Jurassic soft–bottom oyster Crassostrea from Japan

A new species of oyster, Crassostrea tetoriensis , is described from the lower part of the Middle Jurassic–Cretaceous Tetori Group. This is the oldest record of the genus and one of the earliest muddy–bottom–dwelling oysters. Characteristics of the species include an elongate spatulate outline and narrow attachment area, an elongate hinge area with deep umbonal cavity, a reniform adductor muscle scar, and a smooth commissural margin without chomata. The oysters are cemented to each other and constitute bouquet–like aggregates or colonies of shells in muddy deposits of inferred brackish–water origin. The shells contain lenticular or blister–like chambers filled with a chalky deposit. These ecological and structural properties are characteristic of Crassostrea , and may be regarded as the result of adaptation of the sessile, suspension feeding animals to soft muddy substrates. Oysters are a group of bivalves that have lost mobility and live cemented mostly to hard substrates. Crassostrea and its allies returned secondarily to soft bottoms. The find of C. tetoriensis indicates that the return occurred early in the history of oysters, 60 myr after the first appearance of ostreid oysters in Late Triassic.     

Living on an island: characterization of the encrusting fauna of large pectinid bivalves from the Lower Cretaceous of the Neuquén Basin,west‐central Argentina

Exposed mollusc shells may act as benthic islands in soft bottoms, and the analysis of their encrusting faunas provides unique palaeoecological information. In the late Valanginian of the Agrio Formation (Neuquén Basin, west‐central Argentina), the large pectinid Prohinnites acted as a benthic island on soft substrates. Inequivalved Prohinnites adults with small, smooth cementing scars on the right valve suggest that a free reclining life habit followed the epibyssate juvenile and cementing phases. The encrusting fauna on Prohinnites was studied taxonomically and palaeoecologically by means of a quantitative approach. Over 90% of 123 valves presented encrusters. Encrustation was equally common in both valves. Internal encrustation was rare. The left umbonal region was less encrusted probably due to sediment accumulation or early colonization by soft‐bodied taxa. The fauna was composed of 14 encrusting taxa, including oysters, serpulids, sabellids and cyclostome bryozoans. Oysters exceeded 50% of the total abundance, but serpulids and bryozoans were more diverse. Serpulids and particularly oysters showed a gregarious life habit. Few interactions took place among encrusters and most were post‐mortem, involving the overgrowth of already dead oysters. The oysters were early settlers that took advantage of their gregarious behaviour to rapidly cover available hard surfaces. However, they were unable to exclude bryozoans and polychaetes, which settled on the pectinid's valves regardless of the presence of oysters. The studied fauna corresponds to a climax community that was structured by larval abundance rather than by competitive interactions; oysters settled first and replenished themselves while polychaetes and bryozoans settled over or alongside them     

Morphological and structural adaptations to soft substrates in the Early Jurassic monomyarians Lithiotis and Cochlearites

Lithiotis problematica and Cochlearites loppianus are sessile monomyarian bivalves known from the Early Jurassic of the Tethyan region, where they are found standing vertically in the calcareous mud of lagoonal fades. Their shells are characterized by a long cardinal area, ventrally displaced body space, thick attached valve, and a thin, flat free valve of equal length. A functional ligament is present only in juvenile stages of Cochlearites. To grow straight, the ventral end of the shell had to gape, so in both species it was probably the elasticity of the thin free valve that caused the shell to open. In the adult stage the shell grew only towards the venter with an apparently constant growth rate, while the soft body size remained unchanged. The outer shell is composed of a compact aragonitic layer, while the inner part is filled with loose chalky deposits which are thought to have functioned as supporting the soft body and lifting it upwards. These and other morphologic and structural features, as well as the growth pattern, can be explained as an adaptation of a sessile animal to soft muddy bottoms and rapid sedimentation. Some elongated Crassostrea living in similar environments show remarkable morphologic and structural convergence with Lithiotis and Cochlearites.     

A new mode of ammonite preservation – implications for dating of condensed phosphorite deposits

Unusual phosphatic casts of the ammonites Mortoniceras (Subschloenbachia) sp. and Stoliczkaia sp. from the upper Albian condensed phosphorite bed at Annopol, Poland, are discussed in terms of their taphonomic history. These specimens are interpreted as ‘secondary’ external casts of ammonite replicas preserved originally as attachment scars on oyster shells. The following genetic history is suggested for this previously undocumented mode of ammonite preservation: (1) settling of shells of dead ammonites on the seafloor; (2) colonization of these shells by oysters and formation of ammonite replicas on left valves of oysters; (3) dissolution of ammonite shells; (4) reworking and fragmentation of oyster shells; (5) casting of ammonite replicas by phosphatic material; and (6) separation of ammonite casts from oyster shells, either through mechanical disintegration or dissolution of the latter. The specimens studied were formed after dissolution of the ammonite conchs, not prior to this event as in the case of typical ammonite steinkerns (internal moulds). Therefore, they are here referred to as ‘pseudo‐steinkerns’. The time interval between loss of the original ammonite shells and the formation of oyster‐mediated pseudo‐steinkerns may be very extensive. Therefore, the pseudo‐steinkerns may potentially mislead in biostratigraphic dating of condensed phosphorite deposits.     

Chemical stimuli in coral reefs: how butterflyfishes find their food

Animals use sensory stimuli either to assess and select habitats, mates or food, as well as for communication. The present study aimed to understand the behavioural processes enabling several Chaetodon species (butterflyfishes) to locate one of their food sources (epibionts present on pearl oyster shells) at Rangiroa atoll (French Polynesia). Among the five species tested, our 2-channel choice flume chamber experiments identified three species that were attracted to their food source by chemical stimuli. HPLC experiments showed that pearl oysters and epibionts have specific and unique chemical fingerprints, either one or nine specific peaks, respectively. Overall, chemical stimuli are emitted by both epibionts (used directly by Chaetodon auriga, C. lunula and C. citrinellus) and live pearl oysters (used indirectly by C. auriga and C. lunula) to locate their food source. Biosynthesis of these chemical stimuli could be used to artificially attract butterflyfishes to pearl oyster rearing stations in order to increase the natural cleaning of pearl oyster shells and thus reduce one large cost for this aquaculture.     

Interaction of mineral elements in sea water and shell of oysters (Crassostrea virginica (Gmelin)) cultured in controlled and natural systems

The interaction is reported of selected chemical elements (cadmium, calcium, copper, iron, magnesium, manganese, strontium, and zinc) in cultured sea water, with soft tissues, prismatic calcite of the right valve, and foliated calcite of right and left valves of genetically similar American oysters, Crassostrea virginica (Gmelin) grown in a natural habitat and in two environmentally controlled experimental systems (flow-through and recycle). The addition of trace elements as algal nutrients in ambient sea water was reflected in higher concentrations of trace metals in shells and soft tissues of oysters grown in experimental systems. Calcium was relatively uniformly distributed in major regions of valves from the three habitats, even though its concentration fluctuated widely in sea water in experimental systems. Magnesium and strontium were most concentrated in valves of oysters grown in the recycle system (magnesium in the prismatic layer of the shell and strontium in the foliated calcite). Iron was uniformly distributed. Cadmium, copper, manganese, and zinc were most concentrated in the prismatic calcite of valves from the flow-through system. In soft tissues, calcium was more concentrated in oysters from experimental systems than in those from the natural habitat. Manganese was about equally distributed in soft tissues from the three habitats, whereas copper and iron were more concentrated in soft tissues in experimental systems than in the natural habitat, and were many times more concentrated in soft tissues than in valves from all three habitats. As concentrations of magnesium, strontium, mangenese, zinc, and cadmium increased in valves in experimental systems, pigmentation of valves decreased. The study confirmed the capacity of oysters to concentrate several elements in their valves as concentration of these elements increased in ambient sea water and disclosed the heterogeneous distribution of these elements in major regions of the valves.     

The effects of elevated temperature and dissolved ρCO2 on a marine foundation species

Understanding how climate change and other environmental stressors will affect species is a fundamental concern of modern ecology. Indeed, numerous studies have documented how climate stressors affect species distributions and population persistence. However, relatively few studies have investigated how multiple climate stressors might affect species. In this study, we investigate the impacts of how two climate change factors affect an important foundation species. Specifically, we tested how ocean acidification from dissolution of CO₂ and increased sea surface temperatures affect multiple characteristics of juvenile eastern oysters (Crassostrea virginica). We found strong impacts of each stressor, but no interaction between the two. Simulated warming to mimic heat stressed summers reduced oyster growth, survival, and filtration rates. Additionally, we found that CO₂‐induced acidification reduced strength of oyster shells, which could potentially facilitate crab predation. As past studies have detected few impacts of these stressors on adult oysters, these results indicate that early life stages of calcareous marine organisms may be more susceptible to effects of ocean acidification and global warming. Overall, these data show that predicted changes in temperature and CO₂ can differentially influence direct effects on individual species, which could have important implications for the nature of their trophic interactions.     

Molecular analysis of the bacterial communities in the live Pacific oyster (Crassostrea gigas) and the influence of postharvest temperature on its structure

Aims: To evaluate the effect of postharvest temperature on bacterial communities in live Pacific oysters (Crassostrea gigas) using nonculture‐based methods. Methods and Results: Live oysters were compared before and after storage at 4, 6, 15, 20 and 30°C using terminal restriction fragment length polymorphism (T‐RFLP). Bacterial communities in freshly harvested (control) vs stored oysters were significantly different. Changes in bacterial communities at 4, 15 and 30°C observed by T‐RFLP were further investigated by clone library analysis. Members of the Proteobacteria predominated (43·0–57·0% of clones) in control oysters, while storage altered the bacterial profile. At 4°C, Psychrilyobacter spp. (phylum Fusobacteria) predominated (43·8% of clones), while at 15 and 30°C, members of the phylum Bacteroidetes represented 63·0 and 60·2% of clones, respectively. High microbial diversity in oysters was observed, with at least 73 different genera‐related clones among all samples. Conclusions: Changes in the overall bacterial community of Pacific oysters were influenced by storage temperature and would likely not be detected by standard culture‐based methods currently used to assess oyster quality. Certain dominant genera, such as Psychrilyobacter, Polynucleobacter and a bacterial group related to Alkaliflexus, should be further studied as possible indicators for postharvest temperature control. Significance and Impact of the Study: This work is the first report describing the effect of different storage temperatures on bacterial diversity in postharvest live Pacific oysters using molecular‐based methods.     

How did the carrier shell Xenophora crispa (König, 1825) build its shell? Evidence from the Recent and fossil record

The genus Xenophora comprises species of marine gastropods (Cretaceous-Recent) able to add fragments of various origins to their shell surface. Agglutination potentials vary, from species lacking attachments to species completely covered by agglutinated materials, as in the Mediterranean species Xenophora crispa. Here, we analyse Recent and fossil specimens of Xenophora crispa from the Mediterranean area using SEM and XRD, to better understand their biomineralization patterns and the mechanisms leading to the agglutination of shells, bioclasts and lithoclasts, and their evolution in time. We also provide new data on poorly studied gastropod shell microstructures. We conclude that: (1) most of the Xenophora crispa shell consists of an aragonitic crossed lamellar fabric, but fibrous to spherulitic prismatic fabrics, seemingly of calcite, have been found in the columella and peripheral edge (the thickest parts of the shell); (2) attachment of objects is mediated by a prismatic microstructure, indicating that this may be the most functional fabric in attachment areas in molluscs; and (3) the functional purpose of the agglutination in Xenophora crispa may be related to a snowshoe strategy to successfully colonize muddy substrates, coupled with tactile and olfactory camouflage. Indeed, this species secretes in the columella and peripheral edge a less dense and a more organic rich calcitic fabric, possibly to lighten the shell thickest parts in order not to sink in soft sediments and to facilitate the shell raising from the substrate to create a protected feeding area. This behaviour seems to have been maintained by X. crispa over 2 My time span.     

Substrate specificity in the Devonian tabulate coral Pleurodictyum

The tabulate coral Pleurodictyum americanum Roemer has been cited as an example of a host-specific organism occurring exclusively on the shells of gastropods, particularly Palaeozygopieura hamiltoniae (Hall). Examination of over 1600 specimens of P. americanum, from the Middle Devonian Hamilton Group of western New York, reveals additional complexities which require reinterpretation. While substrate selectivity for Palaeozygopieura shells is evident in all 42 subsamples, a variety of other substrates were also utilized by Pleurodictyum including corals, brachiopods, other molluscs and pebbles. Recent scleractinian corals inhabiting soft bottoms show similar substrate preference, selecting for the tubes of live serpulids, or gastropod shells (invariably with a secondary sipunculid host), but also occasionally settling on unoccupied shells or pebbles. Shell surfaces of P. hamiltoniae, preserved as external molds on the Pleurodictyum epitheca, exhibit encrustation by worm tubes and bryozoans as well as borings and mechanical shell damage, suggesting that these were not the shells of live gastropods. However, the invariant aperture-downward orientation and the high degree of selectivity of P. americanum strongly suggest that the shells were occupied by secondary hosts. □ Substrate specificity, commen-salism, tabulate coral, gastropod, sipunculid, Devonian, Hamilton Group, New York.     

Predation by the endemic whelk Tenguella marginalba (Blainville, 1832) on the invasive Pacific oyster Crassostrea gigas (Thunberg, 1793)

The endemic mulberry whelk (Tenguella marginalba) is a common predator on Australian intertidal rocky shores. The introduced Pacific oyster (Crassostrea gigas), found within the natural range of T. marginalba, is potential prey for the whelk. In experiments designed to increase our understanding of predatory behaviour by the whelk on oysters, we found that adult T. marginalba detected C. gigas and increased movement in the presence of oyster prey. Tenguella marginalba showed a preference for smaller C. gigas, but consumed oysters up to 60?mm in shell height. To access oyster flesh, whelks used their radula to drill holes in the oyster’s shell. These holes were on average 0.68?±?0.09?mm in diameter, most frequently located central to the pericardial cavity on the right (upper) valve. Predation was greatest when predator and prey were both submerged, but was unaffected by a diurnal light cycle. When offered a choice among the native Sydney rock oysters (Saccostrea glomerata), mussels (Trichomya hirsuta) or the invasive C. gigas, whelks displayed no preference among prey. We conclude that the invasive oyster C. gigas represents a viable food source for T. marginalba, which may help to slow the spread of this invasive oyster throughout eastern Australia.     

Oyster reef enhancement utilizing gardened oysters in a subtropical estuary

Crassostrea virginica, the eastern oyster, is a native foundational species that inhabits coastal and estuarine ecosystems along the western Atlantic seaboard. Introduction of C. virginica into estuarine areas with limited or no extant populations is gaining popularity as a pro‐active approach for improving estuarine water quality and creating natural wave breaks for shoreline stabilization. Adult oysters, grown by 113 community members under their private docks, were collected and deployed at three county‐owned sites along the Indian River Lagoon within Brevard County, Florida. In this shallow, warm‐water estuary, replicate treatments deployed at each site included bagged adult oysters collected from gardeners in fall 2014, bagged adult oysters from spring 2015 gardeners, bagged blank (clean) shell, and empty plots (control). Prior to deployment, morphometric data (shell length, weight) were collected on all gardened oysters. Morphometric data were then collected quarterly for all surviving and recruited oysters for 18 months. Our monitoring timeframe was sufficient for assessing survival of gardened oysters, but likely not sufficient to understand recruitment patterns. In areas with no recruitment and limited gardened oyster survival, regular enhancement with live oysters would be needed for long‐term success. In areas with natural recruitment, the blank shell treatment was most successful. Lessons learned from this study include: (1) need for better tracking of abiotic variables (e.g. salinity) where gardening occurred, (2) role of seasonality in initial post‐deployment survival, even in a warm‐water estuary, and (3) importance of pilot studies prior to large‐scale gardened oyster deployments.     

Natural cultch type influences habitat preference and predation,but not survival,in reef‐associated species

A shared origin with fresh and dredged cultch and availability via mining have made fossil cultch a commonly used reef restoration substrate. However, important differences in shape and size between whole‐shell cultch and fossil cultch may impact the complexity of reefs constructed from these materials. To determine if these differences may impact the development of restored reefs, we quantified the interstitial space each cultch type provides and constructed reef mesocosms to measure (1) the immediate effects of exposure to each cultch type on mortality of blue crab (Callinectes sapidus) and pink shrimp (Farfantepenaeus duorarum); (2) the tendency of crab, shrimp, and Florida crown conch (Melongena corona) to be found on habitats composed of each substrate type and their position within each in split‐substrate mesocosms; and (3) the influence of cultch type on predation of Eastern oysters (Crassostrea virginica) by crabs and conch. Aggregation of fossil cultch contains more shells and provides less interstitial space than an equivalent volume of whole‐shell cultch. Although immediate mortality following deployment was low and did not differ among cultch types, we found that all species were more likely to be found on fresh cultch over fossil cultch in choice experiments and used each habitat type differently. Cultch type also impacted the size of oysters consumed by crabs in short‐term feeding trials. The structure and traits of habitats created by various materials should be added to the growing list of issues considered when natural communities are to be restored in oyster reefs and other environments.     

Distribution and diversity of the Syllidae (Annelida: Polychaeta) from the Mexican Gulf of Mexico and Caribbean

The objective of this study was to contribute to the knowledge of the syllids distributed in the eastern coasts of Mexico (Gulf of Mexico and Caribbean) taking into consideration their composition, distribution and diversity. Sampling was done at 77 stations including 49 in soft bottoms (terrigenous and carbonate sediments), and 28 in hard bottoms (coralline substrates) sampled in nine reef zones included in three reef systems (Veracruz, Campeche and Cozumel). In all 2084 individuals were collected (13.5% in soft sediments and 86.4% in coralline areas). Forty-five species were identified: 31 in soft bottoms and 38 in corals, with 21 species common to both types of environments. In soft bottoms, the highest species richness and diversity were found in the Carbonate Zone, while syllids were uncommon in the Terrigenous Zone. In hard bottoms, the highest density, species richness and diversity values were found in the Veracruz Reef System, followed by the Campeche Bank and Cozumel Island Reef Systems. The dominant species in soft bottoms were Haplosyllis spongicola, Exogone lourei, Typosyllis cornuta and T. papillosus, whereas T. alosae, T. corallicola, T. cornuta and H. spongicola dominated in hard bottoms. The importance and success of the syllids in both soft bottoms and reef environments, particularly in those of the Gulf of Mexico is emphasized.     

Removal of Toxoplasma gondii Oocysts from Sea Water by Eastern Oysters (Crassostrea virginica)

SUMMARY. Toxoplasma gondii infections have been reported in a number of marine mammals. Presently it is not known how these animals acquire T. gondii from their aquatic environment. The eastern oyster, Crassostrea virginica , has been shown to remove Cryptosporidium oocysts from seawater and a similar phenomenon may be occurring with T. gondii oocysts and marine invertebrates. The present study was done to determine if eastern oysters could remove and retain T. gondii oocysts from seawater. Oocysts of the VEG strain of T. gondii (1 × 10⁶ oocysts) were placed in seawater (32 ppt NaCl) containing live eastern oysters. The infected seawater was removed one day postinoculation (PI) and replaced with fresh seawater. Selected oysters were removed at 1, 3 and 6 days PL Hemolymph, gill washes, and oyster tissue were collected separately at each observation time. The oyster tissue was homogenized and all 3 samples fed separately to mice. Toxoplasma gondii positive mice were observed at each time period. The results indicate that T. gondii oocysts can be removed from seawater by eastern oysters and retain their infectivity. Contaminated raw oysters may serve as a source of T. gondii infection for marine mammals and humans.     

Soft‐bottom tube worms: from irregular to programmed shell growth

Like other secondary soft‐bottom dwellers, serpulid and sabellid tube worms used particular strategies in order to cope with their earlier loss of mobility. This is expressed by the transition from irregular to genomically programmed morphologies of their calcareous shells that guarantee a stable horizontal resting position. In contrast to permanent recliners, however, this attitude was probably only the starting position for active resurrection after the muddy tail of storm sediments had settled on top of the displaced animal.     

Phenotypic plasticity in oysters (Crassostrea virginica) mediated by chemical signals from predators and injured prey

Prey organisms reduce predation risk by altering their behavior, morphology, or life history. Avoiding or deterring predators often incurs costs, such as reductions in growth or fecundity. Prey minimize costs by limiting predator avoidance or deterrence to situations that pose significant risk of injury or death, requiring them to gather information regarding the relative threat potential predators pose. Chemical cues are often used for risk evaluation, and we investigated morphological responses of oysters (Crassostrea virginica) to chemical cues from injured conspecifics, from heterospecifics, and from predatory blue crabs (Callinectes sapidus) reared on different diets. Previous studies found newly settled oysters reacted to crab predators by growing heavier, stronger shells, but that adult oysters did not. We exposed oysters at two size classes (newly settled oyster spat and juveniles ~2.0 cm) to predation risk cue treatments including predator or injured prey exudates and to seawater controls. Since both of the size classes tested can be eaten by blue crabs, we hypothesized that both would react to crab exudates by producing heavier, stronger shells. Oyster spat grew heavier shells that required significantly more force to break, an effective measure against predatory crabs, when exposed to chemical exudates from blue crabs as compared to controls. When exposed to chemical cues from injured conspecifics or from injured clams (Mercenaria mercenaria), a sympatric bivalve, shell mass and force were intermediate between predator treatments and controls, indicating that oysters react to injured prey cues but not as strongly as to cues released by predators. Juvenile oysters of ~ 2.0 cm did not significantly alter their shell morphology in any of the treatments. Thus, newly settled oysters can differentiate between predatory threats and adjust their responses accordingly, with the strongest responses being to exudates released by predators, but oysters of 2.0 cm and larger do not react morphologically to predatory threats.     

PCR-RFLP genotyping for Japanese and Korean populations of Pacific oyster using mitochondrial DNA noncoding region

Production of Pacific oyster Crassostrea gigas in Miyagi and Hiroshima (Japan) has gradually increased, with a marked increase in imported oysters from Goseong (Korea), and then cultured oysters of the Miyagi, Hiroshima, and Goseong populations accounting for most oyster consumption in Japan. In this study, we developed a simple PCR-RFLP analysis of a mitochondrial DNA noncoding region (mtDNA-NCR) for differentiating cultured oysters of these populations. PCR amplification yielded a 818 bp fragment comprising the entire mtDNA-NCR and parts of adjacent tRNA^Cys and tRNA^Asn genes from all the specimens. By use of a wide restriction test using 30 different restriction enzymes, only a single enzyme only, AluI, produced a unique RFLP pattern enabling us to discriminate Miyagi and Hiroshima oysters from Goseong oysters. This difference is probably because of nucleotide alteration at the presumptive AluI recognizing site on position 439 of the mtDNA-NCR. Our simple, robust and cost-effective PCR-RFLP analysis is potentially useful for population genetic investigation of cultured Pacific oyster, particularly when large numbers of specimens must be analyzed.     

Epizootiology of Minchinia costalis in susceptible oysters in Seaside Bays of Virginia's Eastern Shore, 1959–1976

“Seaside Disease” of oysters caused by Minchinia costalis (Haplosporida, Sporozoa) produced annual mortalities on the Seaside of the Delmarva Peninsula along the middle Atlantic Coast from Chesapeake Bay to Delaware Bay, U.S.A. The May–June mortalities occurred from 1959 to 1976 without exception; deaths began in late May, peaked in June, and were usually over by July 1. The pathogen developed rapidly from March to May, and sporulation occurred in connective tissues of all organs in May and June. Exposure to a May–June enzootic was required to obtain infections. The pathogen remained subclinical until late winter of the following year. A sympatric pathogen, Minchinia nelsoni, which kills oysters extensively in lower Chesapeake Bay, was present but caused only minor mortalities. Salinities > 30 parts per thousand seem to favor M. costalis and inhibit M. nelsoni. Prevalences of both diseases in live oysters or gapers are given for 11 of the 18 years monitored.     

Hettangian (Early Jurassic) Dinosaur Tracksites from the Mecsek Mountains,Hungary

A rare example of a North American Jurassic hardground is found in the Carmel Formation of southwestern Utah. The Carmel hard‐ground was formed across a carbonate lagoon from an oolitic shoal seaward to a subtidal shelly facies landward. It has an abundant bivalve fauna consisting of thick layers of encrusters (the oyster Liostrea and the plicatulid Plicatula), borers (the ichnofossil Gastro‐chaenolites with the mytilid Lithophaga often preserved inside), and nestlers (the mytilid Modiolus). A rare soft‐bodied bryozoan (Arach‐nidium) is preserved by bioimmuration in the attachment scars of Liostrea; this is the first bioimmuration recorded from the Jurassic of North America, and the first bioimmuration recorded from a hard‐ground. The phoronid boring Talpina is present in some Liostrea shells; it was apparently excavated after the death of these oysters. The Carmel hardground community does not contain other fossils, such as serpulids, brachiopods, foraminiferans, and skeletal bryo‐zoans, typical of Jurassic hardgrounds elsewhere. It represents a low diversity molluscan community developed in a restricted marine environment.