Age,growth, and mortality of the prosobranch<Emphasis Type="Italic"> Zidona dufresnei</Emphasis> (Donovan, 1823) in the Mar del Plata area,south-western Atlantic Ocean

The volutid snail Zidona dufresnei is a benthic top predator in the Mar del Plata (Argentina) shelf area where it was subjected to unregulated commercial exploitation for more than 20 years. So far there is no stock management, and hitherto even the most basic information on population dynamics of this species is missing. Annual formation of internal shell growth bands visible by x-ray was confirmed by the stable oxygen isotope record in the shell carbonate that reflects seasonal oscillations in water temperature. A Gompertz growth function ( , K=0.211 year^–1, t₀=5.496) fitted 142 pairs of size-at-age data (30 shells) best. Maximum individual production amounted to 26.8 g shell-free wet mass (SFWM) at 115 mm shell length. Based on a size-frequency distribution derived from commercial catches, annual mortality rate of Z. dufresnei was estimated to be 0.61 (±0.21) year^–1.Communicated by O. Kinne, Oldendorf/Luhe     

Patterns of selective predation by juvenile,benthivorous fish on estuarine macrofauna

Benthic feeding on macrofauna was studied in juveniles of the sparids Lithognathus lithognathus and Rhabdosargus holubi in the upper reaches of the Gamtoos Estuary, South Africa. Fish and benthic macrofauna were sampled simultaneously, and the selection of invertebrate prey assessed. Both fish species strongly selected for corophioid amphipods and consumed other benthic taxa in low numbers. R. holubi also exploited aquatic autotrophs, while L. lithognathus had a narrow prey-spectrum, feeding almost exclusively on the tube-dwelling amphipod Grandidierella lignorum. G. lignorum was the most abundant prey species, both in the benthos and the fish's diet. This species also showed prominent behavioural differences between the sexes; males were markedly more active on the sediment surface than females, who rarely left their tubes during the day. Males switched from an infaunal to epifaunal microhabitat in search of receptive females, concurrently increasing their exposure to fish predators. Consequently, L. lithognathus selected significantly more males than female amphipods, causing a marked bias towards females in the sex ratio and age-structure of the amphipod population. Juvenile amphipods were less preyed upon, presumably as a result of lower prey-detection or capture efficiency by the predators. Accepting current notions about predation as an important structuring element for benthic communities, our data also stress the prominence of size-and sex-selective predation in structuring individual prey populations.     

The effects of starvation and acute low salinity exposure on food intake in the Dungeness crab, Cancer magister

Adult Cancer magister make forays into hyposaline estuarine habitats during times of high food abundance. However, as weak osmoregulators, they are poorly equipped to deal with the concurrent demands of osmoregulation and digestion. Therefore, the potential interaction between nutritional status and feeding in a physiologically challenging environment was investigated. Changes in the proportion of crabs feeding, the amount of food consumed, the time spent feeding, and the efficiency with which a meal was consumed were examined in response to the length and severity of hyposaline exposure, and the duration of starvation. Reductions in the (a) number of animals feeding, (b) the amount of food consumed, and (c) the time spent feeding were observed in salinities where C. magister actively osmoregulates the concentration of its internal fluids. Although this reduction in feeding was likely a stress response, the crabs were able evaluate the level of salinity stress: there was a dose-dependent reduction in feeding, and they were able to discriminate between salinities separated by as little as 3.5‰. The likelihood that animals would feed in low salinity increased with starvation. Thus, the aversion to food uptake in physiologically stressful conditions may be overridden by the need to procure nutrients. In the natural environment, we suggest that C. magister are employing an ‘eat and run’ strategy, moving into the estuary, consuming a meal, and retreating to higher salinities to digest.     

Impact of a burrowing deposit-feeder, Monoporeia affinis, on viable zooplankton resting eggs in the northern Baltic Sea

We investigated the effect of different densities of the burrowing deposit-feeding amphipod Monoporeia affinis on the potential for recruitment of zooplankton from benthic resting eggs. Hatching of resting eggs was induced in the laboratory on sliced and resuspended 1-cm depth-sections of sediment cores, collected at six stations in an archipelago area of the Gulf of Bothnia, Baltic Sea. The uppermost 5 cm of the sediment was studied. The most common species that hatched was Eurytemora affinis (Copepoda). Individuals from another copepod genus, Acartia, hatched in significant numbers only in the cores from two stations with low amphipod abundance. Cores from stations with high amphipod densities showed a deeper distribution of emerging E. affinis nauplii compared with stations with few amphipods; the oxidised sediment layer was also deeper at high M. affinis densities than at low. Total (0 to 5 cm strata pooled) number of hatched E. affinis nauplii was independent of amphipod density. This indicates that the effect of M. affinis on E. affinis eggs involves deeper burial due to bioturbation, rather than predation. Decreased benthic recruitment of zooplankton at localities with high M. affinis density is suggested, since more deeply positioned eggs are less likely to hatch. When hatching was induced in intact, non-sliced cores from one station, the number of E. affinis nauplii that hatched was on average 43% of the number that hatched in the upper centimetre of the sliced cores from the same station. This fraction (43%), if applied to the other stations, implied a potential for benthic recruitment of up to 80 000 ind m⁻² for E. affinis. Due to its high abundance, M. affinis is likely to greatly reduce benthic recruitment of zooplankton in this system. Received: 17 September 1999 / Accepted: 23 February 2000     

Examples of post-mortality alteration in Recent brachiopod shells and (paleo)ecological consequences

Post-mortality alteration of brachiopod shells under normal environmental conditions leads to high taphonomic loss, and to a poor contribution to the biodetrital soft sediment. The successive stages of alteration which shells undergo are: (bio)degradation of the organic matrix shell softening structural disaggregation or/and mechanical fragmentation; these processes depend on the shell structure (number of layers) and composition (organic and inorganic components), but very little on environmental conditions, except for the dissolution of inorganic shell constituents. Among the Brachiopoda, three types of alteration occur to different types of shells — Type I: two-layered chitino-phosphatic shell (species ofLingula andGlottidia) displays a rapid degradation of the organic matrix and mechanical abrasion, leading to total disappearance of the shell in 2 to 3 wk; Type II: two-layered carbonate shell, e.g.Terebratulina spp. [but notNotosaria nigricans (Sowerby) which may constitute a fourth shell type], exhibits degradation of the organic matrix of the secondary layer, shell softening, and structural disaggregation leading to shell disintegration in 6 to 7 mo with a concomitant contribution of calcitic microfibres to the sediment; Type III: three-layered carbonate shell, e.g.Gryphus vitreus (Born), undergoes organic degradation of the secondary layer, fragmentation of the anterior two-thirds of the shell, and slow degradation (because of the thick tertiary layer) of the posterior portion of the shell, with dissolution of the inorganic components (mainly in the tertiary layer) which make a relatively minor contribution to the sediment.     

The role of different types of detached macrophytes in the food and habitat choice of a surf-zone inhabiting amphipod

Allorchestes compressa is the dominant macroinvertebrate species in wrack accumulations on surf zones of south-western Australia. These amphipods were provided with a choice of macrophyte material representing brown and red algae and seagrass in a series of preference experiments in the laboratory and field. Feeding experiments showed that A. compressa exhibited a strong preference for particular types of macrophytes (P < 0.01). Amphipods primarily consumed brown algae, with 69–98% of the biomass of Ecklonia radiata and 64% of the biomass of Sargassum sp. lost over the experiments. This study has shown that the amphipod A. compressa exhibits a clear preference for brown algae over red algae and seagrass as food. In terms of habitat preference, tank experiments using a series of pair-wise comparisons showed that, in the absence of fish predators, A. compressa selected seagrass as its preferred habitat over the other types of wrack (P < 0.001). When satiated or starved, between 68 and 83 and 79 and 98% of amphipods were found in Amphibolis and Posidonia, respectively. In contrast, field-cage experiments revealed that A. compressa preferred either mixed wrack, brown algae or red algae over seagrass as a habitat (P < 0.01). The contrasts between results from the laboratory and field suggest that other factors such as the presence of predators, water flow and light could influence habitat choice in the surf zone. This study shows that allochthonous material transported to surf zones from other habitats therefore play different roles in driving secondary production in this shoreline habitat.     

Incidental capture,direct mortality and delayed mortality of sea turtles in Australia's Northern Prawn Fishery

The species composition, catch and mortality rates of sea turtles captured incidentally by the tiger prawn fishery on Australia's northern coast in 1989 and 1990 were estimated by monitoring the fishery's catch. In 1990, the delayed rate of mortality from damage was estimated and the size composition was measured. Five species of turtles were captured: the flatback (Natator depressa, 59% of the total), loggerhead (Caretta caretta, 10%), olive ridley (Lepidochelys olivacea, 12%), green turtle (Chelonia mydas, 8%) and hawksbill (Eretmochelys imbricata, 5%). The turtle catches varied with water depth: the highest catch rates (0.068±0.006 turtles per trawl) were from trawls in water between 20 and 30 m deep, relatively few turtles (10%) were captured in water deeper than 40 m (25% of trawls). Catch rates varied with time of year: the highest catch rates were 0.098 (±0.013) turtles per trawl in winter. There was no significant difference in the overall catch rate (²= 0.047; p=0.8111; df=1) but a significant difference in mortality rate (²= 3.99; p<0.05; df=1) between the two years. The incidence of capture in the commercial fishery was 0.051 (±0.003) turtles per trawl towed for about 180 min, with 0.007 (±0.001) turtles per trawl drowning in the nets. There were no significant differences in the catch and mortality rates between the two years for any of the turtle species except the loggerhead, which had a significantly (² = 11.029; p=0.0013; df=1) lower catch rate in 1990 (0.002±0.001 turtles per trawl) than in 1989 (0.008±0.002 turtles per trawl), and a significantly higher mortality in 1990 (33%) than in 1989 (19%). Catch rates and mortality varied between the species: the flatback had the highest catch rate (0.030±0.002 turtles per trawl) but the lowest mortality (10.9%); the loggerhead had a catch rate of 0.005±0.001 turtles per trawl, and high mortality (21.9%); the olive ridley had a catch rate of 0.006±0.001 turtles per trawl and a low mortality (12.5%); the green turtle's catch rate was 0.004±0.001 per trawl and mortality 12.0%; the hawksbill had the lowest catch rate (0.002±0.001 turtles per trawl) but highest mortality (26.4%). Based on the fishing effort (27 049 d for 1989 and 25 746 d for 1990), we estimate that 5 503 (±424) turtles were caught and returned to the sea in 1989 and 5 238 (±404) in 1990, of which 567±140 drowned in 1989 and 943±187 in 1990. In 1990, an estimated 25% of all captured turtles suffered some non-lethal damage; an estimated 21% of turltes were captured comatose and 4% were injured. We conclude that, considering other threats, trawl-induced drowning is not the major impact on turtle populations in northern Australia, but that measures to reduce drowning and delayed mortality would be desirable.     

Feeding habits of a pelagic amphipod,Themisto japonica

Vertical distribution, chlorophylla (chla) and phaeopigment concentrations in the gut, and natural nitrogen isotope ratio ( ¹⁵N) were investigated for pelagic amphipodsThemisto japonica (Bovallius) collected from the Sea of Japan in July 1987. Differences in diel vertical migration behavior were clearly observed between small and largeT. japonica. Many small (<5 mm body length) amphipods appeared in the phytoplankton-rich shallow layers. Their gut pigment concentrations were higher (mean 0.52 ± 0.15µg chla g^–1 amphipod) than those of large amphipods (mean 0.33±0.14µg g^–1); this implies that the amphipods fed on a large amount of phytoplankton during the early stage of life. The ¹⁵N values of small amphipods were lower (5.7 to 6.3) than those of large amphipods (6.8 to 11.7), reflecting the lower trophic level of small amphipods compared to large ones. The ¹⁵N values for small amphipods were similar to those of herbivorous zooplankton. The amphipods' feeding behavior thus changes from herbivorous to carnivorous as they grow.     

Population biology of Dyopedos monacanthus (Crustacea: Amphipoda) on estuarine soft-bottoms: importance of extended parental care and pelagic movements

The suspension-feeding amphipod Dyopedos monacanthus (Metzger, 1875) is a common epibenthic amphipod that lives on self-constructed “mud whips” (built from filamentous algae, detritus and sediment particles) in estuaries of the northern North Atlantic Ocean. The population biology of D. monacanthus at a shallow subtidal site in the Damariscotta River Estuary (Maine, USA) was examined between July 1995 and July 1997. The resident population at the study site was dominated by adult females during most months of the year. High percentages of subadults were found in late summer/early fall. Often, between 10 and 20% of the adult females were paired with males, and the percentage of ovigerous females varied between 40 and 100%, indicating continuous reproduction. The percentage of parental females varied between 40 and 80% during most months, but dropped to levels below 20% during summer/early fall. The average size of amphipods on their own mud whips was ∼4 mm during the summer/early fall, after which it increased continuously to >7.0 mm in March or April, and then dropped again. In March and April, the average number of eggs and juveniles female⁻¹ was ∼100 eggs and 55 juveniles, while during the summer/early fall the average number of eggs female⁻¹ was <20 and that of juveniles female⁻¹ was <10. Many juveniles grew to large sizes (>1.4 mm) on their mothers' whips in winter/early spring but not in the summer/fall. The average number of amphipods at the study site was low in late summer/early fall (<50 individuals m⁻²), increased steadily during the winter, and reached peak densities of >3000 individuals m⁻² in April 1996 (>1600 individuals m⁻² in May 1997), after which densities decreased again. The decrease of the D.␣monacanthus population at the study site coincided with a strong increase of amphipods found pelagic in the water column. This behavioural shift occurred when temperatures increased and benthic predators became more abundant and active on shallow soft-bottoms, suggesting that D. monacanthus at the study site is strongly affected by predation. The effects are direct (by predation on amphipods) and indirect (by reducing duration of extended parental care and enhancing pelagic movements). Both extended parental care and pelagic movements are important behavioural traits of D.␣monacanthus (and other marine amphipods), and significantly affect its population dynamics. Received: 18 January 1998 / Accepted: 27 May 1998     

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     

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.     

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     

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     

Grazing rates of organic matter and living fungal biomass of decaying Spartina alterniflora by three species of salt-marsh invertebrates

 The pathway for the flow of salt-marsh grass production into marsh food-webs is still not well defined. We compared the abilities of three marsh macroinvertebrates [salt marsh periwinkles, Littoraria irrorata (Say) (=Littorina irrorata), salt-marsh coffee-bean snails, Melampus bidentatus (Say); and a talitrid amphipod, Uhlorchestia spartinophila Bounsfield and Heard] to access standing-dead leaves of smooth cordgrass (Spartina alterniflora Loisel). The invertebrates were incubated with naturally-decaying leaves, and the rates of removal of organic matter and living fungal biomass (ergosterol) were measured. The impact of invertebrate activity upon fungal growth rates was measured as rates of fungal-membrane synthesis (incorporation of radioacetate into ergosterol). The removal rates of organic leaf biomass per mg individual biomass were highest for amphipods (700 μg mg⁻¹ d⁻¹) and lowest for periwinkles (90 μg mg⁻¹ d⁻¹), but the relatively large biomass of the snails made their removal rates per individual greater than those of amphipods. Net removal of ergosterol by all three invertebrates was >50% for yellow-brown (early-decay) leaf blades. For fully-brown (advanced-decay) blades, >50% removal of ergosterol was found only for periwinkles; exposure to coffee-bean snails and amphipods resulted in a net ergosterol reduction of ≤20%. The lower net reduction of living fungal biomass by coffee-bean snails and amphipods may have been due to fungal-growth stimulation (2.3-fold stimulation in coffee-bean snails and 1.5-fold stimulation in amphipods). Grazing by periwinkles did not stimulate fungal growth, possibly because of its high intensity. Grazing by these three salt-marsh shredders may affect marsh-grass shoot-decay in different ways. Periwinkles may abbreviate the period of fungal production, and incorporate the decaying material relatively quickly into snail biomass and fecal-pellet rain to the sediments. Coffee-bean snails and amphipods may enhance and prolong fungal production, along with the formation of fecal-pellet rain. All three invertebrates fed preferentially on leaf blades rather than leaf sheaths, and feeding rates of gastropods were higher during the night than during the day. Received: 25 November 1998 / Accepted: 4 November 1999     

Fish Responses to Experimental Fragmentation of Seagrass Habitat

Abstract: Understanding the consequences of habitat fragmentation has come mostly from comparisons of patchy and continuous habitats. Because fragmentation is a process, it is most accurately studied by actively fragmenting large patches into multiple smaller patches. We fragmented artificial seagrass habitats and evaluated the impacts of fragmentation on fish abundance and species richness over time (1 day, 1 week, 1 month). Fish assemblages were compared among 4 treatments: control (single, continuous 9‐m² patches); fragmented (single, continuous 9‐m² patches fragmented to 4 discrete 1‐m² patches); prefragmented/patchy (4 discrete 1‐m² patches with the same arrangement as fragmented); and disturbance control (fragmented then immediately restored to continuous 9‐m² patches). Patchy seagrass had lower species richness than actively fragmented seagrass (up to 39% fewer species after 1 week), but species richness in fragmented treatments was similar to controls. Total fish abundance did not vary among treatments and therefore was unaffected by fragmentation, patchiness, or disturbance caused during fragmentation. Patterns in species richness and abundance were consistent 1 day, 1 week, and 1 month after fragmentation. The expected decrease in fish abundance from reduced total seagrass area in fragmented and patchy seagrass appeared to be offset by greater fish density per unit area of seagrass. If fish prefer to live at edges, then the effects of seagrass habitat loss on fish abundance may have been offset by the increase (25%) in seagrass perimeter in fragmented and patchy treatments. Possibly there is some threshold of seagrass patch connectivity below which fish abundances cannot be maintained. The immediate responses of fish to experimental habitat fragmentation provided insights beyond those possible from comparisons of continuous and historically patchy habitat.     

Compressibility and shell failure in the European Atlantic <Emphasis Type="Italic">Patella</Emphasis> limpets

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

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     

Population biology and life strategies of Chlorophthalmus agassizii Bonaparte, 1840 (Pisces: Osteichthyes) in the Mediterranean Sea

The population biology and life strategies of Chlorophthalmus agassizii were studied in the Ionian Sea (eastern–central Mediterranean) using the data collected during the experimental trawl surveys carried out from 1995 to 2000. Depth-related trends of both density and size were found. With depth the former decreased while the latter increased. A typical bigger–deeper phenomenon was detected: young-of-the-year individuals occur on the shelf during autumn–winter months and move towards bathyal bottoms as they grow. The sampled population was made up of several size–age groups. The maximum age of 8 years was identified by means of otolith readings. The Von Bertalanffy growth parameters were estimated from the age–length key (L _∞=189.04±5.401 mm; k=0.24±0.021; t _o=−1.20±0.132; φ′=3.94) and modal progression analysis (L _∞=218.33±18.397 mm; k=0.164±0.028; t _o=−1.694±0.171; φ′=3.89). Reproduction of this monoecious fish was observed during summer–early autumn. Considering the female portion of the gonad, the size at attainment of 50% maturity was 115 mm TL. The corresponding age is within the third year of life. The simultaneous occurrence of oocytes in different development stages was shown in the ovary. Both the asynchronous ovary and oocyte size distribution indicate that C. agassizii spawns more than once during a reproductive season (batch spawner). Functional fecundity (on average 3,018 hydrated oocytes) was between 37 and 69% of the absolute fecundity and increased significantly with the individual size. Since adult specimens are mostly distributed on the slope, eggs and larvae develop in epipelagic waters and migrate in-shore where juvenile forms recruit on the shelf. Juveniles migrate ontogenetically towards deeper bottoms and after 2–3 years start to reproduce annually within a life span greater than 10 years.     

Ontogeny of diet shifts by a microcarnivorous fish, Cheilodactylus spectabilis : relationship between feeding mechanics, microhabitat selection and growth

The mechanisms leading to ontogenetic shifts in prey selectivity are examined for the temperate microcarnivore Cheilodactylus spectabilis (Cheilodactylidae) in north-eastern New Zealand. These fish prey on invertebrates associated with benthic turf and foliose algae, using a suctorial feeding mode combined with oral sorting. All sizes of fish feed in the same shallow-water habitat using the same feeding mode. Dietary analysis revealed that while all sizes of fish consumed similar taxa, the relative proportion of taxa consumed reflected fish size. Juveniles consumed mainly gammarid amphipods while large adults targeted ophiuroids, with an abrupt shift from feeding predominantly on amphipods at 250 mm standard length. This dietary shift loosely coincided with the onset of sexual maturity and a change in growth trajectory, although dietary trends did not differ between sexes. Both juveniles and adults were found to select particular taxa from the available turf micro-fauna, with juveniles consuming smaller sizes of amphipods than adults. Microhabitat use was also found to change ontogenetically. Detailed observations on feeding mechanics suggested that size-related changes in suctorial force allowed the exploitation of a broader range of microhabitats with increasing fish size. Received: 20 June 1997 / Accepted: 1 April 1998     

Survival and bioturbation of the amphipod Monoporeia affinis in sulphide-rich sediments

Laboratory experiments were conducted to investigate the survival rate of Monoporeia affinis in sulphide-rich sediment with oxic overlying water, and the effect of amphipod bioturbation on sulphide and oxygen profiles. As long as the oxygen content in the water is high, the amphipods seem to avoid quite high concentrations (>200 μmol l⁻¹) of sulphide in the sediment by creating microhabitats where sulphide is rapidly oxidised. In cores with amphipods, a decrease of sulphide concentration was found in upper layers, while an increase of sulphide was found in deeper layers. Aggregation of amphipods generated pockets of light-brown sediment, characterised by high oxygen concentrations and no sulphide, and their depth was clearly dependent on amphipod density. This indicates that M. affinis has a potential to recolonise sulphide-rich sediments, devoid of macroscopic life, after the overlying water column has become oxygenated. Received: 13 April 2000 / Accepted: 8 September 2000