Incidence and distribution of phototrophic shell-degrading endoliths of the brown mussel Perna perna

The incidence, distribution and infestation sequences of four endolithic cyanobacteria in the shells of the brown mussel Perna perna (L.) were studied along the south coast of South Africa. The incidence of endolith-infested shells varied significantly among the 21 study sites (∼23 to 95%), with highest infestation rates occurring on promontories and headlands as compared to sites within bays. At a smaller scale, the incidence of infested shells also varied with tidal height, being high at the upper tidal levels of mussel distribution and low or absent on the lowshore. The observation that small-scale variability in infestation rates was positively related to water movement suggested that physical damage to the outer protective periostracum of mussels may have facilitated colonisation by endoliths. This hypothesis was supported by the fact that shells with artificially damaged periostraca became infested at a greater rate than did control shells. Once colonisation by the filamentous cyanobacteria Plectonema terebrans, Hyella caespitosa and Mastigocoleus testarum had taken place, endoliths spread throughout the shell, causing progressive shell deformation and damage. Only older shells that were infested by the late successional, cavity-forming Pleurocapsa sp., however, exhibited severe shell deformations, became brittle and eventually fractured near the structurally important site of adductor muscle attachment. Heterotrophic endoliths typically associated with shell degradation in previous studies were extremely rare and if present did not contribute to shell disintegration. Although it is generally perceived that endolithic algae and cyanobacteria are restricted to the surface layers of shells by light limitation within the substratum, it is clear from this study that the interaction of a combination of factors (i.e. erosion of the periostracum, successional sequence of colonists and mechanical properties of the shell) may result in phototrophic endoliths causing severe shell degradation and eventually mussel mortality. Received: 18 December 1998 / Accepted: 18 June 1999     

Lethal and sub-lethal effects of phototrophic endoliths attacking the shell of the intertidal mussel Perna perna

Animals that bore into calcareous material can cause considerable damage to molluscan shells. In contrast, smaller microbial phototrophic endoliths have until recently been thought of as relatively benign. Phototrophic endoliths (primarily cyanobacteria) infest the shells of 50 to 80% of midshore populations of the mussel Perna perna (L.) in South Africa. This infestation causes clearly visible shell degradation, and we record here ecologically important lethal and sub-lethal effects (e.g. changes in growth and reproductive output) of the endoliths on their mussel hosts. Endolith infestation reduced the strength of shells significantly and also affected shell growth. In situ marking of shells, using the fluorochrome calcein, showed that infested and non-infested mussels increased in shell length at the same rate. However, the rate of increase in shell thickness (associated with shell repair) was significantly faster in infested than in uninfested individuals. This increase in the rate of shell thickening was not sufficient to compensate for rapid endolith-induced shell degradation and, around the site of adductor muscle attachment, infested shells were thinner than their uninfested counterparts. The shells of 18% of recently dead mussels had holes induced by endolith erosion. This effect was highly size dependent, and the proportion of mortality due to endoliths rose to almost 50% for the largest mussels. The re-routing of energy due to shell repair had important sub-lethal effects on the reproductive rates of mussels. During the reproductive period, mean dried flesh mass for large (>70 mm), non-infested P. perna was substantially higher than for infested individuals. This difference was almost entirely due to differences in gonad mass, which was approximately 100% higher for non-infested mussels. We conclude that, by attacking the shell, phototrophic endoliths reduce both the longevity and reproductive output of large mussels on the midshore. Received: 26 January 1999 / Accepted: 17 August 1999     

Effects of seawater temperature and pH on the boring rates of the sponge Cliona celata in scallop shells

Warmer, more acidic water resulting from greenhouse gas emissions could influence ecosystem processes like bioerosion of calcifying organisms. Based on summer-maxima values (temperature = 26 °C; pH = 8.1) at a collection site in New York (40°56″ N, 72°30″ W), explants of the boring sponge Cliona celata Grant, 1826 were grown for 133 days on scallop shells in seawater ranging from current values to one scenario predicted for the year 2100 (T = 31 °C; pH = 7.8). High water temperature had little effect on sponge growth, survival, or boring rates. Lower pH slightly reduced sponge survival, while greatly influencing shell boring. At pH = 7.8, sponges bored twice the number of papillar holes and removed two times more shell weight than at pH = 8.1. Greater erosion resulted in weaker scallop shells. This study suggests that lower seawater pH may increase boring rates of C. celata in shellfish, with potentially severe implications for wild and farmed shellfish populations.     

Spatial distribution of bacteria associated with the marine sponge Tethya californiana

Microbial diversity and spatial distribution of the diversity within tissue of the marine sponge Tethya californiana was analyzed based on 16S rRNA gene sequences. One candidate division and nine bacterial phyla were detected, including members of all five subdivisions of Proteobacteria. Moreover, chloroplast-derived Stramenopiles- and Rhodophyta-affiliated 16S rRNA gene sequences were found and Stramenopiles represented the most abundant clones (30%) in the clone library. On the phylum-level, the microbial fingerprint of T. californiana showed a similar pattern as its Mediterranean relative T. aurantium. An interesting difference was that Cyanobacteria that were abundantly present in T. aurantium were not found in T. californiana, but that the latter sponges harbored phototrophic Stramenopiles instead. Surprisingly, the phototrophic microorganisms were evenly distributed over the inner and outer parts of the sponge tissue, which implies that they also reside in regions without direct light exposure. The other phyla were also present in both the outer cortex and the mesohyl of the sponges. These results were confirmed by analysis on the operational taxonomic unit level. This leads to the conclusion that from a qualitative point of view, spatial distribution of microorganisms in T. californiana tissue is quite homogeneous. Thirty-two percent of the operational taxonomic units shared less than 95% similarity with any other known sequence. This indicates that marine sponges are a rich source of previously undetected microbial life.     

Distribution of Trypetesa lampas (Cirripedia,Acrothoracica) in various gastropod shells

The distribution of Trypetesa lampas (Hanoock) and its rates of infection of different species of gastropod shells in the waters of Anglesey were studied. T. lampas was found in depths from 3 to at least 25 fathoms (5 to 46 m). It infected dead shells, mainly of Buccinum undatum L., Colus gracilis (da Costa), Neptunea antiqua (L.) and Natica alderi (Forbes); it was absent from shells occupied by hermit crabs in the intertidal zone. Significantly more B. undatum shells were infected than shells of N. antiqua and Natica alderi, and more of both C. gracilis and N. antiqua than of Natica alderi. Larger shells of B. undatum were more likely to be infected than were the smaller ones. The number of females per shell varied with the shell length raised to the power 2.39 instead of as expected with the square of the length.     

Symbiotic zooxanthellae enhance boring and growth rates of the tropical sponge Anthosigmella varians forma varians

Several species of boring sponges harbor symbiotic zooxanthellae, and it is believed that the symbiont enhances boring activity of host sponges. This hypothesis was tested using manipulative field experiments to assess the effect of intracellular zooxanthella populations on boring rates of the tropical sponge Anthosigmella varians forma varians. Portions of sponge were attached to 60 calcium carbonate blocks of known weight. Three sets of 10 blocks were grown at high light levels and three sets of 10 blocks were grown at low light levels for 105 d in the Florida Keys, Florida, USA. Boring rates, growth rates (lateral growth and within-substratum tissue penetration), and zooxanthella populations were measured at the end of the experiment. Absolute rates of boring and growth of A. varians forma varians were significantly greater when zooxanthella densities were higher. Boring rate and tissue penetration related to final surface area of sponge attachment was also enhanced when zooxanthella densities were higher, suggesting that the symbiont plays a physiological role in the decalcification process. This is in contrast to the role that zooxanthellae play in coral hosts. Based on the results of this study, it appears that the presence of zooxanthellar symbionts has important ecological and life-history consequences for host sponges. Ability to laterally overgrow competitors will be correlated with the size and activity of zooxanthella populations. In addition, the fitness of host sponges will be enhanced by algal symbionts, since greater penetration within substrata will result in an increase in production of tissue that can be converted into storage, feeding and reproductive functions.     

Distribution of encrusting bryozoans and other epifauna on the subtidal bivalveChlamys opercularis

The shells of living queen scallops,Chlamys opercularis (L.), are often completely colonised with epifauna. In this study, the distribution of the major epifaunal groups across the surfaces of left (upper) and right (lower) valves is examined using samples from two commercial queen scallop beds off the Isle of Man. The composition of the epifauna differs across and between the valves and also varies with the age of the scallop. YoungerC. opercularis are commonly encrusted by sponges, but on older shells other epifauna colonise the surfaces. The encrusting bryozoan assemblage found on livingC. opercularis is not as abundant or speciose as on some other substrata such as stones and dead shell. This may be a result of intense competition from other phyla. The distribution of different bryozoan species between the surfaces is examined. Most species are more abundant on the lower valves.     

Spatial–temporal variability of epibenthic assemblages on subtidal biogenic reefs in the northern Adriatic Sea

Spatial distribution and temporal variation of epibenthic assemblages of coralligenous biogenic rocky outcrops occurring in the northern Adriatic Sea (45°04′–45°24′N; 12°23′–12°43′E) were investigated by photographic sampling from 2003 to 2006 at 12 randomly selected sites. The dominant reef-forming organisms were the encrusting calcareous algae (Lithophyllum stictaeforme, Lithothamnion minervae and Peyssonnelia polymorpha), while the main bioeroders were boring sponges (Cliona viridis, C. celata, C. thoosina, C. rhodensis, Piona vastifica) and the bivalve Gastrochaena dubia. Composition of the assemblages varied thorough years and among sites. Spatial heterogeneity, at local and regional scale, prevailed over temporal variation. This variability was related both to the geo-morphological features of the outcrops and to environmental variables. Sites clearly differed in the percent cover of reef builder and bioeroder species while only limited temporal variation within site was found. Some taxa revealed complex intra-site temporal trends. These results provide valuable information on the diversity and variability of epibenthic assemblages of the northern Adriatic coralligenous reefs, essential for the management and conservation of these unique biogenic habitats.     

Shell-size selection by intertidal sympatric hermit crabs

This study evaluated selection for shell size by three species of tropical intertidal hermit crabs, Clibanarius antillensis, C. sclopetarius, and C. vittatus, from species of shells which are frequently used in nature. Crab size and weight were strongly and significantly related to all measured parameters of the selected shells. The strength of these relationships (r² values) depended neither on the crab nor on the shell variables taken into account. The relationships between crab size and the dimensions of the selected shells showed higher r² values than the corresponding relationships with the shells that the crabs had occupied when they were collected (0.482–0.903 in comparison to 0.091–0.652, respectively), indicating that the crabs were occupying sub-optimal shells in nature. Negative allometry was frequently found in the relationships between crab and shell variables, indicating that large crabs select and use proportionally lighter shells than do small crabs. This negative allometry was stronger for the shells used in nature (except for C. antillensis), i.e. larger crabs tended to select heavier shells in the laboratory than in nature. Different allometric relationships were also recorded among the dimensions of shells used in nature and those selected by the hermit crabs in free-access experiments: as shell length increased, the selected shells were heavier and had larger apertures than the shells used in nature. The relationships between crab size and the length and weight of the selected shells did not depend on the species of crab or species of shell, but only on crab size. Therefore, analyses using these variables can be performed without taking the species of crab or shell into account, i.e. data from different crab or shell species can be pooled. The influence of crab and/or shell species was recorded only in the models fitted for aperture length and width, variables which were more related to shell architecture than did shell length or weight. In contrast, if crab weight is used as an independent variable, different crab or shell species can be analyzed together independently of the particular shell parameter. This indicates that crab weight may be less susceptible than crab shield length to shell morphological constraints. Finally, the results indicate that the preferred size of a given shell type chosen by a given hermit crab will depend more on crab size or weight, than on the crab or shell species under consideration, i.e. crab shell-size relationships are not species specific.Communicated by P.W. Sammarco, Chauvin     

Growth in the bivalve Yoldia eightsi at Signy Island,Antarctica, determined from internal shell increments and calcium-45 incorporation

The growth rate of the infaunal nuculanid bivalve Yoldia eightsi at Factory Cove, Signy Island, South Orkney Islands (maritime Antarctica), was estimated from internal shell increments and ⁴⁵Ca incorporation of individuals collected monthly from December 1987 to April 1989. Acetate peels of etched shells revealed clear first-order increments, with less well defined, narrower, second-and third-order increments. The first-order increments were assumed to be annual, although there is no independent confirmation of this assumption. Unfortunately abrasion of the umbo region and the small thin shells of Y. eightsi meant that in no case could a complete sequence of increments be measured realiably on any individual shell. Measurements of 1043 first-order increments from 130 shells where a minimum of two consecutive increments could be detected were therefore pooled, and a population growth curve constructed from a Ford-Walford plot. This indicated a slow growth rate, with a maximum shell height of 22.3 mm (equivalent to a shell length of 35.6 mm) being reached at an age >60 yr. The size-frequency distribution of 1521 individuals pooled from winter (July to October) samples revealed a distinct lack of smaller (younger) individuals, possibly reflecting poor recruitment in areas of dense adult populations. The largest shell recovered in the samples was 33.5 mm in length, with an estimated age of 52 yr. Short-term ⁴⁵Ca-incorporation experiments indicated a mean daily rate of growth increment of 3.8 m for individuals of 12 mm shell height, which matches the proposed annual growth rate if growth is assumed to occur for about 150 d each year and the first-order increments are assumed to be annual.     

The invasion and subsequent die-off of <Emphasis Type="Italic">Mytilus galloprovincialis</Emphasis> in Langebaan Lagoon,South Africa: effects on natural communities

The alien mussel Mytilus galloprovincialis invaded sand banks in Langebaan Lagoon on the west coast of South Africa in the mid-1990s. However, by 2001 these beds had completely died off, with only empty shells and anoxic sand remaining. In an effort to prevent the re-settlement of this aggressive invader, all dead mussel shells were then cleared. This study considered the impacts of the invasion and subsequent die-off on natural benthic communities. Community composition differed significantly between non-invaded and invaded areas (ANOSIM, R = 0.685 and P < 0.01) as the physical presence of mussel beds created a new habitat that promoted invasion by indigenous rocky-shore species. This dramatically increased faunal biomass from 1,132.9 g m⁻² ± 3,454.7 SD to 53,262.4 g m⁻² ± 23,052.6 SD and species richness from 38 to 49 species. Following the die-off of the mussel beds, communities remained significantly different between non-invaded areas and those in which mussel shells remained (ANOSIM, R = 0.663 and P < 0.01). Species richness was significantly greater in non-invaded areas (18 species) than in uncleared areas with remnant shells (four species) (Kruskal–Wallis ANOVA H _2,36 = 10.8964 and P = 0.032), as the previously dominant rocky-shore species became smothered by sediment and the compacted shells formed an impermeable layer excluding sandy-shore burrowing organisms. After the shells were cleared, 50% of the sandy-shore species associated with non-invaded areas returned within 5 months, but community structure still remained significantly different to non-invaded areas (ANOSIM, R = 0.235 and P > 0.05). Invasion thus dramatically altered natural communities and although the subsequent removal of the dead mussel shells appears to have aided recovery, community composition remained different from the pre-invasion state after 5 months.     

Predaceous gastropods regulate new-shell supply to salt marsh hermit crabs

Marsh hermit crabsPagurus longicarpus Say directly acquire new shells as the predatory gastropodMelongena corona Gmelin consumes marsh periwinkles,Littorina irrorata Say. The influx rate of new shells into a salt marsh hermit crab population was measured by marking live periwinkles and daily recovering the shells from hermit crabs over periods of 3 to 6 d. Average rates of new shell acquisition ranged from 4.0 to 23.3 new shells per day from salt marsh areas of 4×10 m. Such consistently high rates contrast with the negligible rates generally assumed for new shell entry into hermit crab populations. The number of new shells acquired each day varied directly with the number of the predatory gastropod,M. corona, present in each study area at both natural and manipulated predator densities. Empty shells on the substrate are usually considered as the primary source of new shells to hermit crabs. However, over 500 empty shells had to be placed on the substrate in a 4×10 m area to provide a daily rate of 20 new shells to the hermit crab population.This is the first in a new contribution series from the Florida State University Marine Laboratory No. 1001     

Ultrastructural analysis of dissolution of shell of the bivalve Mytilus edulis by the accessory boring organ of the gastropod Urosalpinx cinerea

The shell penetrating mechanism of the muricid gastropod Urosalpinx cinerea follyensis Baker was examined with reference to solubilizing effects of the secretion of the accessory boring organ (ABO) on the ultrastructural organic and mineral components of the shell of the bivalve Mytilus edulis Linné. The fine structure of shell etched by the secretion was contrasted with normal shell and shell solubilized artificially by acids, a chelating agent, and enzymes as an aid in interpreting the pattern etched by the secretion. A synoptic series of scanning electron micrographs of representative regions of the normal shell of M. edulis was prepared to serve as a standard for ultrastructural interpretation of the patterns of dissolution. Intercrystalline conchiolin of the mosaicostracal, prismatic, myostracal, and nacreous strata was dissolved as readily as the periostracum by the ABO secretion. In the prismatic region, maximum depth of dissolution of intercrystalline organic matrix occurred when long axes of prisms were approximately perpendicular to the surface being dissolved. Microscopic solubilization of organic matrix noticeably preceded dissolution of mineral crystals, revealing subsurface prisms and lamellae similar in size and form to the distinctively shaped normal shell units. After solubilization of intercrystalline conchiolin, further dissolution by the ABO secretion revealed a variety of what appeared like internal structures in prisms and lamellae. The form of these subunits varied from that of platelets to nodules in prisms and from laths to nodules in lamellae. These intracrystalline patterns of dissolution probably resulted from preferential etching of (a) soluble intracrystalline conchiolin membranes or other internal aggregates of nacrin, (b) heterogeneously distributed trace and minor elements, or (c) from both. Carriker and Williams (1978) hypothesized that a combination of HCl, chelating agents, and enzymes in a hypertonic mucoid secretion released by the ABO dissolve shell during hole boring. The similarity of patterns of dissolution etched by the ABO secretion and those produced artificially by HCl and EDTA in the present study support the hypothesis that these chemicals, or chemicals similar to them, are constituents of the ABO secretion. Lactic and succinic acids and a chitinase-like enzyme were also suggested by Carriker and Williams as possible agents in shell dissolution. Alteration of the shell surface by experimental application of these, and other, chemical agents was not sufficiently comparable to that etched by the ABO secretion to support the suggestion. Preferential dissolution of shell matrix by the ABO secretion is functionally advantageous to boring gastropods because it increases the surface area of mineral crystals exposed to solubilization and facilitates removal of shell units from the surface of the borehole by the radula.     

Epibiosis on cerith shells in a seagrass bed: correlation of shell occupant with epizoite distribution and abundance

The cerith Cerithium atratum (Born 1778) is an abundant gastropod in the seagrass beds at Cabo Frio, Brazil. In order to estimate the ecological importance of cerith shells as a rare hard substratum in the seagrass bed, the abundances of C. atratum and of cerith shells occupied by hermit crabs were quantified. The mean densities of C. atratum and hermit crabs were 1887 and 100 individuals m⁻², respectively, and these provided 0.5 m² shell area m⁻² available for epizoite colonization. The tube-forming polychaete Hydroides plateni (Kinberg 1867) and oyster Ostrea puelchana Orbigny, 1841 were the dominant visible epizoites on inhabited cerith shells. These epizoite populations were compared in order to investigate whether the temporal and spatial patterns in the epibiotic community were related to ecological and behavioral aspects of the occupant species (cerith or hermit crab). Larger cerith shells had a greater abundance of epizoites. Each epizoite showed a preference for a different occupant of the shells (the oyster for C. atratum and the polychaete for cerith shells occupied by hermit crabs). The oyster showed a seasonal pattern in abundance on C. atratum, being more common in fall (March–April). The distribution of the epizoites on the shells depended on the shell occupant species and was probably related to their different foraging activity –C. atratum ploughs half buried through the sediment, while the hermit crab crawls on the sediment surface. In both cases, the activity of the shell occupant was considered to be beneficial to the epizoites, as empty shells and shell fragments did not support a macroepifauna. Received: 1 May 2000 / Accepted: 8 August 2000     

Life history characteristics of a coral reef sponge

Study of the life history characteristics of the common Red Sea sponge Mycale fistulifera (Poecilosclerida: Demospongiae) reveals several traits which may categorize it as an opportunistic species: (1) at least part of the population is reproductively active throughout the year, providing a continuous supply of larvae for settlement; (2) sponges may produce and release a large number (152±39 larvae d^-1) of brooded larvae; (3) released larvae are capable of fast settlement and metamorphosis (minutes to 30 hours post-release); (4) the turnover of the population is high and population size varies with time. From 48 initial sponges plus 94 recruits, only 25 remained alive after 14 months of observation; (5) most members of the population (>70%) have a small body size (<30 cm²); (6) sexual maturity may occur at an early age. These traits facilitate continuous establishment of M. fistulifera in new spaces on the reef; (7) M. fistulifera, preferred substrate is another opportunistic species, the coral Stylophora pistillata which it overgrows.     

Shell choice in<Emphasis Type="Italic"> Pagurus longicarpus</Emphasis> hermit crabs: does predation threat influence shell selection behavior?

Pagurus longicarpus hermit crabs depend on empty gastropod shells for protection against predation. Hermit crabs avoid gastropod shells in which holes have been drilled by naticid gastropods, and hermit crabs forced to occupy drilled shells are more vulnerable to predation by green crabs, Carcinus maenas. In this study, we examined the effect of predator cues on P. longicarpus shell investigation behavior and shell choice. In paired laboratory shell choice trials, we examined hermit crab response to green crab chemical cues. We compared hermit crabs from two sites differing in the percentage of Littorina littorea shells with drill holes. The percentage of time hermit crabs spent occupying intact shells increased significantly in the presence of predator cues. The effect of predator cues on the amount of time hermit crabs spent investigating shells differed between individuals from the two sites. Predator effluent had a marginal effect on the proportion of hermit crabs initially choosing intact shells and within 15 min most hermit crabs in both treatments occupied intact shells due to shell switching. These results indicate that predation cues alter P. longicarpus shell choice behavior favoring intact shells, which provide greater protection. In summary, predation appears to be a key factor influencing hermit crab shell selection behavior.Communicated by T. Czeschlik     

Obligate commensalism of <Emphasis Type="Italic">Curvemysella paula</Emphasis> (Bivalvia: Galeommatidae) with hermit crabs

Curvemysella paula is a markedly crescent-shaped bivalve that lives inside snail shells occupied by hermit crabs. Here, we describe the unique symbiotic life, growth pattern, and reproductive biology of this bivalve, based on specimens collected from the shallow, muddy bottom of the Seto Inland Sea, Japan. C. paula was found attached to columellae in the siphonal canal, mainly of nassariid snail shells occupied by two types of hermit crabs: Diogenes edwardsii (Diogenidae) and Spiropagurus spiriger (Paguridae). The crescent-shaped shell of C. paula is an adaptation to symbiotic life in the narrow interspace between the snail shell and the hermit-crab abdomen. C. paula is a protandric hermaphrodite. In our samples, each host snail shell harbored one (or rarely a few) large female and several males. All the female bivalves settled on the host shells with their anterior end facing outward and benefited from currents created by the hermit crab when feeding. In the muddy bottom, snail shells are a limited resource for both the hermit crabs and symbiotic bivalves. The bivalves benefit from the mobility of the hermit crabs, which prevent the shells from becoming buried in the mud. C. paula represents the only example of obligate commensalism with hermit crabs found in Bivalvia.     

Evidence of nitrification and denitrification in high and low microbial abundance sponges

Aerobic and anaerobic microbial key processes were quantified and compared to microbial numbers and morphological structure in Mediterranean sponges. Direct counts on histological sections stained with DAPI showed that sponges with high microbial abundances (HMA sponges) have a denser morphological structure with a reduced aquiferous system compared to low microbial abundance (LMA) sponges. In Dysidea avara, the LMA sponge, rates of nitrification and denitrification were higher than in the HMA sponge Chondrosia reniformis, while anaerobic ammonium oxidation and sulfate reduction were below detection in both species. This study shows that LMA sponges may host physiologically similar microbes with comparable or even higher metabolic rates than HMA sponges, and that anaerobic processes such as denitrification can be found both in HMA and LMA sponges. A higher concentration of microorganisms in the mesohyl of HMA compared to LMA sponges may indicate a stronger retention of and, hence, a possible benefit from associated microbes.     

Age and growth in <Emphasis Type="Italic">Odontocymbiola magellanica</Emphasis> (Gastropoda: Volutidae) from Golfo Nuevo,Patagonia, Argentina

Growth, age and somatic production of the benthic predator Odontocymbiola magellanica were studied in Golfo Nuevo (42°S 65°W), on the South American Atlantic shelf. Stable oxygen isotope ratios confirmed semiannual formation of internal and external shell growth marks. Mean shell length (SL) of females was 115 and 112 mm for males, while population modal shell-free wet mass (SFWM) was 62.8 g. A Gompertz growth function (SL_∞ = 200 mm, K = 0.197, t ₀ = 5.486) fitted 113 pairs of size-at-age data (12 shells) best. O. magellanica is a long-lived species, reaching up to 20 years of age. The maximum individual somatic production of 29.3 g SFWM per year is attained at 145 mm SL, which corresponds to about 12 years of age. The life span of this volutid seems to be twice compared with other large gastropods. O. magellanica is a valuable and exploitable resource regarding its large size and somatic production, but on the other hand, its slow growth, late maturity and direct development makes it extremely vulnerable to overexploitation.     

Habitat structure,population abundance and the opportunity for selection on body weight in the amphipodEogammarus oclairi

The importance of oyster-shell habitat-characteristics (depth of the shell layer or degree of fragmentation) to the amphipodEogammarus oclairi were studied in Grays Harbor estuary (Washington, USA) in regard to their effect on local density ofE. oclairi, the risk of predation by juvenile Dungeness crabsCancer magister, and the opportunity for selection on body weight during pairing.E. oclairi was the most abundant macrofaunal species in the intertidal oyster-shell assemblages (density range = 20 to 8500 amphipods m^–2), and its density was positively correlated with the depth of the oyster-shell layer (r = 0.85,n = 30,p < 0.005). Field experiments showed that amphipod density was much lower when oyster shells were whole (x = 340.7 ± 72.1,n = 10) as opposed to fragmented [x = 41.4 ± 9.3,n = 10;t _c (Welch's approximatet-test) = 13.0,df = 9.3,p < 0.05]. Densities of a predator, juvenileC. magister, were not affected by depth of the shell layer nor by degree of fragmentation (whole shell,x = 14.6 ± 13.0 crab;fragmented shell,x = 18.7 ± 2.4 crab;t _c = 0.83,df = 9.7,p > 0.05). Predation rate on single amphipods by crabs did not differ between habitat types (whole shells vs fragmented shells), but mating pairs were consumed more often in the whole-shell treatment (whole shell,x = 0.9 ± 0.8, fragmented shell,x = 0.1 ± 0.3). There were no differences in the size of single amphipods (both sexes) consumed between treatments, or in the size of the paired males consumed. Field experiments showed that the opportunity for selection (i) on male body weight increased with increasing amphipod density which, in turn, increased with increasing degree of shell fragmentation (whole-shell treatment,i = 0.0014 ± 0.0002, fragmented-shell treatment,i = 0.3756 ± 0.0338). Large spatial and temporal fluctuations in population abundance complicate the evaluation of the importance of selection in determining traits such as body weight.