alkalinizing agent, pH and time of eggs alkalinization were evaluated. Moreover, time of sperm activation, optimum sperm and oocytes concentration during fertilization, gamete contact time, use of stirring during the fertilization, egg concentration and incubation temperature were examined. Minimum sample size per treatment was also estimated. Exposure of oocytes for 10min to FSW alkalinized with NH4OH at pH 9.0, the use of undiluted sperm pre-activated during 45min and a concentration of 200 oocytesmL(-1), a gamete-contact time of 180min and egg incubation at 18°C during 24h at a concentration of 80 eggsmL(-1) were the conditions allowing maximal embryo-larval development success. With an error of 0.05, a sampling size ≥320 allows a 95% confidence in the estimate. This Patella spp. acute bioassay fulfills a number of important a priori requirements to be used in ecotoxicological studies. Nevertheless, in vitro fertilization requires considerable handling, which may lead to failure in fecundation. Such difficulties are also addressed, in order to facilitate the routine use of this protocol by other laboratories.

This study examines the importance of thermal refugia along the majority of the geographical range of a key intertidal species (Patella vulgata Linnaeus, 1758) on the Atlantic coast of Europe. We asked whether differences between sun-exposed and shaded microhabitats were responsible for differences in physiological stress and ecological performance and examined the availability of refugia near equatorial range limits. Thermal differences between sun-exposed and shaded microhabitats are consistently associated with differences in physiological performance, and the frequency of occurrence of high temperatures is most probably limiting the maximum population densities supported at any given place. Topographical complexity provides thermal refugia throughout most of the distribution range, although towards the equatorial edges the magnitude of the amelioration provided by shaded microhabitats is largely reduced. Importantly, the limiting effects of temperature, rather than being related to latitude, seem to be tightly associated with microsite variability, which therefore is likely to have profound effects on the way local populations (and consequently species) respond to climatic changes.

Exploitation of key consumers can have major consequences for community and ecosystem functioning. Limpets are key grazers exploited in regions such as Macaronesia, southern Africa, Chile and California. Here we describe a field experiment designed to simulate human exploitation of British limpets that are unexploited and used as model populations. Our aim was to evaluate the effects of size-selective harvesting on the composition of the rocky shore community of non-target species. Limpet populations were subjected to simulated exploitation of large size classes for 18 mo at 2 locations in the southwest of England, by systematic removal at 2 different intensities: low and high exploitation compared with unexploited plots. The exploitation of limpets led to establishment of Fucus spp. to differing degrees at each location, but while variation in percentage cover of Fucus spp. decreased over the course of the experiment in unmanipulated control plots, it increased in plots with either low or high exploitation. Multivariate analyses showed that communities at the 2 locations responded differently to the same intensity of exploitation: unmanipulated controls were similar to low-exploitation treatments at Constantine, while at Trevone low-exploitation treatments were similar to high-exploitation treatments. This was mainly due to increases in percentage cover of F. vesiculosus var. evesiculosus with exploitation, indicating that site-specific differences in assemblage structure and the size structure of the harvested populations will determine its assemblage-level responses. Therefore, reductions in density of grazers may have divergent consequences for different rocky shore communities.

The limpet Patella vulgata is a key species of northern Atlantic rocky shore-associated communities, and is commonly considered to be important in regulating populations of canopy-forming Ascophyllum nodosum, through consumption of propagules and young recruits. Although P. vulgata is usually regarded as a non-selective epilithic biofilm grazer, a role in the collapse of established A. nodosum through grazing of adult plants has been repeatedly suggested. Factors controlling the preference of P. vulgata for epilithic biofilm or adult algae are still not clearly established. Here, we test the hypothesis that the diet of P. vulgata is mainly driven by the local availability of food sources. Limpets were sampled along the first 6 metres of an A. nodosum bed–bare rock gradient. Stable isotope ratios of their muscle tissue and digestive glands were measured. The contribution of A. nodosum to the diet of limpets was the highest in the immediate vicinity of macroalgae beds, which confirmed our initial hypothesis. However, the contribution of epilithic biofilm did not match our hypothesis, being the lowest for limpets colonizing bare rock. Instead, these limpets relied on a wide array of sources, including ephemeral green algae, biofilm and drifting A. nodosum fragments. Overall, our results indicate that A. nodosum can be readily grazed by limpets, which challenges the hypothesis that these macroalgae dominate rocky shores due to the absence of strong top-down control exerted by herbivores. Our results also highlight the need to consider the small spatial scale to understand the dynamic of herbivore–algae interactions in natural environments.

Sex change, or sequential hermaphroditism, occurs in the plant and animal kingdoms and often determines a predominance of the first sex. Our aim was to explore changes in sex ratios within the range of the species studied:Patella vulgataandPatella depressa. The broad-scale survey of sex with size of limpets covered a range of latitudes from Zambujeira do Mar (southern Portugal) to the English Channel. Indirect evidence was found for the occurrence of protandry inP. vulgatapopulations from the south of England, with females predominating in larger size-classes; cumulative frequency distributions of males and females were different; sex ratios were biased towards males and smallest sizes of males were smaller than the smallest sizes of females. In contrast in Portugal females were found in most size-classes ofP. vulgata. InP. depressapopulations from the south coast of England and Portugal females were interspersed across most size-classes; size distributions of males and females and size at first maturity of males and females did not differ.P. depressadid, however, show some indications of the possibility of slight protandry occurring in Portugal. The test of sex ratio variation with latitude indicated thatP. vulgatasex ratios might be involved in determining the species range limit, particularly at the equatorward limit since the likelihood of being male decreased from the south coast of England to southern Portugal. Thus at the southern range limit, sperm could be in short supply due to scarcity of males contributing to an Allee effect.

The reproductive cycle and physiological status of five populations of Patella vulgata have been studied along the Galician coast from July 2010 to June 2011. The use of generalized linear model has allowed to obtain parameters such as Day50 and Day95, which helped compare reproductive cycles among populations and allowed to determine the significance of the effect of size and degree of exposure on the start and duration of the cycle. On the other hand, a standardized condition index has been established, which enables the comparison of samples collected in different sites or seasons and composed by individuals of different size. Moreover, the use of a generalized nonlinear least squares model has allowed assessing the temporal variability of the physiological status of the populations and the effect of size and degree of exposure on its different components. This has contributed some useful information for planning a sustainable exploitation of this resource: sheltered areas turned out to be more productive, in terms of biomass. The reproductive cycle starts earlier and its duration is longer in sheltered areas. In addition, organisms showed minimum condition indexes during spawning periods, which makes these periods less suitable for the exploitation, both from a biological and from a commercial point of view. Larger organisms showed longer spawning periods. This highlights the importance of taking size composition into account, seeking to maintain a size structure which enables a reproductive period which is long enough to ensure future recruitment success.

Humans have contributed to phenotypic and demographic changes in their prey from very early on in the colonization of Europe, including the harvesting of shellfish in coastal ecosystems. We estimated trends in population growth (variation in the number of individuals) from DNA sequences of modern specimens in two North Iberian molluscs, top shells (Osilinus lineatus, from 24 sequences and 14 haplotypes) and limpets (Patella vulgata, taken from the bibliography), which were subjected to very different levels of harvesting pressure during the Upper Palaeolithic (~ 20000 to ~ 6000 years ago). The less harvested Osilinus top shells experienced fluctuations in population numbers coincident with climatic oscillations. Patella limpets, which were harvested in greater numbers, suffered clear and uninterrupted decreases in their numbers during the Upper Palaeolithic. These trends coincided with morphological changes in shell size (length or width) in the same direction (i.e., shell size decreased when population size decreased and vice versa). The differing trends seen in taxa subjected to different intensities of harvesting pressure suggest that climate effects were overcome by anthropogenic selection (leading to a smaller average length) in limpets. We suggest that intense fishing pressure may have induced irreversible shell length decreases in the most exploited species.

International audience; Understanding the physiological abilities of organisms to cope with heat stress is critical for predictions of species’ distributions in response to climate change. We investigated physiological responses (respiration and heart beat rate) of the ectotherm limpet Patella vulgata to heat stress events during emersion and the role of seasonal and microclimatic acclimatization for individual thermal tolerance limits. Individuals were collected from 5 microhabitats characterized by different exposure to solar radiation in the high intertidal zone of a semi-exposed rocky shore in winter and summer of 2014. Upper thermal tolerance limits (heat coma temperatures – HCTs, and heart rate Arrhenius break temperatures - ABTs) were determined for individuals from each microhabitat in both seasons under laboratory conditions. While we found a clear seasonal acclimatization, i.e., higher HCTs and ABTs in summer than in winter, we did not find evidence for microhabitat-specific responses that would suggest microclimatic acclimatization. However, operative limpet temperatures derived from in-situ temperature measurements suggest that individuals from sun exposed microhabitats have a much narrower thermal safety margins than those from less exposed surfaces or within crevices. Microhabitat specific thermal safety margins caused by high thermal heterogeneity at small spatial scales and the lack of short term acclimatization will likely shape small scale distribution patterns of intertidal species in response to the predicted increase in the frequency and intensity of heat waves.

Human activities have an impact on extant biotic communities, and may have had just as important an impact in the past. We assess human impact on limpet assemblages during the Upper Palaeolithic in Asturias (north-west Spain). The intensely exploited genus Patella exhibited a marked size decrease and a change in species assemblage composition, substituting the larger species P. vulgata for the smaller P. depressa. The present Patella assemblages in the upper tidal level exhibit the same pattern as those of the Epipalaeolithic (approx. 12 000 to 6000 years before the present). Although climate change may have contributed to such species replacement, spatial differences between close areas with different densities of Palaeolithic human settlements indicate unequivocal human impact. Present Patella species sampled from the region exhibit genetic signatures of past bottlenecks in mitochondrial DNA, which also indicate recent demographic expansion, suggesting that old impacts have been sufficiently important to leave genetic traces in current populations. Copyright © 2011 John Wiley & Sons, Ltd.

Concentration of 99Tc has been measured in fucoids and molluscs, sampled in a sheltered intertidal at the southwest coast of Norway from February to November 2006. The concentrations of 99Tc in molluscs differed significantly between species. The filtering bivalve Mytilus edulis had the lowest concentrations with averages of 2.3–5.9 Bq kg−1 d.w., while the herbivorous gastropods Littorina littorina, Littorina obtusata and Patella vulgata had higher concentrations. P. vulgata and L. obtusata had the highest concentrations, 40–47 and 26–30 Bq kg−1 d.w., respectively. L. obtusata has a specialized habit of living, and prefers to feed on fucoids. P. vulgata can graze extensively on the fucoid Ascophyllum nodosum when available. Fucoids are known to have very high uptake of 99Tc, and this was also found in the present study. The high 99Tc-concentrations of L. obtusata and P. vulgata are most likely due to their habit of feeding on fucoids.

Evidence for the exploitation of marine resources has been identified throughout prehistory but its relative importance has varied. Knowledge for earlier periods is biased by low archaeological visibility, and so the majority of hypotheses support the occurrence of intensification in the exploitation of marine resources during the Holocene. In the Cantabrian region (northern Spain) intensification has been proposed by some authors and rejected by others and there is also debate over when it began. Marine molluscs have been one of the most studied resources to support or reject intensification, and these studies have been mainly focused on variations of shell size through time. The hypothesis supporting intensification suggests that the decrease in molluscan size during the Holocene was caused by human pressure, whilst the alternative hypothesis is that this was induced by climatic and environmental conditions. In this paper, I study change of shell size and age distributions in northern Spain leading up to and after the Pleistocene–Holocene transition (35,000–5700 cal BP) using samples of four gastropod species from 18 shell midden sites. The species studied are the most abundant in the regional shell middens: Patella vulgata Linnaeus 1758, Patella intermedia Murray in Knapp 1857, Osilinus lineatus (Da Costa, 1778) and Littorina littorea (Linnaeus, 1758). The trends observed in both shell size and age structures shows that changes during the Upper Paleolithic may be due to climatic factors, but the constant decrease in both variables during the Mesolithic and the early Neolithic was caused by human pressure.

Shellfish metrical data are a source of information about the exploitation of marine resources in the past. In this study, we propose a methodological approach based on the size structures of different rocky intertidal gastropod species. Three limpet species (Patella vulgata, Patella intermedia and Patella ulyssiponensis) and the toothed topshell Osilinus lineatus are studied from two sites in Cantabrian Spain: La Garma A and Los Gitanos caves over a period of 10 000 years, covering the Pleistocene–Holocene transition. Data are also supplied about a further sea snail species, the periwinkle Littorina littorea (Upper Magdalenian). A reduction in size can be seen, between the upper Magdalenian and the late Neolithic, in the case of the first four species. The explanation for this decline is probably related to the climate change that occurred in the transition between the Late Pleistocene and the Holocene, but it is possible that human impact might also have influenced shell sizes in the Mesolithic and Neolithic.

There is substantial evidence from terrestrial and freshwater systems of species responding to climate change through changes in their phenology. In the marine environment, however, there is less evidence. Using historic (1946–1949) and contemporary (2003–2007) data, collected from rocky shores of south-west Britain, we investigated the affect of recent climate warming on the reproductive phenology of two con-specific intertidal limpet grazers, with cool/boreal and warm/lusitanian centres of distribution. Reproductive development in the southern limpet, Patella depressa, has advanced, on average, 10.2 days per decade since the 1940s, with a longer reproductive season and more of the population reproductively active. The peak in the proportion of the population in advanced stages of gonad development was positively correlated with sea surface temperature (SST) in late spring/early summer, which has increased between the 1940s and 2000s. The advance in peak reproductive development of this species is double the average observed for terrestrial and freshwater systems and indicates, along with other studies, that marine species may be responding faster to climate warming. In contrast, the northern limpet, Patella vulgata, has experienced a delay in the timing of its reproductive development (on average 3.3 days per decade), as well as an increase in reproductive failure years and a reduction in the proportion of the population reaching advanced gonad stages. These results are the first to demonstrate a delay in the reproductive development of a cool-temperate, winter spawner, towards cooler more favourable environmental conditions in response to climate warming. Such a delay in spawning will potentially lead to trophic miss-matches, resulting in a rapid nonlinear decline of this species.

The effect of short-term (5 days) exposure to CO2-acidified seawater (year 2100 predicted values, ocean pH = 7.6) on key aspects of the function of the intertidal common limpetPatella vulgata(Gastropoda: Patellidae) was investigated. Changes in extracellular acid–base balance were almost completely compensated by an increase in bicarbonate ions. A concomitant increase in haemolymph Ca2+and visible shell dissolution implicated passive shell dissolution as the bicarbonate source. Analysis of the radula using SEM revealed that individuals from the hypercapnic treatment showed an increase in the number of damaged teeth and the extent to which such teeth were damaged compared with controls. As radula teeth are composed mainly of chitin, acid dissolution seems unlikely, and so the proximate cause of damage is unknown. There was no hypercapnia-related change in metabolism (O2uptake) or feeding rate, also discounting the possibility that teeth damage was a result of a CO2-related increase in grazing. We conclude that although the limpet appears to have the physiological capacity to maintain its extracellular acid–base balance, metabolism and feeding rate over a 5 days exposure to acidified seawater, radular damage somehow incurred during this time could still compromise feeding in the longer term, in turn decreasing the top-down ecosystem control thatP. vulgataexerts over rocky shore environments.

The importance of external and internal population processes in determining variation in reproductive output and variation in population size were quantified with model simulations for open populations of the sequentially hermaphroditic limpet Patella vulgata using field data from the Isle of Man and South West Ireland. Cross-correlation analyses of model outputs and elasticity analyses show that population dynamics are dominated by the effects of large females, and that recruitment adds little to reproductive output. However, populations experiencing low but highly variable recruitment appear male limited and recruitment pulses carrying young males into the population are correlated to reproductive output with a 2–5-year lag. We conclude that pulses in recruitment can be a major structuring force in these limpet populations, but site-specific post-recruitment processes will determine the relative importance of recruitment to population dynamics and the lag between recruitment and reproductive output.

Laboratory investigations of environmental factors controlling early development and larval survival of free-spawning marine invertebrates are often limited by the difficulty of inducing spawning under laboratory conditions. Patellid gastropod species must be dissected in order to obtain the gametes from the gonads, often yielding unfertilizable immature eggs. The artificial maturation of oocytes through NaOH-alkalinization methods was investigated in order to improve laboratory experiments on larval ecology of Patella depressa and Patella vulgata. Maturation was accelerated above pH 9.0. Alkalinizing oocytes at pH 9.5 significantly enhanced in vitro fertilization rates, especially at higher sperm concentrations. pH 10 was harmful to eggs. Sperm dilution experiments using alkalinized oocytes suggest the use of sperm concentrations proximate to 106 spermatozoa ml−1.

Climate change has strong potential to modify the structure and functioning of ecosystems, but experimental field studies into its effects are rare. On rocky shores, grazing limpets strongly affect ecosystem structure and their distribution in NW Europe is changing in response to climate change. Three limpet species co-occur in SW Britain (Patella vulgata, Patella ulyssiponensis and Patella depressa) on open rock and in pools. Shores in Ireland are similar, but currently lack P. depressa. It is anticipated that P. depressa will expand its range into Ireland as the climate warms, but we currently lack an empirical basis to predict the consequences of this change. Recent studies show that increasing abundance of P. depressa on British shores has been accompanied by a decline of P. vulgata suggesting interspecific competition. In this study, a new experimental framework was used to examine the potential for P. depressa to affect P. vulgata on Irish shores. P. vulgata was experimentally transplanted into enclosures on open rock and in pools in both Ireland and Britain. In pools, treatments also included transplanted P. ulyssiponensis to mimic natural assemblages. Growth and mortality of P. vulgata were measured over 6 months with no differences between Ireland and Britain. In Britain, P. vulgata caged in pools with transplanted P. depressa and P. ulyssiponensis showed reduced growth, compared with when caged in pools with P. ulyssiponensis alone. There was no effect of P. depressa on the growth rate of P. vulgata on open rock. Results indicate that if the range of P. depressa extends into Ireland, it would reduce the growth of P. vulgata where it co-occurs with P. ulyssiponensis in pools. The framework used here provides a field-based approach that could be used to examine the impacts of climate-induced range expansions on the structure and functioning of other ecosystems.

Summary 1On thermally stressful rocky shores, small, slow-moving ectotherms such as limpets exhibit morphological characteristics such as high-spired and heavily ridged shells which may reduce the likelihood of reaching stressful or lethal body temperatures. 2The effects of shell height and shell surface area on predicted limpet body temperatures were tested with a previously developed heat budget model. The model was parameterized with morphological data from three species (Lottia gigantea, Patella vulgata and Siphonaria gigas), which differ dramatically in their morphology and in the body temperatures they are likely to reach in the field. 3Limpet models and standard cones with higher height : length ratios lost heat to convection more readily than models with lower spired shells. 4Heavily ridged shells lost heat to convection more readily than smoother shells, but this effect was only pronounced at high wind velocities. 5When the heat budget model parameters were applied to a real environmental data set, the model predicts that maximum body temperatures and cumulative thermal stress vary among species. These differences are related primarily to the height : length ratio of the shell, and to a lesser extent to the presence of ridges. 6These results suggest that some intra- and interspecific variation in limpet morphology may be phenotypic or evolutionary responses to variation in environmental temperatures. Our findings are supported by observed patterns of limpet morphological variation across natural thermal gradients.

Highly mobile aquatic predators are known to forage in the intertidal during periods of immersion. There is limited quantitative information, however, on the extent to which these predators influence the abundance of grazing molluscs which are known to have a key role in structuring intertidal assemblages. Our preliminary video observations revealed that crabs and small fish were abundant on shores in southwest England during high-tide. We then used manipulative experiments to quantify the effect of small mobile aquatic predators on the abundance of limpets (Patella vulgata L.). On the lower shore at two moderately sheltered rocky shores three treatments were established: complete cage, partial cage (cage control) and uncaged (natural condition). The complete cages excluded all predators. The partial cage treatment allowed full access to small predators and the uncaged treatment allowed access to all predators. After two months, limpet abundance in uncaged and partial cage treatments had declined by around 50% compared to the complete cage treatment. Population structure also changed with survival of larger individuals being greater than smaller individuals in the open and partial cage treatments compared to the complete cage treatment. The effects of excluding predators were consistent at small (meters) and large spatial scales (kilometres) and hence, it would appear that the outcomes of our research are generally applicable to similar shores in the region. To explore the mechanism behind the differential effects of predators according to prey size, we compared the detachment force required to remove limpets of differing sizes from the shore. This was around four times greater for larger individuals than for smaller ones indicating that smaller limpets were more vulnerable to predation. These effects were also consistent between locations. Subsequent laboratory observations showed that the crabs Carcinus maenas (L.), Necora puber (L.) and Cancer pagurus (L.) which are locally abundant predators of limpets, had differing handling behaviour but were all highly efficient at removing limpets from substratum. Hence, shell width and attachment force appeared to be critical factors influencing the vulnerability of limpets to predation by these crabs. Limpets are known to control the abundance of macroalgae on shores in the North-east Atlantic and so our conclusions about the role of mobile predators in regulating the abundance of these grazers are important to our broader understanding of the ecology of these shores.

The characteristic variability of grazing has potential consequences for intertidal productivity and community structure, particularly as many of the underlying functional relationships are thought to be non linear. As a first approximation, it can be hypothesised that grazing is patchy over short time periods before a more uniform coverage is established over longer time scales. This prediction is supported by relatively short term observations previously made of limpet foraging. We used eight arrays of wax disks on each of four shores to test the hypothesis that grazing is patchy in the short term, but that this pattern is lost as the pattern of grazing averages out over longer time scales. Wax disk arrays were exposed for two weeks at a time for a period of six weeks in 2001 and in 2002 using the same set of disk holes each time. Grazing at the same disk location could therefore be measured over two weeks and over longer periods by averaging successive deployments. We used all three successive deployments to estimate the average grazing at each disk location over a six week period in 2001 and 2002. All six deployments were used to characterise the pattern of grazing at longer time scales. The spatial pattern of grazing in arrays was summarized using semivariogram analyses. For two-week deployments, the average standardized semivariance of grazing for disks separated by 20 cm was less than one. This pattern implies spatial autocorrelation of grazing at this scale. There was no support for the hypothesis that small scale patchiness in grazing would disappear over time. The average strength of spatial autocorrelation increased when data were integrated over longer periods. A preliminary analysis indicated that the degree of autocorrelation within arrays increased with grazing intensity at short time scales. Surface roughness disrupted autocorrelation of grazing over both short and long time scales. The persistent patchiness of grazing is likely to have implications for biofilm productivity, particularly on smoother shores. Keywords: Patella vulgata; semivariogram; spatial autocorrelation; topography; wax disk peer-reviewed

Limpets, predominantly Patella vulgata, have been associated with damaged or reced- ing canopies of Ascophyllum nodosum. Although damage results from limpet grazing, the benefits that limpets gain from this behaviour are unclear as A. nodosum is thought to be well defended from grazers by anti-herbivore compounds. In the present study, P. vulgata individuals were enclosed at densities between 80 and 320 m -2 at 2 sites within Strangford Lough, Northern Island. Limpet growth and limpet survival were compared between unsubsidised controls and treatments in which limpet diets were subsidised by fronds of A. nodosum. When subsidised, limpet residual growth rates were significantly higher and mortality was lower than in unsubsidised control treatments. Individual limpets consumed a similar amount of A. nodosum regardless of limpet density. Higher densities of limpets, therefore, consumed more A. nodosum per replicate. The effects of A. nodosum in maintain- ing limpet densities could resonate through sheltered rocky communities. The importance of a macroalgal subsidy in supporting limpet populations may have been underestimated or overlooked in earlier studies. Therefore, the extensive and productive macroalgal canopies that characterise many sheltered temperate rocky shores could be more sensitive to increased limpet abundances than previously thought.

Biodiversity is defined by the scales at which organisms coexist. Coexistence at larger spatial scales may be underpinned by segregation at smaller spatial scales or temporal variability in behaviour and/or distribution. Limpets of the genus Patella are keystone grazers controlling the growth of macro-algae on many rocky shores. This study examined the distribution and abundance of P. vulgata and P. ulyssiponensis in relation to habitat and to each other at a range of spatial scales. Limpets were sampled on four shores on each of the east and west coasts of Ireland, in two different transects (10 m apart) at each of three shore heights in winter 2003 and summer 2004. P. vulgata and P. ulyssiponensis coexist around the coast of Ireland, but on shores where they co-occur, the pattern of coexistence begins to break down at the level of shore height: P. ulyssiponensis were more abundant on the low-shore than mid-shore, and P. vulgata were distributed evenly throughout the low and the mid-shore. At the level of habitat (pools vs. open rock), the two species exhibited a high degree of segregation: P. vulgata of all sizes were more common on open rock and P. ulyssiponensis of all sizes were more common in pools. The presence or absence of P. ulyssiponensis had no effect on the distribution of juvenile P. vulgata with respect to pools. Unexpectedly, however, a greater proportion of adult P. vulgata were found in pools at sites where P. ulyssiponensis were present in the pools than at sites where they were absent. The results of this study emphasize the need for multiple-scale analyses of the distribution and abundance of organisms to understand the organisation of natural assemblages.

The occurrence of hybridisation between limpet species of the genus Patella has always been a contentious issue. Although a previous allozyme study reported high differentiation and no hybridisation between Patella vulgata Linnaeus, 1758, Patella depressa Pennant, 1777 and Patella ulyssiponensis Gmelin, 1791 along English shores, the recent finding of an mtDNA haplotype of P. depressa in a P. vulgata individual raised new doubts on this issue. To further study the possibility of hybridisation between limpet species and their level of genetic differentiation, ten allozyme loci were screened using starch gel electrophoresis for P. ulyssiponensis, P. depressa, P. vulgata and Patella rustica Linnaeus, 1758, from the Atlantic coast of the Iberian Peninsula. Our results show high differentiation between species, which could be clearly separated into different clusters with a Bayesian clustering algorithm. No significant signs of hybridisation were detected between any of the four species. Thus, the hypothesis of hybridisation between P. vulgata and P. depressa across their sympatric distribution is not supported. Two sympatric clusters were recovered within P. vulgata that could be related to Hardy-Weinberg disequilibrium found in locus MPI. Finally, due to the high level of intraspecific variability, the studied loci are interesting tools for the analysis of population structure and stock identification.

This study devised a staging system for, and monitored, the gonad development of the limpet species Patella vulgata and Patella ulyssiponensis on the South West coast of Ireland using histological techniques. Maturation began in the males of both species in January and in the females it began in March. There was no statistical difference in gonad development between sexes and between species. Spawning in the male P. vulgata occurred from September to December 2003 and in September and October 2004. In female P. vulgata spawning occurred from October to December 2003, no spawning of females was observed in 2004. In male P. ulyssiponensis spawning occurred in November and December 2003 and from September 2004 to December 2004. Spawning was observed from November 2003 to January 2004 and in September 2004 in female P. ulyssiponensis. Sex ratios also varied between the species and between months sampled. Nevertheless more males were observed in both species.

Opportunistically, the limpet Patella vulgata will switch from microphagous grazing to feeding on attached macroalgae or, a novel finding, on accumulations of stranded seaweeds. Macroalgal food is ingested primarily at night during emersion, when the thalli come to lie on the rock surface and are sufficiently damp. Monocultures of Ascophyllum nodosum are not immune to the destructive effects of limpets as previously thought. Patella vulgata can graze them down to complete disappearance, thus pushing Ascophyllum back to refuges where P. vulgata is absent or rare. Such refuges were found along a wide range from very sheltered to strongly wave-exposed rocky shores in Brittany, France. By a combination of its macrophagous and microphagous feeding habits, P. vulgata can control mid-tidal macroalgal assemblages in a similar fashion to the action of sea urchins upon kelp communities inhabiting the lower intertidal and shallow sublittoral zones of rocky shores.

Through their grazing activities limpets have an important role in controlling macroalgal abundance and as a result the structure and dynamics of rocky shore assemblages. Using two congeneric limpet species, with different biogeographic distributions, and whose ranges are expected to alter with climatic warming, we separated the magnitude of their grazing activity over time and the subsequent consequence for macroalgal growth. The northern/boreal limpet, Patella vulgata (L.), consistently grazed more than the southern/lusitanian limpet, P depressa (Pennant), particularly during spring and summer when P depressa was reproductively active. Individuals of Fucus vesiculosus (L.) that settled during this time were able to grow to a size where they escaped the grazing activities of P. depressa, resulting in mature F vesiculosus being present in all P. depressa treatments. In contrast, P vulgata, which was not reproductively active during this period, exhibited no reduction in its grazing activity and prevented macroalgae from growing in experimental treatments. It therefore appears that P. vulgata has a stronger role, than P depressa, in controlling macroalgal abundance on shores of southwest Britain. We present a conceptual model highlighting the direct and indirect interactions between these two limpet species and F. vesiculosis. If as predicted, under current warming scenarios, P depressa becomes the dominant limpet on shores of southwest Britain there will be subsequent changes in rocky shore community structure and ecosystem functioning. Our research emphasises that even closely related species with similar ecological niches can exhibit different behaviours that fundamentally alter their biological interactions with other organisms leading to idiosyncratic responses to predicted changes in climate. (C) 2007 Elsevier B.V All rights reserved.

The distribution of most species is expected to alter in response to climate change. Pre- dictions for the extent of these range shifts are frequently based on 'climate envelope' approaches, which often oversimplify species responses because many do not consider interactions between physical and biological factors. The local persistence of some species, however, is likely to be strongly modulated by microhabitat-forming organisms. Using congeneric patellid gastropods with northern/ boreal and southern/lusitanian distributions, we have demonstrated how the loss of habitat-forming macroalgal species could modify species responses to climate change. The northern limpet Patella vulgata preferentially aggregates beneath Fucus spp. When Fucus vesiculosus was experimentally removed, to simulate a decline in macroalgal abundance in response to climatic warming, P. vulgata suffered increased mortality or relocated home scars, often to nearby Fucus spp. patches. In contrast, the southern limpet P. depressa did not aggregate beneath Fucus spp. and showed no response in terms of movement or mortality to the loss of F. vesiculosus. Based on these results, we predict that the loss of Fucus spp. will influence the relative abundance of these 2 limpet species, particularly at the distributional limit of Fucus spp. In addition, differences in the aggregative behaviour of these limpet species will result in changes in the spatial distribution of grazing in the intertidal, with likely consequences for community dynamics. These outcomes could not be anticipated from predictions based on direct responses to temperature alone, highlighting the need for biotic and abiotic factors to be incorporated into predictions of species responses to climate change.

Specimens of Patella intermedia, Patella rustica, Patella ulyssiponensis, and Patella vulgata were analyzed for shell and radula characteristics. Shell growth in P. rustica and P. ulyssiponensis was basically isometric, indicating that shell shape was constant during growth. On the contrary, shell growth in P. intermedia and P. vulgata was positively allometric, indicating that as shells increased in size, the base became more circular and the cone more centred and relatively higher. Radula relative size increased in the order P. ulyssiponensis, P. vulgata, P. intermedia and P. rustica, and had negative allometric growth in all species, indicating that radula grew less as shell increased in size. Data reported in the literature estimated that the lowest risk of dislodgment for a limpet is associated with a centred apex, and a (shell height)/(shell length) or (shell height)/(shell width) ratio of ca. 0.53. However, as reported for other limpets, in all four studied Patella species, shells were more eccentric and flat than this theoretical optimum. Data reported in the literature indicated that, in limpets, decreasing the (shell base perimeter)/(shell volume) or (shell surface area)/(shell volume) ratios by increasing size results in lower soft body temperature and desiccation. In the present study, P. rustica shells displayed the lowest ratios, and P. ulyssiponensis shells, the highest. Considering that the former species lives at high shore levels, and the latter species at low levels, it appeared that shell shape in European Atlantic limpets can be directly related to resistance to desiccation and high temperature stresses. Radula relative size (in relation to shell height) also increased with increasing level in the shore, suggesting that this might be due to a decreasing abundance of algae with increasing shore level.

Germ cell commitment and meiosis initiation are among the multitude of physiological roles of retinoic acid (RA) in vertebrates. Acting via receptor-mediated transcription, RA induces the expression of meiotic factors, triggering meiosis. Contrasting with vertebrates, invertebrate RA metabolism is scarcely understood. Still, some physiological processes appear to be conserved. Here we set to evaluate the role of retinoids in the gonad maturation process of the marine gastropod Patella vulgata. We found that retinoid concentration in gonadal tissue, namely RA, varies between breeding and resting specimens, with maxima attained in the latter. Additionally, we isolated and quantified the expression of both the retinoic acid receptor (RAR) and the retinoid X receptor (RXR) in gonads. In view of the stability of retinoid receptor expression, we suggest that the balance of RA levels operates through the enzymatic control of synthetic and catabolic processes. Overall, the reported data are supportive for a developmental role of RA during gonadal maturation in P. vulgata, which should be addressed in other protostome lineages.

Grazing mollusks are used as a food resource worldwide, and limpets are harvested commercially for both local consumption and export in several countries. This study describes a field experiment to assess the effects of simulated human exploitation of limpets Patella vulgata on their population ecology in terms of protandry (age‐related sex change from male to female), growth, recruitment, migration, and density regulation. Limpet populations at two locations in southwest England were artificially exploited by systematic removal of the largest individuals for 18 months in plots assigned to three treatments at each site: no (control), low, and high exploitation. The shell size at sex change (L 50: the size at which there is a 50:50 sex ratio) decreased in response to the exploitation treatments, as did the mean shell size of sexual stages. Size‐dependent sex change was indicated by L 50 occurring at smaller sizes in treatments than controls, suggesting an earlier switch to females. Mean shell size of P. vulgata neuters changed little under different levels of exploitation, while males and females both decreased markedly in size with exploitation. No differences were detected in the relative abundances of sexual stages, indicating some compensation for the removal of the bigger individuals via recruitment and sex change as no migratory patterns were detected between treatments. At the end of the experiment, 0–15 mm recruits were more abundant at one of the locations but no differences were detected between treatments. We conclude that sex change in P. vulgata can be induced at smaller sizes by reductions in density of the largest individuals reducing interage class competition. Knowledge of sex‐change adaptation in exploited limpet populations should underpin strategies to counteract population decline and improve rocky shore conservation and resource management.

Along with habitat loss and climate change, anthropogenic inputs to the biosphere rep- resent major threats to biodiversity; unfortunately, many of the studies necessary to understand these potential impacts have been carried out in laboratory systems with little or no ecological relevance. Understanding the impact of contaminants is dependent on the knowledge of their ecological conse- quences in the environment rather than reliance on laboratory studies. We assessed the impact of pulse contamination of copper on the behaviour and physiology of the limpet Patella vulgata L. on rocky shores. Toxicological methods were used in conjunction with ecological methods to elucidate possible effects of copper on cellular viability and phagocytosis (measures of cellular and immunolog- ical health), tenacity and grazing behaviour of P. vulgata. Exposures lasted for 10 d, and were repli- cated on 2 shores, and at 2 times. Ecological effects were assessed by the deployment of wax discs to quantify grazing intensity of contaminated limpets during this period. We also tested the effects of copper on tenacity as a measure of vulnerability to predators or to wave action. Phagocytic activity was unaffected by the addition of copper, as was the grazing intensity of P. vulgata. Tenacity was significantly reduced due to the copper pulse exposure, as was cellular viability. Copper significantly affected cellular viability; this was also reduced, although to a lesser extent, in limpets sampled from control plaster block replicates, suggesting unidentified influences from the plaster itself. Copper levels were significantly elevated in limpets sampled from the copper treatment, suggesting this as a useful method of toxicant dosing in the field. Ecological effects were mixed. There was no change in grazing intensity, but a significant reduction in tenacity, with implications for limpet mortality and thus rocky shore ecosystem processes.

The specimens of Patella intermedia, Patella rustica, Patella ulyssiponensis and Patella vulgata were analysed for morphological and morphometric characters, and for the resistance to compression and crushing to a force applied at the apex. Shell shape in these species ranged from the high symmetrical cone, with a rounded base of P. rustica, to the flat, asymmetrical, narrow-pear-shaped base of P. ulyssiponensis. P. intermedia and P. vulgata showed intermediate morphologies. Shell thickness increased linearly with age, but differed in the four limpets. P. rustica had the thickest shells, and P. ulyssiponensis and P. vulgata had the thinnest shells. P. intermedia displayed intermediate shell thickness. Considering deformability and toughness, P. intermedia shells usually needed the highest force to compress in height, the highest pressure to collapse, and were appreciably deformed at collapse. On the opposite side, P. ulyssiponensis shells usually needed the lowest force to compress in height, the lowest pressure to collapse, and were much less deformed at collapse. P. intermedia shells were therefore the most deformable and tough, and P. ulyssiponensis, the most stiff and fragile. P. rustica and P. vulgata shells displayed intermediate behaviour. However, numerical simulations based on the finite element method using the experimentally determined shells’ geometry and thickness, but considering similar shells’ material and structure in the four species, predicted that shell toughness should be decreased in the order P. rustica >> P. intermedia > P. vulgata >> P. ulyssiponensis. P. rustica shells’ geometry (a high and centred cone) and thickness (very thick) were therefore, theoretically, the most fitted for shells to resist crushing by compression. Yet, in the experimental tests, they were not the most resistant shells. It was concluded that resistance to crushing was not a direct function of shell morphology and morphometry, but appeared to be mainly determined by shell deformability. This is most probably related to differences in the internal composition and architecture of the shell in the four species. By comparison with data reported in the literature on the pressures normally exerted by ocean waves, it was concluded that these limpets have resistances to crushing in far excess to pressures normally endured in nature, being therefore unlikely that these species are crushed by the action of even very strong waves on shores. Hard objects, like logs and boulders, can be hurled onto the shore by waves, and constitute a much greater threat to limpets than the waves themselves. The high toughness of the limpet shell can be related to resistance to the impact of incidental hard objects. There was no direct relationship between the habit preferences of these limpets and the resistance to crushing. Other factors are involved in the distribution of these species in the shores.

Laboratory experiments were undertaken to examine fertilization success in the intertidal prosobranch limpets Patella ulyssiponensis and Patella vulgata. Alkalization of eggs (10 min in pH 9.0 sea water) prior to fertilization trials improved fertilization rates greatly. Fertilization success was found to be a function of sperm concentration, gamete age and contact time. Sperm concentration needed for optimum fertilization success in vivo ranged between 105 and 107 sperm ml−1 for both species although at higher concentrations the number of normally developing trochophore larvae decreased. For P. vulgata, sperm longevity (at a concentration of 106 sperm ml−1) did not exceed 6 h, whereas eggs of both species were fertilizable for up to 12 h. Maximum fertilization success occurred after 15–30 min gamete contact time. The Vogel et al. (Math Biosci 58:189–216, 1982) fertilization kinetics model is developed to allow for non-complete fertilizations under optimal sperm concentrations, and a new parameter fitting technique is developed to improve estimates of fertilization success for short gamete contact times.

Canopy-forming macroalgae are key species on temperate rocky shores. However, there is a lack of understanding of how the relative balance of physical and biological factors controls the establishment and persistence of intertidal macroalgae. Here we present an integrated study of the relative importance of wave-induced forces and grazing for the recruitment and survival of the canopy-forming intertidal macroalgae Fucus vesiculosus and F. spiralis. A set of overtopped breakwaters provided a nearly unconfounded gradient in wave exposure between seaward and landward sides. A biomechanical analysis was performed based on empirical measurements of maximum drag forces in breaking waves, a model of long-term maximum wave height, and the breaking stress of Fucus spp. The estimated maximum flow speed (7-8 m/s) on the seaward side of the breakwaters was predicted to completely dislodge or prune Fucus spp. larger than approximately 10 cm, while dislodgment was highly unlikely on the landward side for all sizes. Experimental transplantation of Fucus spp. supported the biomechanical analysis but also suggested that mechanical abrasion may further limit survival in wave-exposed locations. Experimental removal of the limpet Patella vulgata, which was the principal grazer at this site, resulted in recruitment of Fucus spp. on the seaward side. We present a model of limpet grazing that indicates that limpet densities5-20 individuals/m2 provide a proximate mechanism preventing establishment of Fucus spp., whereas wave action2 m/s reduces persistence through dislodgment and battering. In a conceptual model we further propose that recruitment and survival of juvenile Fucus spp. are controlled indirectly by wave exposure through higher limpet densities at exposed locations. This model predicts that climate change, and in particular an increased frequency of storm events in the northeast Atlantic, will restrict fucoids to more sheltered locations.

Planktonic cyprid larvae of the intertidal barnacle, Semibalanus balanoides (L.) and the infralittoral/sublittoral Balanus crenatus Brug., are readily captured in small, chambered, passive traps deployed on rocky shores. Several structural developments of a previous trap design improved urea (killing solution) retention and capture of cyprids of both species. The design adopted for assessment of the larval supply/settlement relationship of the focal species, S . balanoides , included additional internal baffles and the replacement of the cylindrical trap opening with a coned aperture. Three sizes of cone aperture area were compared, with the intermediate size (1 cm 2 ) being optimal for measuring larval supply over the full range of wave action on local shores. Most daily catches included larvae that were recognizable as having been dead before they were trapped, and had recently passed through the guts of grazing limpets ( Patella vulgata L.). One site showed that ∼32% of trapped S . balanoides were so-called ‘faecal’ cyprids. An inability to distinguish these in trap samples would lead to a significant over-estimation of larval supply. Preliminary data indicated that S . balanoides larval supply to the intertidal is enhanced by onshore winds (or perhaps increased wave turbulence), whereas B . crenatus supply was positively correlated with offshore winds (or decreased turbulence). A clear ‘supply/settlement’ relationship for S . balanoides at six sites along ∼40 km of coastline was derived from 1 cm 2 coned trap catches and daily counts of settlement on ceramic tiles. There were no instances of disproportionately high settlement in relation to larval supply, but at five sites there were days when settlement apparently failed. The identification and explanation of settlement success and failure is fundamental to understanding the importance of larval supply in structuring marine benthic populations and communities.

Patella vulgata is a boreal cold temperate species and is the dominant limpet in northern Europe. Few works have focussed on the population genetics of this species. Therefore, the aim of this work was to assess the degree of genetic and morphological differentiation of P. vulgata on a macroscale by using 20 allozyme loci and 6 morphological variables. Samples were taken from the following locations: Dingle Peninsula (Southwest Ireland), Port Erin (Southwest Isle of Man), St. Bees Head (north Cumbria, England), St. Agnes Head (north Cornwall, England), Cellar Beach (south Devon, England), Whitley Bay (north Newcastle-Upon-Tyne, England), Sines (Portugal), and Pointe de Chanchardon, La Rochelle (Bay of Biscay, France). Morphological variables were analysed by the multivariate Canonical discriminant analysis. Genetic variation was assessed by diversity measures such as polymorphism and heterozygosity; genetic subdivision of P. vulgata population was determined by the estimator θ of F ST, and the genetic similarity between populations was measured by Nei’s genetic identity. No significant morphological differentiation was observed among samples. Moderate genetic population subdivision was observed (θ = 0.137±0.074) despite great geographic distances. The minimum genetic identity observed was between Ireland and France (I = 0.942) and maximum was observed between Portugal and north-east England (I=0.998). Two main groups were shown by UPGMA cluster analysis (I = 0.965). One formed by Irish, Manx, north Cumbria, and curiously, south Devon samples, while the second includes Portuguese, French, north-Newcastle-upon-thyne, and north Cornwall samples. No association (g = 0.956; p>0.050) was found between pair-wise genetic divergence and geographic distance separating subpopulations, mainly due to an unexpected pattern of genetic heterogeneity found in Southwest England.

Conventionally, direct interactions between species are considered to be the most important biological factors determining community composition, structure and stability. However, it has been suggested that the indirect interactions occurring between species may be as important. One area of ecology where the direct effects of one species on another have been well studied is in the rocky intertidal. Examination of the effect of the presence of P. vulgata (limpets) and L. littorea (periwinkles) on the settlement and development of S. balanoides (cyprids/barnacles), over a cyprid settlement season and some six months later, in four different treatments (limpets only, limpets and periwinkles combined, periwinkles only and control (no animals)) revealed the following: (1) that the presence of limpets increased cyprid settlement and recruitment success above treatments containing no limpets; (2) that cyprid settlement and success were greatest on the limpets-only treatment, followed by the limpets-and-periwinkles treatment, then by the control treatment and then by the periwinkles-only treatment; (3) that the initial effects observed in the treatments were reflected in the long-term community structure; (4) that the effects of the treatments were independent of variations in algal biomass between treatments, i.e. the effects were not indirectly mediated through a second species (host); (5) that cyprid mortality was greatest on the periwinkles-only treatment; (6) that the source of the effect of limpets on cyprid settlement appeared to originate indirectly through the action of their residual pedal mucus trails. It is concluded that periwinkles can affect the settlement and success of barnacles directly through biological disturbance (i.e. surface ablation). However, although limpets may have a direct negative effect on barnacle settlement and success, at low to medium densities, limpets can positively indirectly influence the cyprid settlement and success. This effect operates at a factor greater than that afforded by the direct negative effects of periwinkles in a mixed-species treatment. These results illustrate how the indirect effects of one species on another can have a more important structuring effect than those derived from direct effects alone.

By the end of the 21st century models of climate change predict that the air temperature over most of the British Isles will increase by between 2 and 3 °C and sea-level will rise by 40–50 cm. Over that period it will become windier and mean wave height will increase, as will the frequency of storms. These changes in climate and weather will impact the intertidal zone of the UK and will cause distribution changes in many of the common invertebrate species that live there. Where these changes are severe they may well impact on patterns of distribution of ducks and wading birds. In the British Isles a number of organisms live close to their geographical limits of distribution. Some of these species might be expected to extend their range as climatic restraints are relaxed. Species currently limited by cool summers or winter cold will move northwards. In most cases the effects on the distribution of waterbirds will be small. For example, the replacement of the Northern Limpet Patella vulgata by the Southern Limpet P. depressa is unlikely to adversely affect Eurasian Oystercatchers Haematopus ostralegus. Of wider concern is the possibility that as climate warms the abundance and productivity of brown algae will decrease. This is likely to have two significant effects for waders. First, it would represent a loss of potentially rich feeding grounds for species such as Ruddy Turnstone Arenaria interpres that feed on small easily desiccated invertebrates living on or below the seaweed. Secondly, as algae die or are broken away the resulting debris is exported to sediment habitats where it considerably boosts the in situ production of bacteria at the base of the food web. An increase in sea-level will only have a major impact on the extent of rocky shore invertebrate communities where shore topography prevents the upward migration of the biota. Where a seawall limits shores, for example, biological production will be curtailed as the area available for colonization decreases. Increases in the size of waves and the frequency of storms will mimic increasing exposure and there will be a significant reduction in algal production in areas that are affected.