Nearest neighbour analysis of spatial dispersion of intertidal prosobranch gastropods within two substrata

Summary The patterns of dispersion of Nerita atramentosa, Bembicium nanum and Cellana tramoserica within two major substrata, the encrusting alga Peyssonelia gunniana and bare rock were analysed from quadrat samples taken at low tide, over 1 year. Distances to the nearest members of each species from random points were analysed to determine spatial patterns of dispersion. The frequencies of nearest neighbours of each species to each animal in the sampled area were analysed to determine the degree of spatial segregation and symmetry in the three species. Nerita atramentosa and Bembicium nanum showed significant aggregated (clumped) dispersion in both substrata. Cellana tramoserica, however, was dispersed at random on Peyssonelia, and regularly dispersed on bare rock. There was no correlation between index of dispersion and density, or time of year, in any species. No useful tests for segregation and symmetry between Cellana and the other two species could be made because of the difference in dispersion of this species. Nerita and Bembicium were significantly segregated on both substrata, indicating that the microhabitat requirements of the two species are different. The two species were symmetrical on both substrata, indicating that there is no tendency for members of one species to be more isolated than members of the other species.Possible explanations of the differences in spatial pattern shown by the three species are discussed, and experimentally testable hypotheses are proposed to clarify the importance of physical and biological factors in the spatial patterns of distribution of the three species.     

Movements of intertidal gastropods

Movements of marked individuals of Nerita atramentosa Reeve, Austrocochlea constricta (Lamarck), Bembicium nanum (Lamarck), and Cellana tramoserica (Sowerby) were recorded at various sites in New South Wales in 1974 and 1975. A model of random movements during feeding at high tide was proposed, i.e., that animals wandered in random directions and the distances moved were exponentially distributed. Appropriate statistical tests for departures from such a model are given. The types of departure tested were non-random directions, non-exponentially distributed dis- tances, different distances in different directions, and correlated distances or directions of movement between days. With two exceptions, the movements of the animals over short periods were adequa- tely described by the assumption of random movements. One exception was A. constricta in one locality where movements of the sample were limited by it being in a large rock-pool and the other was the limpet C. tramoserica which displayed homing behaviour, so that the population could be divided into homers and non-homers. Movements of non-homing limpets were random.The mean distance moved by N. atramentosa was not significantly different from that of A. con- stricta. The mean distances moved by these two species were greater than those of B. nanum and the non-homing C. tramoserica, which were not different from each other. Adult and one-year old N. atramentosa did not differ significantly in mean distance moved, but both moved more than juveni- les. These patterns of movement are discussed with respect to hypotheses about feeding in these species.The model of movement was used to predict the effect of different percentage cover in small pools on the movement of A. constricta, which shows a preference for this micro-habitat. Increased availability of small pools reduced the mean distance moved by this species.     

Field experiments with “cageless” methods to manipulate grazing gastropods on intertidal rocky shores

Excluding particular species or guilds of grazing gastropods from experimental areas on intertidal rocky shores is a manipulation frequently used by marine ecologists. Manual removals, cages or fences are classical methods, but they usually require large areas or multiple procedural controls to avoid experimental artefacts. In order to minimise some of those constraints, alternative methods have been developed and are increasingly being used. The effects and limitations of these methods are, however, generally assumed and hardly ever tested. The experiment described here was designed to compare the effectiveness of manual removals, copper paint and sticky barriers in excluding intertidal grazing gastropods from experimental areas on a natural sandstone platform in South-eastern Australia. The response of the grazing assemblage was evaluated for individual species and collectively (integrating all species) by direct counts of individuals and from radular marks on wax discs. The observed patterns varied considerably among species. As predicted, barriers of copper paint effectively excluded the limpet Cellana tramoserica, for several weeks, while having no apparent effect on co-occurring species of grazing gastropods. Sticky barriers were also effective in excluding C. tramoserica, but for shorter periods of time. Contrary to expectations, however, sticky barriers were generally incapable of deterring the coiled snails Bembicium nanum and Nerita atramentosa. Barriers of copper paint are, therefore, a cost-effective way to manipulate diversity (densities of individuals, number and composition of species) and function (intensity of grazing) in this type of assemblage. Care must be taken to clarify the underlying mechanisms and to identify adequate procedural controls.     

Competition between Cellana tramoserica (Sowerby) (gastropoda) and Patiriella exigua (Lamarck) (asteroidea), and their influence on algal standing stocks

Caging experiments were undertaken in the field to test if competition occurs within and between the limpet Cellana tramoserica (Sowerby) and the starfish Patiriella exigua (Lamarck). Both species suffer from intraspecific competition, even at natural densities, and increased density reduces body weight and survival in Cellana tramoserica and growth rate in Patiriella exigua. Cellana tramoserica depresses the growth of Patiriella exigua, while the starfish unexpectedly seems beneficial to Cellana tramoserica, increasing its body weight. The difference in these effects may be due to the divergent methods of feeding used by the two species. C. tramoserica digs into the substratum and removes most available microflora, while Patiriella exigua everts its stomach onto the rock face and can only remove superficial or loosely attached microflora and detritus, and leaves a mucous web which may supplement the diet of Cellana tramoserica. Even at low densities C. tramoserica can completely prevent macroalgae from developing, while Patiriella exigua is unable to do so, although it shows the rate of development. Macroalgae grew in all the cages which contained only P. exigua, but in spite of this the starfish still suffered from intraspecific competition, probably because they cannot feed on macroalgae. Cellana tramoserica also competes by interference, and on contact with Patiriella exigua it extends its mantle and pallial tentacles, both of which may release mucus. P. exigua retreats from Cellana tramoserica, and those parts of its body that have been touched by the limpet may temporarily be immobilised and become wrinkled up. It is suggested that this interference behaviour by C. tramoserica is derived from a similar but more intense reaction that it has to invertebrate predators. Patiriella exigua lacks a pelagic larva and does not have a refuge outside the range of Cellana tramoserica. Other factors must thus be sought to explain the continued coexistence of Patiriella exigua with Cellana tramoserica. It is suggested that although C. tramoserica has a competitive effect on Patiriella exigua, it (and other grazers) may be necessary for P. exigua, preventing macroalgal growth and maintaining a surface suitable for P. exigua to feed on.     

Effects of body size and water temperature on grazing rates of four intertidal gastropods

Abstract Rates of radular activity were measured in four species of Australian gastropods. The species were Austrocochlea constricta, Bembicium nanum, Cellana tramoserica, and Nerita atramentosa. Work was done in Botany Bay at Cape Banks, NSW, Australia from September 1989 to June 1990. A hydrophone was used to record radular raspings of snails in situ. Averages based on all observations showed Cellana had the fastest rate with 80 rasps/min, and was followed by Austrocochlea (71 rasps/min), Bembicium (57), and Nerita (39). Faster rates were associated with warmer water temperature and smaller body sizes in all species except Bembicium. The potential for these differences in radular activity affecting competitive interactions is discussed.     

Analysis of patterns of dispersion of intertidal prosobranch gastropods in relation to macroalgae and rock-pools

Summary The patterns of dispersion of Nerita atramentosa, Austrocochlea constricta, Bembicium nanum and Cellana tramoserica are analysed from quadrat samples taken between June 1972 and June 1973. In each quadrat, the proportion of animals in each of two substrata, the encrusting alga Peyssonelia gunniana and small pools, was recorded, with the proportional area of cover of each substratum, height on the shore, density of the species and the time of sampling. Multiple regression and covariance analysis showed that, in all cases, there was a linear relationship between proportion in a substratum and proportional area of substratum. No other variables were significant.Juvenile Nerita and Austrocochlea showed preferences for both substrata. Adult Nerita and all Bembicium were dispersed at random with respect to both substrata. Adult Austrocochlea were randomly dispersed with respect to Peyssonelia, but showed a preference for water. Cellana showed a preference for water and against Peyssonelia. These microhabitat preferences are discussed in relation to previously described effects of substrata on the densities of these four species.     

Vertical and seasonal patterns in competition for microalgae between intertidal gastropods

Grazing by the snail Nerita atramentosa and the limpet Cellana tramoserica caused similarly great reductions in abundance of microalgae — measured by direct counts and by estimation of chlorophyll analyses. A smaller snail, Bembicium nanum, caused smaller reductions of microalgal resources, compared with ungrazed areas. These results were consistent with the competitive abilities of these three species. Chlorophyll concentrations in samples of grazed rocks were reliable estimates of the nature and abundance of food available to the grazers.Inter- and intra-specific competition amongst Nerita and Cellana were investigated at various densities in experimental cages. To examine the effects of different availability of food resources, the experiments were repeated at three heights on the shore (abundance of food decreases with height) and during autumn/winter and spring/summer periods of the year (less food is available during summer).Density-dependent mortality of Cellana was caused by the presence of other limpets, or of Nerita. Mortality was greatest at higher levels and during the spring/summer and was significantly, negatively correlated with mean chlorophyll concentration in the experimental cages. The only exception was that all limpets suddenly died in cages at the highest level during the summer period, which cannot be explained solely by competitive interactions. Nerita showed no density-dependent mortality during the short periods of these experiments. Tissue-weights of Nerita declined with increased density, but the effect of Cellana was not as great as the intraspecific effect of Nerita. Snails retained weight better at lower than at higher levels, and during the autumn/winter which is consistent with the availability of food. Tissue weights of both Nerita and Cellana were positively correlated with chlorophyll concentrations inside the cages in both seasons investigated.These experimental results demonstrate that intensity of competition will vary from place to place and time to time according to the densities and mixtures of the grazers, and according to the availability of microalgal food.     

Intertidal zonation of prosobranch gastropods: Analysis of densities of four co-existing species

The densities of populations of Nerita atramentosa Reeve, Austrocochlea constricta Lamarck, Bembiciuin nanum (Lamarck), and Cellana tramoserica (Sowerby) were recorded in 0.25 m² quadrats on landward and seaward halves of a sheltered and a moderately exposed rock platform from June 1972 to June 1973. The data have been subjected to variance and multiple regression analysis. Differences between densities of each species on the two shores and in the two areas of each shore are discussed with respect to the regressions on littoral height. Densities of Nerita on the sheltered shore, Austrocochlea and Bembicium on both shores, and Cellana, on the exposed shore, increased with height. The density of Cellana on the sheltered shore decreased as height increased. The density differences in the landward and seaward areas on the two shores were attributable to the effect of height for Nerita and Austrocochlea. Height was the major variable affecting the density of these two species on the sheltered shore. Bembicium and Cellana densities on both shores, and Austrocochlea density on the exposed shore, are not primarily determined by height.On the sheltered shore, Nerita and Cellana increased in density with increasing cover of water in rock pools. On the exposed shore, Bembicium and Cellana became sparser with increasing water cover. Bembicium on both shores increased in density with increasing cover of the alga Peyssonelia, as did Cellana on the exposed shore. Cellana and Austrocochlea tended to increase in density with Nerita on the sheltered shore. Cellana tended to be sparse where Bembicium was dense on the exposed shore. The biological effects of substrata and other species on the densities of each species remain unexplained.Significant time effects were found for variations in the densities of Nerita and Cellana on the sheltered shore and of Austrocochlea and Cellana on the exposed shore. The seasonal change in density of Nerita was parabolic with a minimum in summer, and is consistent with the known period of recruitment of this species. The lack of significant time effects on density of Austrocochlea on the sheltered shore is consistent with the known continuous recruitment of this species. The declining density of Austrocochlea on the moderately exposed shore was found, by size-frequency analysis, to be due to reduced recruitment compared with surrounding shores. The density of Cellana on both shores showed seasonal changes, with a maximum in summer, which is consistent with the known period of recruitment in Cellana. The lack of seasonal changes in density of Bembicium is briefly discussed.This type of multifactor analysis identifies biological interactions affecting variation in density of these species. It also corroborates the results of other ecological investigations and determines the relative importance of a variety of physical and biological factors in the distribution of density of intertidal gastropods.     

Phenotypic flexibility in the metabolic response of the limpet Cellana tramoserica to thermally different microhabitats

Temperature determines all physiological responses by limiting cellular reaction rates. Daily temperature variation differs between microhabitats, which means that subpopulations of the same species may respond differently to temperature. The aim of this study is to determine how physiological responses to temperature of the limpet Cellana tramoserica differ between limpets from variable and from stable thermal environments. Oxygen consumption and anaerobic and aerobic metabolic capacities were measured over a range of temperatures in limpets from thermally stable and variable field sites in summer and winter, and in laboratory acclimation treatments. Limpets from both variable and stable sites, showed acclimatisation of anaerobic and aerobic potentials. Limpets from stable environments, but not from variable environments, showed increased oxygen consumption in winter. Comparison of field and laboratory data showed that temperature was the signal for acclimatisation. The physiological response of C. tramoserica to temperature depends on season and microhabitat. Care must therefore be taken when conducting interspecies comparisons of response to temperature to address the confounding effects of phenotypic plasticity. Differences in physiological response to temperature in phenotypically flexible species like C. tramoserica may simply reflect individual reactions to immediate environmental conditions.     

Patterns of adult abundance in Chthamalus stellatus (Poli) and C. montagui Southward (Crustacea: Cirripedia) emerge during late recruitment

The aim of this study was to investigate when adult distribution patterns are established in the barnacles Chthamalus stellatus and C. montagui. Adult ‘zones’ were identified by analysing field counts of both species at mid and upper shore heights. Monthly collections of cyprids, < 1 month old metamorphs and recruits (all metamorphosed individuals older than approximately 1 month) were made for C. stellatus and C. montagui in natural barnacle beds at six shores in SW Ireland. This was carried out over one year in 1996/1997, using a hierarchical sampling design. Abundance of total recruits (0-3 months old) was compared between adult zones after the main settlement season had ended. In addition, scales of variability in 0-3 month recruitment into adult zones were compared between the species at two scales: shores (1000s of metres) and sites within shores (10s of metres). Older recruits of each species, up to 11 months of age, were also compared between adult zones.The majority of settlement (measured as attached cyprids) occurred between August and October 1996. In October, there was no effect of adult zone on the abundance of total (0-3 month) recruitment up to that point in either species. Despite this homogeneity in recruitment between adult zones, significant spatial variation was found in 0-3 month recruits of both species at both of the scales examined. In C. stellatus the amount of variation associated with the larger scale (shore) was more than twice that of sites or of the residual variation (replicates within sites). 0-3 month recruitment in C. montagui was also most variable at the scale of shores but the residual variability (between replicates within site) was of similar magnitude to that of shores. Variability in 0-3 month C. montagui recruitment was relatively low at the scale of sites.There was a small but consistent input of recruits to adult zones over 9 months of the year, complicating the assessment of when adult patterns were set-up in these species. By June 1997, characteristic patterns of adult dominance had been established at all shores. Settlement had completely ceased by this time and individual barnacles were potentially 11 months old. Neither settlement nor early recruitment are significant in determining adult zonation patterns in these species. Instead, differential mortality patterns in individuals up to the age of 11 months are implicated in determining patterns of distribution of both species.     

Ecology of the intertidal snail Littorina acutispira Smith

Littorina acutispira Smith, a minute gastropod of < 3 mm shell height, lives at great densities in pools and on rock-surfaces at the highest levels of sea-shores in New South Wales. Populations from pools and dry areas were sampled on two shores for 18 months to investigate seasonal changes in density, size-structure, rates of growth and reproductive biology of the snails. Densities of snails increased between February and May, due to an influx of juveniles, and then declined until the following February, when they increased again. The decrease in density was due to the death of the largest snails at the end of summer, and the mortality of medium-sized snails between June and January. Longevity was estimated as 1–2 yr, but most individuals died by ≈ 16 months from settlement on the shore. Newly-settled snails grew to merge in size with those of the previous year's population by winter. During the summer months, the rate of growth of snails from a sheltered shore was greater than that of snails on a shore exposed to wave-action. Laboratory experiments revealed that this could be attributed to the presence of better quality food, or food in greater abundance on the sheltered shore, compared with the exposed shore.During winter months, but not during the summer, snails from dry areas grew more slowly than those from pools. An experiment demonstrated that some snails from dry areas might be able to compensate for reduced periods of feeding by being able to feed faster when submersed. This could not explain the differences in natural rates of growth.L. acutispira bred from October–November to March–April. Spawning in the laboratory was greatest during late summer (January to March). The percentage of mature oocytes in the gonads was small in winter and increased in early summer. Among the largest-sized snails, females outnumbered males. Two experiments, on unsexed and pre-sexed snails, demonstrated that the biased sex-ratio of the largest snails was due to faster growth by females.There was a greater density of snails on the exposed shore, which was correlated with the presence of barnacles. When barnacles were removed from experimental areas, the density of the snails declined within 24 h. This suggested that barnacles provided a refuge from wave-shock, rather than shelter from desiccation or high temperature. In laboratory experiments, snails were exposed to higher temperatures and less humidity than they would normally encounter on the shore. There was negligible mortality of small or large snails after 24 h of these conditions.This minute species grows quickly, recruits annually and has a short life-span. This type of life-history is discussed in comparison with similar small species from other habitats.     

Breeding and recruitment in a population of the New Zealand starfish Stichaster australis (Verrill)

The annual reproductive cycle of the New Zealand starfish Stichaster australis (Verrill) was determined at Maori Bay on the west coast of Auckland. S. australis has a clearly defined summer breeding season, closely repeated from year to year. Changes in the gonad shown by histological sections confirmed this cycle.S. australis juveniles reared in the laboratory were maintained on Mesophyllum insigne (Foslie) Adey substrata and the growth rates and feeding behaviour of individual starfish were determined. Settlement on the shore was from May to July each year. When the time of spawning is considered this implies a planktotrophic larval life of ≈ 6 months. This is considerably longer than laboratory studies on larval development would suggest.Growth of yearly settlement cohorts on a nursery site on the shore used in conjunction with laboratory results gave a fairly clear picture of growth after settlement. Growth rates followed a typical sigmoid curve, growth being slow initially and becoming more rapid later. The numbers of juveniles recruited to nursery areas vary from year to year but mortality following settlement appears to be low. It was found that juvenile Stichaster australis graze Mesophyllum insigne exclusively until they reach ≈ 0.8 cm in diameter (7–8 months old) when they may occasionally predate juvenile Perna canaliculus (Gmelin). The incidence of carnivorous feeding gradually increases until juveniles are ≈ 2.0–2.5 cm in diameter (15–18 months old) by which time they are exclusively carnivorous on small P. canaliculus. As growth continues juvenile starfish gradually migrate from nursery areas to adjacent reefs where there are dense beds of adult P. canaliculus. Starfish of this species become sexually mature when they reach ≈ 5–8 cm in diameter.     

Interactions of Patella and macroalgae with settling Semibalanus balanoides (L.)

Biological interactions affecting densities of settling and newly-settled Semibalanus balanoides (L.) have been investigated by manipulative field experiments on the Isle of Man.The effects of sweeping by fucoid clumps of different species and Patella browsing have been compared on moderately-exposed shores. Patella allowed barnacle settlement by preventing growth of competing green algae, but reduced post-settlement densities. Small clumps of Fucus spiralis L., F. vesiculosus L, and F. serratus L. all reduced settlement considerably more so than limpets. F. serratus had the greatest sweeping effect.Interactions between macroalgae and Semibalanus balanoides have been investigated at all levels on sheltered shores and low down on more exposed shores. In the Fucus spiralis and F. vesiculosus zones, post-settlement numbers were higher than in adjacent areas where the canopy was removed. Barnacles did not settle readily in the Ascophyllum zone in either experimental or control areas. Settlement occurred in the upper part of the Fucus serratus zone in experimental areas where the canopy was removed but not in control areas. No settlement occurred in either treatment or control areas lower in the F. serratus zone. At all levels on the shore fucoid canopies seemed to reduce cyprid settlement, but the effect was greatest amongst F. serratus where there was total prevention. High on the shore the effect of enhanced post-settlement survival under the canopy outweighs reduction of cyprid settlement thus there are greater numbers in the controls. Competition with red algal turfs was shown to set the lower limit of the barnacle zone on a vertical pier face.     

Variability in the reproductive effort of the limpet,Cellana tramoserica

Summary The reproductive effort of the limpet, Cellana tramoserica was investigated in high, mid, and low intertidal areas of the rocky shore as well as in a subtidal population. The reproductive effort of individuals within a population was independent of age. There was, however, a noticeable difference among populations with respect to the relative amount of gonad material spawned per year. The individuals from the subtidal population had the greatest reproductive effort, spawning the equivalent of 101% of their somatic weight per year, while the low, mid and high intertidal populations spawned the equivalent of 66%, 20%, 51%, respectively.Experimental transplants showed that individuals from all three intertidal populations increased their reproductive effort to match that of subtidal individuals. It was unlikely, therefore, that the observed patterns were the result of genetic differences among the populations. The reproductive effort of the populations was also not correlated with the rate of mortality of the adults in the four populations. The observed differences could be explained by a combination of the availability of food and intraspecific competition. The flexibility in the reproductive pattern of Cellana is hypothesised to be a result of this species having a dispersive larval stage, which may result in selection for plasticity of reproductive allocation, depending upon the environmental conditions at the site of settlement and growth to adult size.     

Effects of prey density,prey size and predator size on rates of feeding by an intertidal predatory gastropod Morula marginalba Blainville (Muricidae), on several species of prey

As a prerequisite for models of foraging behaviour of the whelk, Morula marginalba Blainville (Muricidae), the effects of variation in density of prey on the rate of feeding of the predator were examined in field conditions for three coexisting species of prey. Densities of prey used were those at which the prey, two limpets and a barnacle, occurred naturally in the rocky intertidal habitat.Large limpets, Cellana tramoserica (Sowerby) can resist attacks by predatory gastropods by raising the mantle over the outside of the shell. These experiments showed that no C. tramoserica were killed by Morula marginalba even at very great densities and with no alternative prey present. For the small limpet Patelloida latistrigata (Angas), one of the whelk's most highly preferred prey, juveniles were eaten 1.4 times as fast as adults. Fitting the random predator equation gave greater attack coefficients and shorter handling times for juvenile than adult limpets.Sizes of both predator and prey affected rates of eating barnacles, Tesseropora rosea (Krauss), but not in a simple way. Whelks of 15-mm aperture length ate adult barnacles 4.2 times faster than did 12-mm whelks, but there was no significant difference in the rates at which the two sizes of snail ate juvenile barnacles.Rates of feeding on T. rosea and Patelloida latistrigata increased significantly with prey density. These results form a basis for including the density of prey in models of spatial dispersion of the predatory gastropod Morula marginalba.     

Measure of stress response induced by temperature and salinity changes on hatched larvae of three marine gastropod species

To better understand the cascade of molecular reactions leading to delayed development and mortality of early life stages of marine intertidal gastropods, in response to temperature and salinity changes associated with climate change, three biomarkers: total antioxidant capacity, lipid peroxidation and lysosomal stability were investigated on hatched larvae. Encapsulated embryos of three marine gastropod species (Bembicium nanum, Siphonaria denticulata and Dolabrifera brazieri), which have already proven responsive to thermal and osmotic variations, were exposed to six combinations of temperature (22 °C and 30 °C) and salinity (25‰, 35‰ and 45‰) until the larvae hatched. Time to hatching was affected by salinity and temperature in all three species. High salinity (45‰) generally retarded the hatching process although the response was species-specific for temperature. Total antioxidant capacity and lipid peroxidation were also highly species-specific with the general trend showing that these biomarkers were adversely affected by high temperature (30 °C) at salinities of 25‰ and 45‰. Bembicium nanum lysosomal destabilisation increased significantly with an increase in temperature and salinity (30 °C and 45‰) and this was associated with delayed development and increased mortality. Investigations on the additional biomarker, lysosomal stability, gave a clearer picture of the numerous and complex molecular and cellular mechanisms leading to mortality and underdevelopment in response to environmental stress for this species. As few differences were observed in the enzymatic biomarkers total antioxidant capacity and lipid peroxidation between hatched larvae and the previously investigated encapsulated embryo response to thermal and osmotic stress, it is suggested that further studies could be undertaken using embryos encapsulated in egg masses, as it is less time consuming than working on hatched larvae.     

The grazing effect of limpets on the macroalgal community of a rocky intertidal shore

Abstract Abundance of macro-algae in the mid-littoral zone on a Victorian intertidal rocky shore varied seasonally, algae being rare in Summer and common in Winter and Spring. Field experiments demonstrated that, of two species of grazing limpets (Cellana tramoserica and Siphonana diemenensis), only Siphonaria had a major effect on the abundance of foliose algae (e.g. Scytopsiphon lomeniarid) and neither species had a great effect on the encrusting algae (Ralfsia spp.). There was no evidence of competition for food between the two species of limpets, in contrast with results found for similar species in New South Wales.     

Larval settlement can explain the adult distribution of Mytilus californianus Conrad but not of M. galloprovincialis Lamarck or M. trossulus Gould in Moss Landing, central California: Evidence from genetic identification of spat

We investigated the spatial distribution of adult and newly settled mussels (Mytilus galloprovincialis Lamarck, Mytilus trossulus Gould and Mytilus californianus Conrad) on the shore at Moss Landing, California to test the hypothesis that adult distributions are a result of settlement patterns. Adult M. californianus were most abundant on a wave-exposed rocky jetty and adults of Blue mussels (M. trossulus and M. galloprovincialis) were more abundant inside the protected Moss Landing harbor. Using taxon-specific polymerase chain reactions, we monitored recruitment during continuous 1-2 week intervals on fibrous scrubbing pads for 12 months in 2002-2003. All mussel species settled in greatest numbers on the exposed jetty, and Blue mussels settled in greater numbers there than did M. californianus. Because Blue mussels settled abundantly where their adults were rare, post-settlement mortality appeared to be the strongest influence on adult distribution. In contrast, M. californianus settled mostly in their adult habitat.     

Intraspecific variation in the population dynamics and growth of the limpet,Cellana tramoserica

Summary Populations of the limpet Cellana tramoserica (Sowerby) from high, mid, and low intertidal regions, and from a subtidal zone, were studied at Cape Banks (N.S.W.), Australia. Individuals from the subtidal population had the largest mean and maximum shell size, the low and high shore populations were intermediate, while the midshore population had both the smallest mean and maximum size. The density of adults showed the reverse trend: the midshore region had the greatest adult population density while the subtidal population had the smallest density. The density of juveniles and recruits was negatively correlated with tidal height in the intertidal areas, but the density of both age-classes was smallest in the subtidal region. The rate of growth of individuals was negatively correlated to the adult density of a site; individuals from the subtidal population grew the fastest, while those from the midshore region grew the slowest. The level of adult mortality of the four populations was similar, with an annual rate of between 50–60%. Juvenile mortality did, however, differ among populations; reduced percentages of juveniles reached adult size in the mid and low shore populations than in the highshore and subtidal populations.The expected lifetime fecundity of individual females differed among the four populations. Subtidal individuals were expected to spawn 40 times the amount of gonad material during their life than were individuals from the midshore population. Individuals from the high and low populations probably spawn 10 times more than those from the midshore region. Differences in the fecundities of individuals were not likely to be the result of genetic differences because experimental transplants and manipulations of density showed that individuals from all the intertidal populations could increase their growth rate to match that of subtidal individuals. The implications of the difference in lifetime fecundity among populations are discussed.