Microgeographical shell variation in Littorina striata, a planktonic developing periwinkle

Littorina striata is a strictly Macaronesian, intertidal periwinkle with planktonic development. The species produces both nodulose and smooth shells, which co-occur at Ilheu de Vila Franca do Campo, a drowned crater situated about 1000?m off the south coast of São Miguel, Azores. The present work describes and analyzes the shell variation, temporal change and ecological distribution of the two shell types at this crater over a 3-year period. Nodulose shells were more common in the sheltered lagoon inside the crater, while smooth specimens dominated the outside of the wave-exposed crater. Moreover, nodulose specimens were smaller and weighed less compared to smooth ones. However, regardless of morphotype, shells from the lagoon had a smaller aperture and were less globose than those from the outside. Within an exposure regime, smooth specimens had a larger aperture. These patterns remained constant over time. They are tentatively interpreted as functional adaptations to thermal stress and wave exposure. The mechanisms that maintain these patterns are still unknown. Yet, aperture height of specimens transplanted from the lagoon to the outside increased markedly over a period of 5 months, whereas no comparable changes were observed in other experimental groups (i.e. transplanted from the outside into the lagoon, transplanted within the outside and transplanted within the lagoon; the latter two transplant groups being the blancos). This suggests that at least the aperture size in L. striata may be an ecophenotypically plastic trait.     

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     

Shell ecophenotype in the blue mussel (<Emphasis Type="Italic">Mytilus edulis</Emphasis>) determines the spatial pattern in foraging behaviour of an oystercatcher (<Emphasis Type="Italic">Haematopus ostralegus</Emphasis>) population

When feeding on blue mussels (Mytilus edulis), oystercatchers (Haematopus ostralegus) either stab into the mollusc’s gaping valves or hammer through its dorsal or ventral shell. Whilst the selectivity of hammering and stabbing oystercatchers for specific prey morphologies has been well studied, the way in which the effects of environment on M. edulis morphology can in turn affect feeding methods of H. ostralegus is very poorly understood. Based on morphological analyses on randomly selected shells from three intertidal zones, this study failed to detect differences in morphology or distribution of dorsally and ventrally hammered shells but confirms the finding of previous authors that hammering oystercatchers select thinner mussels than stabbing birds. Additionally, we show that this difference in optimal prey morphology can lead to spatial patterns in oystercatcher feeding behaviour. Whilst at the low intertidal and higher mid intertidal zones, characterised by comparatively thick shells, most empty shells had apparently been stabbed, hammering was the dominant feeding behaviour at the lower mid intertidal zone, where shells were thinner. Preference of hammering birds for smaller mussels was not ubiquitous. Sagittal shell shape was predominantly influenced by allometric growth effects and had only minor effect on prey selection. All oystercatchers preferred less inflated mussels, with the degree of shell inflation gradually increasing with higher intertidal elevation. Our results illustrate the importance of small-scale patterns in prey ecophenotypes in determining the distribution and feeding dynamics of wading birds.     

Enzymatic and morphological criteria for distinguishing between Cardium edule and C. glaucum of the Portuguese coast

Cardium edule L. and C. glaucum Bruguière are two sibling species of cockles commonly found in littoral systems along the Portuguese coast, where they sometimes occur as mixed populations. The problem of morphological separation of these two cockle species has not been completely solved. Indeed, the distinctive morphological criterion proposed by Brock after identifying the specimens by means of electrophoretic patterns of hepatopancreatic malate dehydrogenase was not effective when applied to Portuguese Cardium spp. The object of the present study was to establish morphological discrimination criteria for distinguishing Portuguese C. edule and C. glaucum on the basis of initial identification of individuals through the enzymatic criterion established by Brock. Cockles were collected from 1988 to 1991 at 16 estuarine and lagoonal systems along the Portuguese coast. Hepatopancreatic extracts were submitted to horizontal starch gel electrophoresis following Brock's procedure. The malate dehydrogenase patterns obtained were identical to those found by Brock. Clear interspecific differences in the leucine aminopeptidase and esterase isozyme systems were also detected. The morphological study of the shells involved 7 morphometric and 9 qualitative and semiquantitative characters. Canonical variate analysis of the morphometric characters showed that these could not be used for identification purposes. However, a clear separation between the two species was obtained by a principal-component analysis of qualitative and semiquantitative characters. The characters mainly useful for distinguishing between the two species were valve profile, shape of ventral valve junction, type of calcareous scales and shell colour.     

Spatial structure of communities on dead pen shells (<Emphasis Type="Italic">Atrina rigida</Emphasis>) in sea grass beds

Delimiting communities in marine habitats is difficult because co-occurring species often have different life histories and the life stages experience the environment at different spatial scales. The habitat of a particular community is embedded within a larger habitat or ecosystem with many species shared between the focal community and the larger system. Pen shells (Atrina rigida) are large bivalves that, once the mollusk dies, provide shelter for motile species and hard substrate for settling larval invertebrates and egg-laying fishes. In St. Joseph’s Bay, Florida (29°45′N, 85°15′W), pen shells are the most abundant source of hard substrate, especially inside sea grass (Thalassia testudinum) beds, where they reach densities of 0.1–4.0 m⁻². This study, which was conducted from May to August 2005, measured the overlap in species densities between dead pen shells and the surrounding sea grass communities at eight sites to determine the discreteness of the pen shell communities. Of the 70-epibenthic taxa recorded, 66% were found on the pen shells but not in the surrounding sea grass habitat. Community structure, which varied among shells within sites and among the eight sites, could be related to sea grass characteristics such as blade density and length either directly (e.g., inhabitants of pen shells directly benefit from the surrounding sea grass) or indirectly (e.g., pen shells and sea grass both benefit from similar factors such as current and nutrients). Pen shells were randomly distributed at several spatial scales within the 15 × 15 m sites as were many motile species. Two exceptions were the shrimp, Palaemon floridanus and the amphipod, Dulichella appendiculata, whose distributions were clumped. Most of the sessile species had clumped distributions, tending to be very abundant when they were present. These pen shell communities provide an opportunity for experimental studies of factors affecting species diversity on small, discrete, naturally occurring habitats. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Richness,abundance and shell use of subarctic and arctic hermit crabs

Hermit crabs are an obvious, common and abundant feature of global shallow water environments but almost nothing is known of their ecology at their extreme latitudinal ranges, the Arctic and southern South America. In this study, we report the first investigation on hermit crabs and their use of a key resource, gastropod shells, in a subpolar and a polar environment—amongst the most rapidly changing places on earth. Hypothesised low levels of richness were confirmed by surveys—only Pagurus pubescens was found in western Spitsbergen and only three pagurids were found in northern Norway. At the northern-most of their extent, hermit crabs were fairly common but Arctic abundances (1–5 m⁻²) were an order of magnitude lower than in many warm temperate or tropical localities. Along the open coast of Spitsbergen, the occurrence of P. pubescens was infrequent and very patchy, but it was more abundant in the fjords. In Isfjorden, the largest studied fjord, the population of P. pubescens was mainly represented by small individuals. Spitsbergen P. pubescens used few shell types and >87% just occurred in Margarites or Buccinum shells. The proportion of the gastropod shells, used by P. pubescens, which were damaged, was high and increased with shell size. These hermit crabs are at the edge of the range for both their species and Anomura. The extremes of their location are reflected by: They probably represent the least rich assemblage, with the lowest and most patchy typical abundances reported to date and are amongst the smallest individuals using the least diverse and most damaged gastropod shells.     

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     

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.     

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.     

Using shell middens to assess effects of fishing on queen conch ( Strombus gigas ) populations in Los Roques Archipelago National Park, Venezuela

Queen conch, Strombus gigas, is a commercially important gastropod that has been exploited throughout the Caribbean islands for thousands of years. Shell middens in the region are the physical record of a long-term fishery and their study can provide valuable information on selectivity patterns followed by fishermen and on resulting morphological shifts reflected by shells. In this study, we surveyed 27 middens located at Los Roques, Venezuela, to assess pre-Columbian and modern fisheries and measure their impact on local populations of queen conch. Pre-Columbian middens, covering a period of approximately 350 years of exploitation (1160–1540 A.D.), were mostly composed of adult shells (89%) and mean length of catch was estimated at 22.4 ± 0.2 cm (mean ± SE). A decrease in mean length of catch was observed throughout the modern fishery regime, estimated at 22.2 ± 0.3 cm in 1950–1971 and 20.0 ± 0.3 cm in 1990–1995. Higher proportions of immature individuals ranging from 48 to 67% were found in modern middens. Additionally, a decrease in mean length of mature individuals was detected throughout the modern fishery regime. The appearance of younger and smaller specimens is considered as a sign of heavy exploitation of local populations of queen conch during modern fisheries. Pre-Columbian middens in contrast, permitted to establish a relative baseline from which to compare values registered throughout modern times.     

Slowest of the slow: latitudinal insensitivity of burrowing capacity in the bivalve <Emphasis Type="Italic">Laternula</Emphasis>

Low temperature limits the rate of biochemical reactions and aerobic scopes of cold water ectotherms. To compensate for this limiting effect, animals living in cold environments often possess physiological or morphological adaptations to maintain vital functions. Cross-latitudinal comparison of aerobic capacities is one method to test which factors constrain activity in thermally distinct environments particularly when congeneric studies are carried out on related species with conservative ecology and habitat. Burrowing is a major aerobic activity of bivalve molluscs that is described here for the first time for the tropical mangrove species Laternula truncata and Laternula boschasina and then compared with their Antarctic congener Laternula elliptica. About 80% of L. truncata (16.3–46.1 mm shell length) and 63% of L. boschasina (11.3–27.7 mm shell length) buried within 24 h at 28°C. The burrowing rate index (BRI = [³√wet weight/time to bury]×10⁴) ranged between 1.1 and 20.2 for L. boschasina and 1.1–32.9 for L. truncata. These values are 2–3 orders of magnitude less than other tropical bivalve molluscs and are amongst the lowest recorded for any bivalve. Comparisons with the Antarctic L. elliptica showed little or no differences in BRI (Q ₁₀ of 1.0–1.2 for specimens of the same size). This is contrary to the general pattern over a wide range of bivalves, where BRI increases with a Q ₁₀ of between 2.9 and 6.4 between high latitudes and the equator. L. elliptica has 25–30% longer relative foot length than tropical congeners of the same size, which could be a morphological adaptation compensating for reduced burrowing speeds in a colder environment. Burrowing rates within the genus Laternula could, however, also be maintained by differing habitat, ecological and physiological constraints on burrowing capability.     

Precipitation of aragonite by calcitic bivalves in Mg-enriched marine waters

To understand the relative importance of biological versus physicochemical control over biomineralization, we have tested if the chemical composition of the medium (i.e., the Mg/Ca ratio) can change the mineralogy of mollusk shells. The shells of mollusks are made of calcite and/or aragonite, which are by far the most common CaCO₃ polymorphs. Several species of bivalves with predominantly calcitic shells have been cultivated in artificial seawater with a Mg/Ca molar ratio within the range of 8.3–9.2, well above the present value for seawater (5.2). Four out of six species used (the scallop Chlamys varia, the oyster Ostrea edulis, the saddle oyster Anomia ephippium and the mussel Mytilus edulis) survived long enough to secrete significant amounts of calcium carbonate. The deposits (sometimes extensive) formed on the interior shell surfaces were predominantly aragonitic. Three individuals of C. varia also increased their length by adding new shell at the margin. Contrary to the internal shell deposits, these margins were high-Mg calcite. This implies that the marginal mantle is able to exert a more strict control on the secreted mineral phase than the mantle facing the internal shell surface. This is the first report on an in vivo experimentally forced switch in bivalve shell mineralogy, from calcite to aragonite due to a change in water chemistry.     

Climatic effects on the growth of a temperate reef fish from the Southern Hemisphere: a biochronological approach

Increments in the hard parts of marine organisms (otoliths, skeletons, shells) can provide long-term chronologies of growth analogous to tree rings. For the first time in the Southern Hemisphere, we use a dendrochronological (tree-ring analysis) approach to develop a multidecadal chronology of growth for a temperate reef fish, Girella tricuspidata, from the coast of northern New Zealand. Growth patterns in the otoliths of this species were strongly synchronous among individual fish over a period spanning 27 years (1980–2006). We then compared our otolith chronology to climatic records and found strong positive correlations of growth with sea surface temperature, and weak negative correlations with the multivariate El Nino Southern Oscillation (ENSO) index. Strongest correlations were found between summer sea surface temperature and otolith growth. This relationship was consistent across all years and explained 44 % of the variation (y = −2.0 + 0.1785 × temperature, r ² = 0.4367, P = 0.0002) in the G. tricuspidata growth chronology. Our study illustrates how otolith chronologies provide remarkable records of annual growth patterns over decadal time scales that will be useful for forecasting the likely effects of climate change on marine ecosystems.     

Growth and sexual maturity of the northern propellerclam (<Emphasis Type="Italic">Cyrtodaria siliqua</Emphasis>) in Eastern Canada,with bomb radiocarbon age validation

The northern propellerclam Cyrtodaria siliqua is a common bycatch in the Arctic surfclam, Mactromeris polynyma fishery on Banquereau Bank in Eastern Canada. Samples of the propellerclam from this exploited fishery were used to determine the life history characteristics of the population. The age structure of the population is dominated by old animals to ages exceeding 100 years. We validated the age estimates for the propellerclam through analysis of bomb-produced radiocarbon in the shell growth increments deposited before, during and after the atmospheric atomic bomb testing periods of the 1950s and 1960s. Radiocarbon from shells with presumed birth dates between the late 1950s and 1970s clearly reflected the sharp increase in oceanic radiocarbon attributable to previous nuclear testing, indicating that age estimates based on shell increment counts are accurate. Estimates of von Bertalanffy growth parameters revealed that the growth rate of the population was relatively rapid for the first 20 years of life, slowing down to very low growth rates thereafter. Sexual maturity was estimated as being reached at 28.6 mm in length and 4.7 years in age. Size–weight morphometric relationships were also calculated.     

Stable isotopes (δ18O and δ13C) in Spirula spirula shells from three major oceans indicate developmental changes paralleling depth distributions

Stable isotopes (δ¹⁸O and δ¹³C) were measured in successive chambers of the aragonitic shells of the small deep-sea squid Spirula spirula (Linnaeus 1758) (class Cephalopoda, subclass Coleoidea, order Sepioidea, family Spirulidae) to determine whether their depth distributions change with age. The spiral shells, ranging in diameter from 18 to 23 mm (30–38 chambers), were collected between 2000 and 2006 from beaches in six widely separated locations in three oceans, the Atlantic (Tobago and Canary Islands), Indian (Madagascar, Maldives, and Perth, Australia), and Pacific Oceans (Ulladulla, Australia). The patterns for both isotopes were highly correlated in specimens from all six sites. The δ¹⁸O data suggest that after hatching at depths >1,000 m at temperatures of 4–6°C, the squid migrate into shallower, warmer waters at 12–14°C at depths of 400–600 m. Subsequently, the increasing δ¹⁸O values suggest a migration back into somewhat cooler, deeper habitats. The δ¹³C values also revealed three ontogenetic stages in all six specimens, including a major shift from positive to negative values, which probably corresponds to sexual maturation, the initiation of reproduction, and concomitant changes in diet. In three of the six specimens (from Tobago, Canary Islands, and Maldives) a fourth embryonic stage (not detected in the oxygen data) was accompanied by markedly less positive δ¹³C values in the first few chambers. These data, combined with the scanty life history information from previous studies of S. spirula, can be used to compare the habitat requirements of related extant and fossil cephalopod genera. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Marker technique for investigating gut throughput rates in coral reef fishes

Artemia sp. shells were evaluated to determine their accuracy for tracing the passage of algal filaments through the gut of the damselfish Pomacentrus amboinensis Bleeker, 1868 (family Pomacentridae), an omnivorous coral reef fish. An automatic faeces-collection apparatus enabled the quantitative collection of markers and faeces in the laboratory. Defecation rates were similar for light and heavy doses of Artemia sp. shells and controls, indicating no detrimental effects of Artemia sp. shells on the gut throughput rate of P. amboinensis. In addition, similar rates and patterns of the passage of Artemia sp. shells and the algal markers Enteromorpha sp. and Lyngbya sp. indicated that Artemia sp. shells provide a reliable representation of the throughput rate of algal filaments. The mean throughput time of P. amboinensis was 4.6 h ±0.3 SE, with a modal recovery time of 4 h. Laboratory throughput estimates were validated by comparing the distribution patterns of Artemia sp. shells in the dissected gut of specimens administered markers in the laboratory and field. In addition, the retention of markers in the stomach of P. amboinensis suggested a likely site of prolonged processing. Received: 24 October 1996 / Accepted: 18 March 1997     

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     

Evidence of a 2-day periodicity of striae formation in the tropical scallop Comptopallium radula using calcein marking

The periodicity of striae formation in the tropical scallop Comptopallium radula (Indo-West Pacific Ocean) was investigated with an in situ marking technique, using the calcein fluorochrome. To minimize scallop stress caused by excessive handling, in situ benthic chambers were used for marking experiments. Once marked, scallops (shell height range: 38.4–75.8 mm) remained on site in a large benthic enclosure and were collected at regular time intervals to count new striae formed after marking, over a period of 3 months. A 3-h exposure period with calcein (150 mg l⁻¹) was sufficiently long to create a detectable mark in nearly all shells. It was, however, impossible to count the striae in 48.2% of the shells (mainly large specimens) because of a very small growth after marking. Lack of significant mortality during the experiments indicated that tested calcein concentrations were not lethal. A decrease in shell growth rate was observed after marking but the respective impacts of calcein toxicity and changes in environmental conditions could not be discriminated. Our results suggest that in situ calcein marking inside benthic chambers is suitable for shell growth studies of scallops, provided the latter are not too old. After marking, the juvenile C. radula formed an average of one stria every 2.1 days in summer. Reports of 2-day periodicity in biological rhythms are rare. Striae formation in C. radula may be controlled by an endogenous oscillator, synchronized by an environmental cue acting as a zeitgeber, such as seawater temperature or sea level pressure, both of which exhibit 2-day variations in the Pacific Ocean. As in many other scallop species, C. radula forms striae periodically under natural conditions, but this study shows that in pectinid juveniles, this periodicity can deviate from a daily cycle. These results suggest that C. radula shells have tremendous potential for recording environmental conditions during periods ranging from months to a few years and with a resolution of 2 days.     

Seasonal variation in movement,aggregation and destructive grazing of the green sea urchin (<Emphasis Type="Italic">Strongylocentrotus droebachiensis</Emphasis>) in relation to wave action and sea temperature

Hydrodynamic forces are an important determinant of subtidal community structure, particularly when they limit the distribution and foraging ability of mobile consumers. We examined the effect of wave action on the rate of movement and destructive grazing of a kelp bed by the green sea urchin (Strongylocentrotus droebachiensis) under field conditions. We measured density and rate of advance at fixed intervals along ∼100 m of a grazing front over 1 year, and quantified individual movement rates in the barrens 5–10 m behind the urchin front using a time-lapse videography. Seasonal variation in the mean rate of advance of the front (range: 0–4 m month⁻¹) was explained by changes in urchin density at the front (120–360 individuals m⁻²), which in turn varied inversely with significant wave height (0.5–2 m). Water temperature (0.8–17.6°C) had no effect on the rate of advance or on urchin density (aggregation) at the front, except when temperature exceeded 17°C. Movement of individual urchins also was affected by wave action: we observed a significant decrease in speed and displacement of urchins with increasing significant wave height. Wave action had no effect on the proportion of urchins moving or the degree of linearity of their movements. We propose that the decrease in urchin density at the front associated with increased wave action, results from de-aggregation, which reduces the risk of dislodgement, combined with a reduction in urchin movement in barrens, which supplies new urchins to the front.