Feeding ecology of a size-structured predator population,the South American sun-star Heliaster helianthus

Patterns of feeding in a population of Heliaster helianthus (Lamarck), a common and dominant species of starfish indigenous to the Pacific South American coast, were investigated in an intertidal habitat in central Peru from October 1986 to April 1987. The H. helianthus population comprised individuals of 3.5 to 30.2 cm body size (diameter) with two modal size classes. The number of rays ranged between 18 and 40, and individuals with 31 to 33 rays accounted for ca. 42% of the total population. There was a higher rate of increase in ray number with body size amongst small individuals(<13.0 cm diam). H. helianthus is capable of feeding on more than one prey item at a time (average of 5.6 to 13.2 prey items handled, with several predators observed to hold >100), and both the number of prey individuals captured and the total prey biomass were significantly correlated with predator size. Amongst a total of 1132 feeding observations, the largest number of predators (an average of 85.4% of those feeding) were preying on the mussel Semimytilus algosus whilst another mussel, Perumytilus purpuratus, ranked second with 21,9% of predators feeding. The proportion of S. algosus in the diet increased from 65.4% in the smallest predator size-group (10.9 cm diam) to 91.2% in the largest (19.0 cm). In contrast, P. purpuratus and barnacles were more highly represented in the diet of small H. helianthus. The smallest size-group (10.9 cm) had low dietary overlap with larger sizes and less specialized prey utilization. Two geographically separated populations of H. helianthus in Peru and Chile showed contrasting patterns of prey utilization. S. algosus and P. purpuratus comprised 85.5 and 6.5% by number in the diet of the Peruvian population, respectively, whilst corresponding figures for the Chilean population were 8.3 and 60.5%, with barnacles attaining a higher share (22.6%). However, the total number of prey individuals per feeding predator was almost the same in Peru and Chile, with 10.0 and 10.7 individuals, respectively. H. helianthus individuals of different sizes occupy slightly different microhabitats within the intertidal area, which, coupled with differential spatial distribution of prey species, results in the predator population being able to utilize a wide range of resources.     

Local and meso-scale patterns of recruitment and abundance of two intertidal crab species that compete for refuges

Interference competition for limited habitat or refuges is known to produce density-dependent mortality and generate patterns of micro-habitat distribution. While in mobile species the outcome of interference at a local scale can usually be determined from differences in body size and behavior, the population-level consequences of such interactions vary depending on rates of settlement and recruitment at a site, which are not directly correlated to local reproductive success. Previous experimental studies in central Chile demonstrated that interference competition for refuges is the primary factor driving microhabitat segregation between the predatory crabs Acanthocyclus gayi and Acanthocyclus hassleri, with the latter species monopolizing galleries inside mussel beds and excluding A. gayi to rock crevices. Between April 2001 and March 2006 we quantified monthly recruitment rates in artificial collectors at 17 sites over 900 km of the central coast of Chile. Results show that recruitment rates of A. hassleri are almost two orders of magnitude lower than those of A. gayi, and that they are tightly and positively correlated among sites across the region, suggesting that at scales of kilometers larval stages of these species are affected by similar oceanographic processes. Total crab densities per site were also positively correlated between species and strongly associated to mussel cover, with overall low crab densities at all sites where mussel cover was lower than about 60%. At all sites with mussel cover >60%, the ratio of A. gayi to A. hassleri density progressively decreased from recruits (2.6) to juveniles (0.5) to adults (0.04), overcoming initial differences in recruitment rates. The relative success of the inferior competitor at sites with low mussel cover does not appear to provide a potential mechanism favoring regional coexistence through dispersal to other sites (“mass effects”), because their densities were lower than at sites of high mussel cover. Yet, at many sites of low mussel cover the dominant competitor is virtually absent, allowing A. gayi to attain larger population sizes at the scale of the region. Thus, the factors limiting the dominant competitor from successfully utilizing other microhabitats seem to be the most critical factor in promoting both local and regional coexistence between these species.     

Depression of feeding and growth rates of the seastar Evasterias troschelii during long-term exposure to the water-soluble fraction of crude oil

To test the effect of petroleum hydrocarbons on predation by the seastar Evasterias troschelii (Stimpson, 1862) on the mussel Mytilus edulis (L.), we exposed the predator with the prey to six concentrations of the water-soluble fraction (WSF) of Cook Inlet crude oil. Seastars and mussels were collected at Auke Bay, Alaska, in November 1980. During a 28 d exposure in a flow-through system, seastars were more sensitive to the WSF than mussels: the LC₅₀ for the seastars was 0.82 ppm at Day 19 and, although no mussels were exposed to WSF for more than 12 d, none died. Daily feeding rates (whether in terms of number of mussels seastar^-1 d^-1 or dry weight of mussels seastar^-1 d^-1) were significantly reduced at all concentrations above 0.12 ppm. At 0.20, 0.28 and 0.72 ppm WSF, daily feeding rates (in terms of dry weight of mussels) were, respectively, 53, 37, and 5% of the control rate; at the two highest concentrations (0.97 and 1.31 ppm WSF), the seastars did not feed. Seastars at concentrations greater than 0.12 ppm WSF grew slower than individuals from the control group and the 0.12 ppm-treatment group combined. These laboratory results show that E. troschelii is more sensitive to chronic low levels of the WSF of crude oil. The possibility that such oil pollution could reduce predation and permit M. edulis to monopolize the low intertidal zone of southern Alaska remains to be studied.     

Prey capture by Pisaster brevispinus (Asteroidea: Echinodermata) on soft substrate

Observations on the subtidal seastar Pisaster brevispinus (Stimpson) indicate that this predator can extend the central tube feet into sand-mud substrate for a distance roughly equal to the radius of the seastar. Field and laboratory evidence demonstrates the use of the elongated tube feet by the asteroid for the capture of burrowed prey items. A relationship between predator size and the size of juveniles of the deep-burrowing clam Tresus nuttallii taken as prey is a product of the ability of larger seastars to extend the tube feet further into the substrate and capture deeper prey. The role of P. brevispinus as an important subtidal predator along the Pacific Coast of North America is discussed.     

Prey defense,predator preference,and nonrandom diet: The interactions between Pycnopodia helianthoides and two species of sea urchins

Interactions between the predatory sea star Pycnopodia helianthoides (Brandt, 1835) and two of its natural prey, the sea urchins Strongylocentrotus purpuratus (Stimpson, 1857) and S. franciscanus (Agassiz, 1863), are examined with regard to predator preference, predator diet, and prey defenses. The sea star is able to detect both species of sea urchin upstream in a Y-trough, but does not consistently choose one over the other (i.e., no preference). However, when the sea star is presented with equal numbers of similar-sized specimens of the two species of sea urchin, its diet is markedly nonrandom, since S. purpuratus is eaten almost 98% of the time. The defensive responses of the two species of sea urchin differ in form and effectiveness. S. franciscanus employs its long spines as defensive weapons, pinching the rays of an attacking sea star. This defensive response is more effective than the pedicellarial response used by S. purpuratus. The nonrandom diet of the predator seems to result primarily from prey defensive responses that differ in effectiveness, rather than from an intrinsic, behavioral preference of the predator at an earlier stage in the predator/prey interaction.     

Population genetic structures of the fissiparous seastar Coscinasterias acutispina in the Sea of Japan

The morphological characteristics and the population genetic structures of the fissiparous seastar Coscinasterias acutispina were investigated for eight sites in the Sea of Japan in order to clarify the presence of sexual and asexual reproduction. Morphological observation based on arm length showed that fission was common at all eight sites examined, indicating the likely production of clonal individuals. A random amplified polymorphic DNA (RAPD) marker was used to detect clones arising by fission and to assess gene flow among sites. A simulation approach using RAPD data revealed the presence of clonal individuals at almost all sites, suggesting the existence of asexual reproduction. The result of phylogenetic analysis according to RAPD genotype showed no relationship between genetic and geographic distances. Considering the limited movement ability of seastar species during the adult phase, these observations suggest the existence of marked gene flow among sites, due to dispersal of planktonic larvae produced by sexual reproduction. These observations suggest that multi-locus genotypic compositions depend on the relative amounts of recruitment from sexual and asexual reproduction in each population.     

Quantitative analysis of foraging behaviour in a field population of the South American sun-star Heliaster helianthus

Foraging behaviour in the South American sunstar Heliaster helianthus (Lamarck) was investigated quantitatively on a subtropical rocky shore in central Peru. H. helianthus feeds mainly on two mussel species, Semimytilus algosus (Gould) and Perumytilus purpuratus (Lamarck). A sequence of foraging behaviour was described and observations were made of the timing of attacks on mussel beds by sun-stars. Although H. helianthus is capable of foraging out of water, an unusual trait for asteroids, the timing of such foraging appears to be well adjusted to avoid the risk of prolonged heat and desiccation. Foraging activity began 4 h 52 min before high tide and 3 h 19 min after high tide, with a mode between 2 h 30 min and 2 h before high tide. Fifty-two percent of all foraging activity began between 3 h 30 min and 2 h before high tide, while only 12.4% began after high tide. This suggests that H. helianthus mainly relies upon changes in the rate of tidal increase as a cue to begin foraging. Foraging activity ceased between 3 h 40 min before and 4 h 29 min after high tide, with 48% ending between 2 and 0.5 h before high tide. The duration of foraging ranged from 19 to 190 min, with values between 30 and 80 min accounting for 67.4% of all observations. The median duration was 62 min. No significant correlation was detected between the time when foraging activity commenced and its duration. The intensity of foraging activity varied on consecutive days, with a general pattern of decreasing intensity after a day of relatively high activity. Foraging location in relation to a mussel bed was analysed on a marked, 8 m stretch of rocky shore. The numbers of foraging H. helianthus observed on different sections of the shore were related neither to the width of the mussel zone nor to the vertical position of the lower edge of the mussel zone, indicating that sun-stars do not rely upon these factors to assess prey availability and that ideal free distribution with regard to prey abundance does not occur on the spatial scale examined.     

Thermal tolerance of Strongylocentrotus purpuratus early life history stages: mortality, stress-induced gene expression and biogeographic patterns

In this study, we examined the differential thermal tolerance of Strongylocentrotus purpuratus early life history stages by comparing high temperature–induced mortality and the relative levels of the stress-induced gene, hsp70, between S. purpuratus embryos and larvae from adults collected throughout the species range. There was no significant difference between gastrulae and 4-arm plutei mortality from all sites examined. Furthermore, there was little variability in temperature tolerance across the biogeographic range as southern gastrulae and 4-arm plutei exhibited similar tolerances to northern individuals. Relative levels of hsp70 mRNA expression did not differ overall between the two developmental stages at each site. Across sites, all gastrulae and 4-arm plutei exhibited maximum hsp70 expression at approximately 25°C; however, the range of hsp70 expression was narrower in southern individuals, suggesting they are living closer to their upper thermal limit than northern individuals.     

A battery-operated,continous-flow enclosure for metabolism studies in benthic communities

Chemoreception was studied in Luidia clathrata (Say), collected from Old Tampa Bay, Florida, USA, in June 1983. L. clathrata has chemotactic responses to compounds associated with its preferred dietary items. This seastar responded most strongly to L-cysteine, L-cystine, L-isoleucine and L-glutamic acid, compounds associated with fleshy animal foods. Sugars (sucrose, maltose and lactose), compounds associated with plant and detrital foods, were also stimulatory, but less so than the aminoacids. The seastar showed a strong response to the freshtissue component, betaine. The excretory product ammonium, indicative of metabolic activity of live prey, elicited a weak response. Chemotactic responses were concentration-dependent. Concentration-response curves for L-cysteine, L-isoleucine, and L-glutamic acid are steeply sloped and have response-dose₅₀ values within the range of 1×10^-8 and 1×10^-5 M. L. clathrata is adapted to sensing concentration gradients at levels which could be expected in its environment. Distance chemoreception is an important component of both search behavior and food discrimination for this seastar.     

β-glucanase activity in the seastar Pisaster ochraceus

Specimens of Pisaster ochraceus, an intertidal carnivorous seastar, were collected in northern Puget Sound at Post Point, near Bellingham, Washington, USA, in November 1993, and used in experimentation through September 1994. Ammonium sulfate fractions (60% precipitates) from excised pyloric caeca (digestive glands) possessed -glucanase activity directed toward such -linked substrates as carboxymethylcellulose (CMC), partially-digested cellulose (cellodextrins) and the synthetic substrate p-nitrophenyl--D-glucoside (PNDG). pH-activity and pH-stability maxima were at pH 6 and 5, respectively. No hydrolytic activity was detectable on native cellulose, native chitin or on cationic-modified celluloses such as DEAE-cellulose. Gel filtration (Sephadex G-100) of this 60% precipitate followed by DEAE-Sephadex column chromatography revealed heterogeneity of -glucanase activity, with an indication of up to 11 distinct fractions with variable activity directed toward the substrates indicated above (CMC, cellodextrins and PNDG). These fractions displayed differing ratios of hydrolytic activity on these substrates. None of the isolated fractions displayed activity toward either native cellulose or native chitin. Gut extracts from the edible mussel (Mytilus edulis), a common prey species of P. ochraceus, were shown to possess partially digested cellulose (cellodextrins). These cellodextrins were further degraded by seastar -glucanases in vitro. The digestion of the gut contents of ingested prey provides a possible adaptive explanation for the presence of -glucanase activity in a carnivore.     

Long-term change in mussel (Mytilus californianus Conrad) populations along the wave-exposed coast of southern California

Mussel (Mytilus californianus) populations were studied throughout California to determine changes over the past few decades, and, in particular, to determine if declines reported for southern California have occurred outside of the region. We compared mussel cover, biomass, and bed thickness reported in historic studies in the mid-1970s to 1980s with measurements made in 2002. Mussel cover and biomass in southern California declined markedly over the past few decades with a mean cover loss of 31.2% (40.2% loss) and biomass loss of 25.1 kg m⁻² (51.3% loss). Changes in mussel bed thickness were not as strong as cover and biomass, but also appeared to decline over time. Declines were limited to the southern California region, since mussel cover, biomass, and bed thickness remained unchanged or increased at sites in central and northern California. Causes for mussel declines in southern California are unknown, but may include human visitation, increased sea surface temperatures, and pollution.     

Population biology and antipredator defenses of the shallow-water Antarctic nudibranch Tritoniella belli

The circumpolar nudibranch Tritoniella belli Eliot occurs in abundance in shallow-water benthic communities of McMurdo Sound, Antarctica. Density estimates based on belt transects averaged collectively 0.46 individuals m⁻² at three study sites between depths of 6 and 30 m in November 1996. At two of the sites, population densities increased linearly between 18 and 30 m depth (up to 0.7 and 1.15 individuals m⁻² at 30 m depth). Individuals at all sites were rare or absent at depths shallower than 12 m. Size frequencies of individuals at the sites were similar, and a pooled analysis revealed a unimodal distribution skewed highly towards juvenile size classes. This suggests both recent recruitment and constant rates of mortality across size classes. The relationship between foot length and wet weight best fits an exponential growth equation, indicative of an allometric growth pattern. Distribution of T. belli in the field suggests that it is a habitat and diet generalist. Potential invertebrate predators include sea anemones and seastars, both of which co-occur in abundance in McMurdo Sound. Laboratory experiments indicate that the sea anemone Isotealia antarctica can capture and ingest T. belli. However, 70% of T. belli that are captured escape from the tentacles or, following ingestion, are rejected from the gastrovascular cavity. The seastars Odontaster validus, Perknaster fuscus, and Acodontaster conspicuus, avoid contact with T. belli, but if forced into contact with mantle tissues, retract their tube-feet. Mucus secreted from the mantle tissues, coated on to the tips of glass rods, and presented to seastar tube-feet, causes significantly longer tube-foot retraction times than control rods. Moreover, pieces of freeze-dried krill coated with mantle mucus are consumed significantly less often than untreated control pieces of krill by a benthic scavenging fish (Pseudotrematomas bernacchi). Employing seastar tube-foot retractions as a bioassay, we found the bioactive compound(s) are soluble in ethyl acetate, indicating they are lipophilic or moderately hydrophilic in nature. Chemical defenses in the mucus of T. belli probably contribute to its high abundance in Antarctic benthic communities. Received: 6 October 1997 / Accepted: 24 March 1998     

Rats on the run: removal of alien terrestrial predators affects bush rat behaviour

Predators can strongly influence the microhabitat use and foraging behaviour of prey. In a large-scale replicated field experiment in East Gippsland, Australia, we tested the effects of reduced alien red fox (Vulpes vulpes) and alien wild dog (Canis lupus familiaris) abundance (treatment) on native bush rat (Rattus fuscipes) behaviour. Bush rats are exposed to two main guilds of predators, namely mammalian carnivores and birds of prey. Tracking rat movements using the spool-and-line technique revealed that, in treatment sites, rats used ground cover, which provides shelter from predators, less often than at unmanipulated fox and wild dog abundance (non-treatment sites). In treatment sites, rats more frequently moved on logs where they would have been exposed to hunting foxes and dogs than in non-treatment sites. Furthermore, in treatments, rats showed a preference for understorey but not in non-treatments. Hence, bush rats adapted their behaviour to removal of alien terrestrial predators. Giving-up densities (GUDs) indicated no treatment effects on the marginal feeding rate of bush rats. Interestingly, GUDs were higher in open patches than in sheltered patches, suggesting higher perceived predation risk of bush rats during foraging at low versus high cover. The lack of treatment effects on GUDs but the clear response of bush rats to cover may be explained by the impact of predators other than foxes and wild dogs.     

Wave exposure effects on population structure and recruitment in the mussel <Emphasis Type="Italic">Perna perna</Emphasis> suggest regulation primarily through availability of recruits and food,not space

Recruitment and population structure of Perna perna in low shore mussel beds were investigated over 15 months at six sites along the south coast of South Africa. Initial, subjective classification of sites as wave exposed or wave sheltered (three of each) was confirmed using the dissolution of cement blocks to measure average water flux and dynamometers for maximum wave force. Recruitment occurred throughout the year, but recruit (1–5 mm) densities were significantly higher from January to April 1996 on both shore types. Recruit densities were positively correlated with adult (>15 mm) densities for both shore types (P < 0.05) but the correlations were extremely weak (r ² < 0.06 in each case). In areas with 100% cover, adult size (mean and maximum lengths) was greater on exposed sites, but density showed the reverse and was negatively correlated with maximum wave strength (r = −0.84). Despite differences in adult densities and sizes, biomass, which is a product of the two, showed no significant difference between the two shore types (ANOVA P > 0.05). Thus wave exposure dramatically affects density, recruitment and mussel size, but not recruitment timing or biomass where there is 100% cover, and mediates a three-way interaction among food supply, larval supply and intraspecific competition for space. In contrast to shores with saturation recruitment, mussel biomass here appears to be limited by recruit supply and constraints of food, especially on sheltered shores, while density is regulated through intraspecific competition for space primarily on exposed shores and at small spatial scales.     

Settlement and recruitment of sea urchins (Strongylocentrotus spp.) in a sea-urchin barren ground and a kelp bed: are populations regulated by settlement or post-settlement processes?

I sampled recruitment of very small sea urchins (Strongylocentrotus spp.) by using the anesthetic magnesium chloride to remove individuals from substrata collected in sea-urchin barren grounds (barrens) and kelp beds at Naples Reef near Santa Barbara, California, USA. Preliminary sampling found low numbers of newly settled individuals(<0.6 mm test diam) from April–July in 1984 and 1985, and in April, 1986. In early May, 1986, I found many newly settled seaurchins (0.3 to 0.6 mm, 5 to 17 d old), and I compared the densities of the cohort on several types of natural substrata in barrens and kelp-bed habitats. Newly settled individuals of both purple sea urchins (S. purpuratus) and red sea urchins (S. franciscanus) were present in similar, high densities (1 000 S. purpuratus m^-2) on foliose red algal turf, a dominant substratum ofthe kelp bed, and on crustose coralline algae, the dominant substratum of an adjacent barrens. Larvae of S. purpuratus reared and tested in the laboratory showed high rates of settlement on both red algal turf and on crustose coralline algae, but significantly lower rates on rock. Larvae also settled in response to a partiallypurified extract of coralline algae. The reduced settlement on natural rock surfaces relative to either algal treatment and the significant settlement in response to the extract of coralline algae indicate that larvae discriminate between natural substrata and probably respond to a settlement cue other than, or in addition to, a simple microbial (bacterial) film. The similar densities of young recruits of S. purpuratus on dominant substrata of barrens and kelp bed show that, at least in this case, differential settlement cannot explain the high densities of sea urchins in the barrens habitat. Movement between barrens and kelp bed is unlikely given the small sizes of the newly recruited sea urchins relative to the large distances often involved. Reduced post-settlement mortality of newly settled individuals in the barrens remains the most likely mechanism leading to the higher densities of sea urchins in barrens relative to kelp-bed habitats.     

Feeding ecology of Liocarcinus depurator (Decapoda: Portunidae) in the Ria de Arousa (Galicia,north-west Spain): effects of habitat,season and life history

The diet of the portunid crab Liocarcinus depurator (L.) in soft-bottom areas of the Ría de Arousa (Galicia, north-west Spain) was studied by analysis of stomach contents, and by comparison of habitats of contrasting abundance, composition and diversity of prey communities. Monthly samples were taken from July 1989 to June 1990 (N=3747) in mussel (Mytilus galloprovincialis) raft-culture areas and central channel zones. Ninety-two food items were identified: crustaceans, molluscs, polychaetes, ophiuroids and fishes were the dominant prey. The raft epifauna and the mussels constituted the dominant group of prey in the raft areas and inner channel station (the epifaunal crab Pisidia longicornis comprised 30 to 52% of the diet), whereas the benthic macroinfauna was the dominant food component in the mid-outer channel (polychaetes constituted 32% of the diet). In the epifaunal community, diet selection for P. longicornis was positive and for amphipods was negative, whereas within the infauna diet selection for bivalves and ophiuroids was positive and for polychaetes negative or neutral. The seasonal importance of P. longicornis and M. galloprovincialis varied in the diet of L. depurator parallel to seasonal activities connected with mussel culture (seeding, thinning, harvesting) that determine the availability of prey from the rafts. Major changes in the composition of the diet related to the life history of L. depurator occurred during ontogeny. In the raft zones, consumption of mussels and fishes increased with increasing body size, while the consumption of Natantia, non-decapod crustaceans, gastropods, polychaetes Pectinariidae and plants decreased with increasing crab size. Maximum consumption of P. longicornis was by L. depurator individuals of intermediate size (15 to 44 mm carapace width). Stomach fullness (measured as dry weight of food) increased with increasing body size, but with negative allometry. During the late pre-moult and early post-moult periods L. depurator did not consume any food; maximum stomach fullness was recorded for the late post-moult and intermoult stages. Female food consumption decreased during egg incubation. Food consumption was greater in autumn and winter than at other seasons; this appears to be related to seasonal changes in reproduction and moulting.     

Ontogenetic changes in small-scale movement by recruits of an exploited mussel: implications for the fate of larvae settling on algae

Many South African populations of the brown mussel Perna perna have been depleted through over-exploitation by subsistence harvesters. This is problematic because recovery after disturbance is very slow, partly because recruits are largely associated with adult mussels. However, unlike large recruits of 3.5–10 mm that exhibit spatial structure related to that of adults, a very high proportion of settlers and small recruits (0–3.5 mm) occur on foliose algae. We tested the hypothesis that recruits on algae move to adult mussel beds after a period of growth, with the null hypothesis that they die at a certain size. We conducted an indirect field study comparing the ratios of large to small recruits in 100% cover mussel patches at locations with high and low algal cover. A second laboratory experiment analysed whether the size of recruits on algae affects their active movement behaviour in response to nearby mussel patches. Large/small recruit ratios were slightly, but not significantly greater in high than low algal cover locations. Both small (2–2.5 mm) and medium (4.5–5.5 mm) recruits remained on algae and moved very short distances throughout the laboratory experiment, while larger recruits (9–10 mm) moved significantly further distances and more often into mussel patches. The results suggest that very large recruits are able to migrate actively to nearby mussel patches, indicating ontogenetic shifts in this behaviour. However, the absence of a significant difference in ratios between field locations with high and low algal cover suggests many large recruits are accidentally dislodged from the algae and presumably die. Thus settlement of P. perna onto algae is likely to be wasted, with consequences for sustainable management of the mussel resource.     

Sex ratio and hermaphroditism in Oregon intertidal populations of the echinoid Strongylocentrotus purpuratus

Examination of large numbers of the echinoid Strongylocentrotus purpuratus (Stimpson) in the years 1969, 1970 and 1971 indicated that the sex ratio at Yaquina Head, Oregon (USA) was significantly different from 1:1, with males predominating. During 1968, 1970 and 1971 at Boiler Bay and Whale Cove, Oregon, the sex ratio did not differ significantly from 1:1 but, in 1969 at Boiler Bay, the proportion of males was 0.61. Two functionally hermaphroditic individuals were found among the 1,354 Yaquina Head echinoids examined, but none were found at Boiler Bay in an equal number of individuals. The hypothesis is proposed that S. purpuratus is a labile gonochorist, with a multiple, autosomal, sex-determining mechanism whose expression may be influenced by the environment.     

Is habitat amount important for biodiversity in rocky shore systems? A study of South African mussel assemblages

Habitat-forming species on rocky shores are often subject to high levels of exploitation, but the effects of subsequent habitat loss and fragmentation on associated species and the ecosystem as a whole are poorly understood. In this study, the effects of habitat amount on the fauna associated with mussel beds were investigated, testing for the existence of threshold effects at small landscape scales. Specifically, the relationships between mussel or algal habitat amount and: associated biodiversity, associated macrofaunal abundance and density of mussel recruits were studied at three sites (Kidd’s Beach, Kayser’s Beach and Kini Bay) on the southern and south-eastern coasts of South Africa. Samples, including mussel-associated macrofauna, of 10 × 10 cm were taken from areas with 100 % mussel cover (Perna perna or a combination of P. perna and Mytilus galloprovincialis) at each site. The amount of habitat provided by mussels and algae surrounding the sampled areas was thereafter determined at the 4.0 m² scale. A number of significant positive relationships were found between the amount of surrounding mussel habitat and the abundances of several taxa (Anthozoa, Malacostraca and Nemertea). Likewise, there were positive relationships between the amount of surrounding algal habitat and total animal abundance as well as abundance of mussel recruits at one site, Kini Bay. In contrast, abundance of mussel recruits showed a significant negative relationship with the amount of mussel habitat at Kayser’s Beach. Significant negative relationships were also detected between the amount of mussel habitat and species richness and total abundance at Kidd’s Beach, and between amount of mussel habitat and the abundance of many taxa (Bivalvia, Gastropoda, Maxillopoda, Ophiuroidea, Polychaeta and Pycnogonida) at all three sites. No threshold effects were found, nor were significant relationships consistent across the investigated sites. The results indicate that the surrounding landscape is important in shaping the structure of communities associated with these mussel beds, with significant effects of the amount of surrounding habitat per se. The strength and the direction of habitat effects vary, however, between shores and probably with the scale of observation as well as with the studied dependent variables (e.g. diversity, abundance, mussel recruitment, species identity), indicating the complexity of the processes structuring macrofaunal communities on these shores.     

Predator-prey relationships between the rock lobster Jasus lalandii and the mussel Aulacomya ater at Robben Island on the Cape West Coast of Africa

The natural diet and mode of feeding of the rock lobster Jasus lalandii (H. Milne Edwards) was determined in a rock-lobster sanctuary near Cape Town, South Africa. Field observations were tested and confirmed by means of aquarium studies. Rock lobsters feed mainly upon ribbed mussels Aulacomya ater (Molina), which comprise the largest component of the sessile benthic fauna. Mussel remains were found as the major constituent in 97% of the rock-lobster stomachs examined. The density of rock lobsters averaged 8,100 per hectare (0.81 m^-2), while mussel biomass averaged more than 5 kg (wet whole weight) m^-2 within the same depth range (12 to 30 m). More than 80% of the mussel biomass comprised large individuals between 60 and 90 mm in length. Large rock lobsters (mainly males) were capable of feeding on all sizes of mussels, although many of these were inaccessible to predation. Smaller rock lobsters became progressively more limited in the size range of mussels which they could crack open and consume. Competition between rock lobsters for small mussels appeared to be intense, as mussels of suitable size for feeding were generally in short supply to most of the rock-lobster population. Hence, feeding and growth rates of rock lobsters are likely to be affected by the relative population densities of predator and prey. Growth rates of rock lobsters could be limited by food supplies even in areas where mussel biomass is comparatively large.