Habitat segregation and temporal variation in some tropical intertidal populations

Observations were made to document habitat segregation and seasonal changes in density for the most common intertidal organisms at a study site on the Pacific coast of Costa Rica. The pulmonate limpets Siphonaria gigas (Sowerby) and Siphonaria maura (Sowerby) were more abundant in the mid-high- and high-intertidal zone, whereas the keyhole limpet Fissurella virescens (Sowerby) was more abundant in the mid zone. Coiled gastropods were found in the mid- and mid-high-intertidal zone but were absent or scarce in the high zone. Barnacles and coralline algae were found throughout the intertidal zone and foliose algae were mostly present in the splash zone. Abundance of all species changed over the 2-yr period of the study. S. gigas and S. maura and the barnacle Chthamalus fissus Darwin exhibited well-defined fluctuations in density related to recruitment. No recruitment was observed for F. virescens or Tetraclita panamensis Pilsbry and populations of both species declined. Changes also occurred for coiled gastropods and were probably related to movement of adults. Coralline and foliose algae were more abundant during the rainy season than during the dry months. Far from being constant, populations on these tropical shores were highly dynamic with temporal changes rivaling those in the temperate zone.     

Limpet grazing on a physically stressful Patagonian rocky shore

Many theories of consumer control of communities have come from studies conducted in relatively benign, temperate zone rocky intertidal systems. Here, we examine gastropod grazing and the maintenance of bare space on a dry, wind-swept rocky shore of Patagonia, Argentina. Two limpet species are the primary intertidal grazers. Siphonaria lessoni dominates mid and high intertidal zones, while Nacella magellanica dominates the lower zone. In all zones, limpet densities are positively correlated with bare space and the occurrence of cracks. Tethering experiments revealed that: (1) physical stress sets the upper distribution limit of both limpets, (2) predators, such as oyster catchers, regulate Nacella populations and may restrict them to cracks and vertical surfaces, and (3) desiccation stress appears to drive similar crack distribution patterns of Siphonaria in the upper intertidal. Experimental removal of limpets in each intertidal zone indicated that limpets have: (1) no detectable effect in the high intertidal where physical forces dominate community organization, (2) weak impacts at mid-elevations as grazing only limited the abundance of fleshy algae with physical forces again dominating community structure, and (3) relatively stronger, but still weak impacts in the low zone. These results suggest that grazing impacts on Argentine rocky shores are weak in comparison to the physical stresses (e.g. high winds, low humidity) that largely determine structure in this system. The dominance of physical forcing in this system occurs despite having similar grazer densities to other temperate, but comparatively wet, rocky shorelines (e.g. British Isles) where top-down control is strong.     

Interspecific Competition for Shelters in Territorial and Gregarious Intertidal Grazers: Consequences for Individual Behaviour

Experiments have shown that interspecific interactions within consumer guilds can alter patterns of distribution, abundance and size of species. Plastic behavioural responses can be modulated by agonistic interactions. In many cases, consumers compete for space and shelters, and these interactions change the manner in which they exploit food. This study investigates the consequences of competition in the spatial and temporal organization of behaviour of intertidal grazers, which share algal resources and the use of rock crevices while resting, but exhibit different body sizes, spatial behaviour and foraging modes. We evaluate interaction strength between small gregarious Siphonaria lessoni and the larger territorial keyhole limpet Fissurella crassa and between S. lessoni and the medium-size gregarious chiton Chiton granosus. Using field manipulations and artificial arenas in the laboratory, we tested whether the use of crevices, micro-spatial distribution and activity are modified by the density of conspecifics and the presence of heterospecifics. Our results show that small-scale spatial segregation observed in the field between S. lessoni and C. granosus result from species-specific differences in habitat use. In turn, we found evidence that spatial segregation between F. crassa and S. lessoni results from highly asymmetric interference competition in the use of shelters. The presence of F. crassa reduced the use of crevices and growth rates of S. lessoni. Effects on growth rates are assumed to result from exposure to harsh environmental conditions rather than food limitation. Thus, neither gregarious behaviour nor differences in activity were sufficient to prevent competition with the larger grazer. Our study illustrates the importance of competition for shelters, which results in behavioural changes of the smaller-sized species, and how these plastic responses can translate into differences in growth rates. Use of shelters can thus be modulated by environmental conditions in a species-specific as well as an interactive manner within consumers’ guilds.     

Grazing patterns in Siphonaria gigas (Mollusca,Pulmonata) on the rocky Pacific coast of Panama

The pulmonate limpet Siphonaria gigas is the most abundant molluscan grazer in the mid zone on rocky, wave-exposed shores of the Pacific coast of Panama. Erect macroalgae and sessile invertebrates are rare; crustose algae cover ～90% of the rock. The relative abundance of a common blue-green algal crust (Schizothrix calcicola?) is negatively correlated with Siphonaria's abundance. Large-scale removals of the limpet cause rapid increases in percent cover of Schizothrix and concomitant decreases in other crusts, but no changes in the abundance of erect algae or sessile invertebrates. Removing Siphonaria also (1) increases recruitment of crustose algae and barnacles onto new rock and plexiglass substrata, and (2) decreases the abundance of a calcified form of Schizothrix. Harsh conditions during daytime low tides and foraging by fishes at high tide control the microdistribution of most of this region's mobile and sessile benthic organisms. Wave action and substratum heterogeneity modify these constraints: Siphonaria is rare or absent in sheltered areas, especially on homogeneous surfaces, and is most abundant at wave-exposed sites. However, at extremely wave-beaten sites, Siphonaria and other benthic consumers are rare and ineffective. Crustose algae are reduced in abundance and space is dominated by erect macroalgae and/or barnacles. These normally rare species can outcompete crusts only when thermal or desiccation stress and the effects of benthic consumers and fishes are drastically reduced.     

Differential survival of macroalgae to digestion by intertidal herbivore molluscs

This study evaluates the capacity of algal propagules to survive digestion by seven intertidal herbivore molluscs. Thirty individuals of Littorina peruviana, 36 of Siphonaria lessoni, 35 of Collisella ceciliana, and 25 each of C. zebrina, Chiton granosus, Fissurella crassa and F. limbata were collected at different dates between October, 1983 and April, 1984. About half of the individuals of each species were examined for gut contents and the other half was used to provide faecal pellets for culture. Diet of these grazers ranged from 8 macroalgal species in Collisella zebrina to 17 in Fissurella limbata. In total, macroalgal propagules of 56% of the 27 algal species found in these gut contents survive digestion. There was no positive correlation between diet and number of algal taxa growing in the respective faecal cultures. Survival through Littorina peruviana and Chiton granosus was low (20–30% of the algal species consumed) while it was high (75–83%) through the two species of Collisella. No significant correlation was found between frequency of algae in the gut contents and its frequency in the faecal cultures. Algal capacity to survive digestion was almost exclusively restricted to opportunistic species, especially in the Chlorophyta and Phaeophyta. It seems that survival of algal propagules through the digestive tract of generalist grazers is a rather random phenomenon.     

An intertidal limpet species as a bioindicator: Pollution effects reflected by shell characteristics

Mollusc shells have been widely used for monitoring the bioavailability of contaminants in the aquatic environment. The present work examined malformations among the shells of the limpet Siphonaria lessoni from heavily polluted, light polluted and unpolluted sites in Argentina. Data on shell shape, thickness, dry weight, microstructure and semi-quantitative elemental composition was evaluated as well as soft tissue dry weight. Shells from the heavily polluted site were significantly (p < 0.001) thicker than those from other areas. SEM (scanning electron microscopy) analysis of thickened shells revealed the presence of globular malformations on inner shell surfaces. On heavily polluted shells, elemental composition analysis by EDS (electron dispersive spectroscopy) of such malformations indicated concentrations three times higher of carbon and four times lower of calcium and oxygen than the control. Light polluted shells presented concentrations two times lower of calcium. In addition, soft tissues were lighter at the heavily polluted site (p < 0.001). These data demonstrate the sensitivity of this abundant and widely distributed intertidal limpet to aquatic pollutants, and support the use of this limpet as a potential biomarker.     

Impacts of a ‘black tide’ harmful algal bloom on rocky-shore intertidal communities on the West Coast of South Africa

Algal blooms commonly occur along the South African west coast. In March 1994 a dense bloom developed within St Helena Bay. Its subsequent decay caused near-shore hypoxia and elevated hydrogen sulphide levels, leading to it being termed a ‘black tide’. The bloom caused immediate massive intertidal mortalities (95% reduction of biomass), significant changes in community structure, and declines in richness and diversity. Only four taxa were not diminished: ephemeral algae, the gastropod Afrolittorina knysnaensis, the anemone Bunodactis reynaudi and the false limpet Siphonaria capensis. This limpet species subsequently increased at impacted sites relative to controls, probably reflecting reduced competition from other grazers. Ephemeral algae flourished during early recolonisation while grazers were depleted, but declined as grazers re-established. Manipulative experiments demonstrated that grazers do have this capability. Perennial algae also flourished, but lagged behind ephemerals, possibly contributing to their decline. Predators took longest to recover. Differences in the responses of species reflect differing tolerances, zonation patterns and biological interactions. Communities at control sites remained unchanged over the four-year study, and those exposed to a low intensity of the bloom were little affected, but communities exposed to its full intensity failed to recover fully within four years. Increasing frequency of harmful algal blooms along the west coast of South Africa could, therefore, significantly alter rocky-shore communities if intervals between bloom events become shorter than recovery times.     

Sewage pollution and extinction risk: an endangered limpet as a bioindicator?

The mollusc Patella ferruginea, endemic to the Mediterranean, is the most endangered marine species on the list of the European Council Directive 92/43/EEC and it is under serious risk of extinction. In spite of the low abundances and restricted distribution of this limpet, important populations have been found in the harbour of Ceuta, north Africa. The main objective of the present study was to characterise, for the first time, the effects of sewage pollution on P. ferruginea and related limpet species, and to evaluate the potential value of these limpet assemblages as bioindicators, using univariate and multivariate analyses. Physicochemical parameters and limpets were sampled in nine stations located at 0, 1, 2, 4, 8, 16, 32, 64 and 128 m away from the discharge point of a sewage effluent in Ceuta harbour. The stations closer to the outfall (0, 1, 2, 4 and 8) were characterised by higher values of turbidity, phosphate and ammonia in the water column, and organic matter, faecal coliforms and faecal Streptococci in sediments. A total of six limpet species were found and studied (Patella ferruginea, P. caerulea, P. nigra, P. rustica, P. ulyssiponensis and Siphonaria pectinata); the number of limpet species increased with increasing distance from the outfall, while diversity and evenness reached the highest values at intermediate sites. Siphonaria pectinata and P. caerulea were the most resistant and abundant species, while P. ferruginea was the most sensitive species to sewage pollution, only found at stations from 32 to 128 m. The distribution of this endangered limpet seems mainly affected by the pollution gradient, and not by the competition with the remaining limpets. The results of this study should be taken into account in future programmes of management and conservation of P. ferruginea.     

Causes of rarity and range restriction of an endangered, endemic limpet, Siphonaria compressa

The pulmonate limpet Siphonaria compressa is South Africa's most endangered marine mollusc. It is endemic to just two localities: Langebaan Lagoon on the west coast, and Knysna Estuary on the south coast, and occurs only on the eelgrass Zostera capensis. In Langebaan Lagoon, eelgrass has fluctuated substantially over the last 34 years, and S. compressa has twice approached extinction. S. compressa is largely confined to the lower edge of the eelgrass beds there, being replaced higher up by another small gastropod, Assiminea globulus. We explored the physical and biological factors underlying the limpet's narrow habitat, using field observations, translocations, caging and transplant experiments. Abundance of S. compressa was positively correlated with Z. capensis cover and negatively correlated with shore height. When moved to the upper portions of the eelgrass bed, S. compressa had lower rates of persistence and survival than in the lowest zone. The lower limit of zonation for S. compressa was set indirectly by bioturbation by the sandprawn Callianassa kraussi, which excluded eelgrass from intertidal sandbanks. Transplants of eelgrass into the sandbanks proliferated provided C. kraussi was experimentally eliminated, and supported densities of S. compressa 20-fold greater than in control eelgrass beds, suggesting that high-shore eelgrass beds to which S. compressa is normally confined are suboptimal for the limpet. A. globulus showed patterns opposite to those of S. compressa: its persistence and survival were greatest in the upper zone and it actively avoided the lower sections of these beds and never colonised eelgrass transplanted into sandflats lower on the shore. There was no evidence that competition between S. compressa and A. globulus influenced the zonation or abundance of either species. Rarity of S. compressa and its endangered status seem dictated by its extremely narrow and temporally changeable habitat-range, which is defined by physical stress in the high-shore and bioturbation by C. kraussi in the low-shore. Fluctuations in eelgrass abundance and limitation of S. compressa to just two localities add substantially to the risks of extinction for this embattled stenotypic limpet.     

Intertidal and Subtidal Fouling Assemblages in a Patagonian Harbour (Argentina, Southwest Atlantic)

The composition of the early stages of intertidal and subtidal fouling assemblages in Comodoro Rivadavia harbour (Argentina, 45°52′ S, 67°28′ W) and the influence of shore level and season on their structure were analysed. At the beginning of each season, stones were glued to the substratum with epoxy putty and distributed along 4 vertical transects at intervals of 20 m, at 3 levels: upper intertidal, middle intertidal, and subtidal. Substrata remained in the field for 84–100 days. A total of 48 samples (4 seasons × 3 levels × 4 replicates) were analysed. Species richness increased with depth, with 6 taxa in the upper intertidal, 23 in the middle intertidal and 31 in the subtidal. Seasonal differences in richness were less distinct. Green, red and brown algae were the dominant groups. Invertebrates were mainly represented by filter-feeding, sessile organisms, such as cheilostome bryozoans, spirorbid polychaetes and acorn barnacles. The barnacle Balanus glandula and the bryozoan Cryptosula pallasiana were the only non-indigenous species found in this study. Highly significant differences in structure among shore levels and seasons were evidenced by a two-way ANOSIM test. The upper intertidal is characterized by the filamentous green algae Urospora penicilliformis and Ulothrix flacca. The barnacle Balanus glandula is the most abundant species in the middle intertidal. The subtidal is defined mainly by the presence of the spirorbid polychaetes Paralaeospira levinseni and Romanchella perrieri, and the keyhole limpet Fissurella radiosa. Ordination of samples by season was less clear than by shore level.     

Competition between laboratory populations of green leafhoppers,Nephotettix spp. (Homoptera: Cicadellidae)

Intra- and interspecific competition between laboratory populations of four green leafhoppers, Nephotettix spp. was studied in the laboratory under three different temperature regimes of 24°C, 27°C and 30°C. For the single-species population of the three tropical species, the equilibrium density increased as the temperature increased. On the other hand, for the temperature species N. cincticeps, the highest equilibrium density was at the intermediate temperature and the lowest at high temperature. Interspecific interactions between two tropical (N. virescens vs. N. nigropictus), a tropical and a temperature (N. virescens vs. N. cincticeps) and a rice-feeding and a grass-inhabiting (N. virescens vs. N. malayanus) Nephotettix species were also studied in the laboratory at the three temperature regimes. Temperature differentially affected the outcome of competition between two Nephotettix species. Between N. virescens and N. nigropictus, the latter was more successful over the former at low and intermediate temperatures, while the former was more successful at high temperature. Between N. virescens and N. cincticeps, the temperate species inhibited the growth of the tropical species at low temperature while the tropical species inhibited the growth of the temperate species at high temperature. At intermediate temperature, the population of N. virescens persisted at a slightly higher density over the population of N. cincticeps. Between the rice-feeding N. virescens and the grass-inhabiting N. malayanus, regardless of temperature the population density of the latter was greatly reduced and later became extinct while the population of the former continued its growth. These consequences of competition between two Nephotettix species conformed fairly well to those predicted by theLotka-Volterra model using demographic parameters specified for each species.     

Demography, competitive interactions and grazing effects of intertidal limpets in southern New Zealand

Population dynamics and the effects of intraspecific competition on limpet growth and maintenance of bare patches were investigated for the intertidal limpet Cellana ornata (Dillwyn) at a boulder-dominated site and on a rocky platform near Kaikoura (South Island), New Zealand. Distribution and abundance patterns of C. ornata were described in relation to other biota and tidal height. C. ornata occurred almost exclusively in patches devoid of macroalgae, particularly in the mid-tidal zone. Both adult and juvenile limpets were most abundant on the tops of boulders, where their numbers were positively correlated with barnacle cover, which averaged 77%. The size structure and growth patterns of C. ornata were different between populations. Mark-recapture studies showed that the slopes of annual growth increments regressed on initial sizes were similar at both sites but that the annual increments on the platform were about 6 mm greater than on boulders. Growth virtually ceased at 27 mm for limpets at the barnacle-dominated boulder site and at 40 mm at the platform site. Recruiting cohorts had 20% survival on boulders and 37% on the platform during their first year. The largest size classes at both sites had around 57% annual survival. To test the effects of varying limpet densities on the growth and mortality of limpets and the maintenance of bare patches, densities of C. ornata were experimentally increased at both sites. Beyond a density of 4 per 0.25 m², sizes and survival of limpets were reduced at both sites, but the effect was more pronounced at the boulder site. Limpets at the boulder site were more effective at maintaining bare space than those on the reef platform. Enclosing limpets in plots with and without barnacles showed that C. ornata and a co-occurring species (Cellana denticulata (Martyn)) grazed more effectively and had greater growth in cleared plots. Overall, there was considerable variation in the demographics of C. ornata between populations driven by site-specific characteristics.     

Vertical distribution of chitons (Mollusca: Polyplacophora) in the rocky intertidal zone of central Chile

The patterns of vertical distribution of chitons in various exposed rocky shore habitats of central Chile are described. The density and size of the individuals were determined at different levels on intertidal rock walls, in pools and on boulders. A group of four species was found on rock walls while another group of four species was found in tidal pools and on boulders. The species showed vertical separation on rock walls and segregation according to a range of increasing frequency of water movement in pools and on boulders. Only the two species with largest adult individuals (Acanthopleura echinata (Barnes) and Chitonlatus Sowerby) showed vertical segregation in size. Small-sized species and small individuals of large-sized species were found higher on rock walls or in zones where water movement is less; large chitons occur in zones of greatest water impact. This suggests that water movement may be a significant factor in setting the lower limits of distribution of these herbivores. Some of the species examined were extremely abundant (600·m^-2) with densities of one to two orders of magnitude greater than for other chiton species elsewhere.     

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.     

Effects of light and intertidal position on seedling survival and growth in tropical tidal forests

Reciprocal transplant experiments were used to study the effects of tidal inundation and light level on growth and survival of four species of mangroves in Australia: Avicennia marina (Forsk.) Vierh., Bruguiera gymnorrhiza (L.) Lam., Ceriops tagal C.T. White, and Rhizophora stylosa Griff. Seedlings were planted in the high or low intertidal and in light gaps or under the shade of a closed canopy. Survival and growth of the seedlings were monitored for 30–36 months. Significant differences in survival were found among species, between intertidal zones and due to light level. Averaged across intertidal zones and light level, survival was greatest for Rhizophora and decreased in the order Ceriops, Avicennia, and Bruguiera. For all species survival was greater (P ⩽ 0.001) in the high than in the low intertidal treatment, regardless of light level. Within the high intertidal all species survived better in light gaps than under the canopy. Relative growth of Rhizophora stylosa, Avicennia marina, and Ceriops tagal were greater in the high versus low intertidal and in gaps versus under the canopy. For Bruguiera gymnorrhiza growth was not significantly different between gap and canopy or high and low intertidal. Although Avicennia marina, Bruguiera gymnorrhiza, and Rhizophora stylosa survived and/or grew best in the high intertidal they reach maximum abundance in the low intertidal. Ceriops tagal, however, performed best in the region where it is most abundant, yet even there, it was out-performed by the former three species. These results indicate that the species zonation patterns often observed across the intertidal cannot be explained by physiological adaptation alone. Factors such as propagule dispersal, competition and predation on propagules may also be important.     

Polymorphism and selective predation in Donax faba Gmelin (Bivalvia: Tellinacea)

Donax faba Gmelin is abundant in the upper intertidal zone on wave-beaten sandy beaches throughout East Africa. It maintains a feeding position within the wash zone by migrating up and down the beach with the tide. The species is highly polymorphic for shell colour; 14 morphs are present in the population studied. During migration, D. faba is preyed upon by a crab Ocypode ceratophthalmus (Pallas) and a wading bird, Crocethia alba (Pallas). Predation by the crab is shown to be non-random with respect to morph. The commonest morph, which has a disruptive colouration, is more heavily predated when the population density is low and a non-cryptic, rarer morph when density is high. Selection is therefore stabilizing (or directional) at high density and apostatic at low density. The polymorphism is shown to be stable over the period of the investigation (one year).     

Alternative methods of measuring competition applied to two Australian hermit crabs

Summary This article describes the intertidal hermit crab species assemblage at One Tree Island, Great Barrier Reef, Australia. Competition for shells between the two most abundant species, Clibanarius virescens and Calcinus latens, is studied in more detail. Competition appears to be primarily exploitative. The relative intensities of inter- and intra-specific competition between this pair of species are estimated using two different methods. The first is based on habitat overlap data in conjunction with a mathematical model of shell population dynamics described in Abrams (1980). The second method is more direct, and is based on following the fate of marked empty shells. Results of the two methods are very similar. This supports the validity of assumptions made in applying the first method. The relative amount of interspecific competition between these two species appears to be greater than that for other hermit crab species pairs studied previously.     

Exposure to solar radiation drives organismal vulnerability to climate: Evidence from an intertidal limpet

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.     

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.