Taphonomic windows and molluscan preservation

Recent studies on silicified fossil biotas have suggested that substantial skewing of the molluscan record resulted from early aragonite dissolution in mid-outer carbonate ramp settings. If those rare skeletal lagerstätten are representative, then the quality and completeness of the molluscan record are thrown into doubt. Yet database studies suggest that the bivalve fossil record is actually relatively complete. If so, then biodiversity must be captured by other processes that preserved shells vulnerable to early dissolution, and which operated on a relatively high frequency, i.e., less than the species duration for bivalves.Storm beds, shell plasters and submarine hardgrounds are identified as fossil deposits that can preserve the labile aragonitic component of the fauna and thus represent potential taphonomic windows. Many storm event beds include rich accumulations of shelly benthos. Differences between storm bed faunas and those of the background facies could reflect transportation effects. However, some storm bed assemblages are rich in originally aragonitic infaunal bivalves that are not represented in background facies or more proximal shelf equivalents, and here rapid burial and removal of organic matter by winnowing may be the keys to aragonite shell preservation. Despite Palaeozoic to Cenozoic changes in the thickness and frequency of shell beds that reflect the predominant bioclast producers, shallow infaunas are commonly concentrated together with epifauna in such deposits.Some low energy, organic-rich mud-dominated settings are associated with preservation of aragonitic molluscs. Infaunal bivalves are a prominent component of shell plasters or pavements in such settings, linked to episodic bottom water anoxia. Decaying algal blooms drew the redox boundary up above the sediment–water interface, and brought populations of infaunal bivalves to the surface where they died. Isolated from the oxic taphonomically active zone, the shells were not dissolved and were buried as thin shell layers. In similar settings, aragonitic shells were preserved as moulds through early pyritisation, or even through preservation of original shell aragonite.In oxic environments, bioturbational reworking of surface sediment destroyed moulds of aragonitic shells after early dissolution. In some hardgrounds, these delicate moulds were preserved due to synsedimentary cementation, probably using carbonate released by aragonite dissolution. The examples included here come from both intervals of “calcite” and “aragonite” seas, and it is not possible to assess whether the saturation state (with respect to aragonite) of the ambient sea water played a role in the selective removal of aragonitic shells.While taphonomic windows may have captured the diversity of individual groups, it is clear from quantitative data involving skeletal lagerstätten that the scale of loss from early aragonite dissolution has drastically altered the trophic composition of some fossil assemblages commonly used as the basis for reconstructions of past communities.     

a new ichnogenus for etchings made by cheilostome bryozoans into calcareous substrates

Some encrusting cheilostome bryozoans etch a pattern of small pits into hard calcareous substrates, especially calcitic and aragonitic shells of molluscs. These patterns, herein described as Leptichnus ichnogen. nov., comprise pits which are sub-circular to elongate in cross section and are found in either uniserial ( L. dromeus isp. nov.) or multiserial arrangements ( L. peristroma isp. nov., the type species). Each pit corresponds to the location of a single zooid in the bryozoan colony. The oldest known Leptichnus is Late Cretaceous (Maastrichtian), the trace fossil first becomes common in the Cenozoic, and at least nine modern cheilostome genera produce incipient Leptichnus. Leptichnus can be the only evidence remaining of encrusting cheilostomes following taphonomic or diagenetic loss of their calcareous skeletons. The mechanism by which bryozoans etch into their calcareous substrates is unknown but is almost certain to be chemical and necessitates having windows in the basal walls of the zooids which permit contact with the substratum beneath. Etching may result in better adherence to the substrate, giving protection from abrasion and bioerosion.     

The development of an Early Ordovician hard ground community in response to rapid sea-floor calcite precipitation

The Kanosh Shale (Upper Arenig, Lower Ordovician) of west-central Utah. USA. contains abundant carbonate hardgrounds and one of the earliest diverse hardground communities. The hardgrounds were formed through a combination of processes including the development of early digenetic nodules in clay sediments which were exhumed and concentrated as lags by storms. These cobble deposits. together with plentiful biogenic metrical. were cemented by inorganically precipitated calcite on the sea floor. forming intraformational conglomerate hardgrounds. Echinoderms may have -played a critical role in the development of hardground faunas since their disarticulated calcite ossicles were rapidly cemented by syntaxial overgrowths. forming additional cobbles and hardgrounds. The echinoderms thus may have taphonomically facilitated the development of some of the hard substrates they required. A significant portion of the hardground cements may have been derived from the early dissolution of aragonitic mollusk shells. Kanosh hardground species include the earliest bryozoans recorded on hardgrounds and large numbers of stemmed echinoderms. primarily rhipidocystid cocrinoids. Bryozoans and echinoderms covered nearly equal areas of the hardground surfaces. and there was a distinct polarization between species which preferred the upper. exposed portions of the hardgrounds and others which were most common on undercut. overhang surfaces. The Kanosh Shale hardground fossils combine elements of Late Cambrian assemblages and Middle Ordovician faunas, thus confirming predicted trends in hardground community evolution. especially the replacement of cocrinoids by bryozoans and. to a lesser extent, by other stemmed echinoderms, especially crinoids. The Kanosh community marks the transition from the Cambrian Fauna to The Paleozoic Fauna in The hardground ecosystem. *Carbonate hardgrounds, aragonite dissolution, calcite cement, Echinodermara, Trepostomata, Nicholsonclla. Dianulites. Porifpra. taphonomic facilitation, Utah. Pogonip Group, Kanosh Shale. Ordovician.     

Biotic interactions revealed by macroborings in arctic bivalve molluscs

The presence of organisms whose bodies have low preservation potential may be deduced by searching for the traces produced by them. The addition of predatory gastropods and soft-bodied epizoans to Quaternary marine faunas dominated by bivalves was facilitated by an examination of borings in bivalve shells. Borings attributed to predatory gastropods (ichnogenus Oichnus ) were observed in shells of Astarte spp., Hiatella arctica and Macoma calcarea. Astarte, Hiatella and Macoma were preyed upon in preference to other members of a diverse suspension-feeding bivalve community. Borings attributed to epizoans (ichnogenus Cautostrepsis ) were observed in bivalve shells (Astarte spp. Hiatella arctica ), calcareous algae and limestone clasts. Biotic interactions revealed by trace fossils are employed, for the first time, to reconstruct the trophic structure of arctic Quaternary marine benthic faunas. ▭ Arctic molluscs, palaeoecology, Oichnus, Caulostrepsis.     

Molecular Evolution of Mollusc Shell Proteins: Insights from Proteomic Analysis of the Edible Mussel Mytilus

Shell matrix proteins (SMPs) that are embedded within calcified layers of mollusc shells are believed to play an essential role in controlling the biomineral synthesis and in increasing its mechanical properties. Among the wide diversity of mollusc shell textures, nacro-prismatic shells represent a tremendous opportunity for the investigation of the SMP evolution. Indeed, nacro-prismatic texture appears early in Cambrian molluscs and is still present in the shell of some bivalves, gastropods, cephalopods and very likely also, of some monoplacophorans. One key question is to know whether these shells are constructed from similar matrix protein assemblages, i.e. whether they share a common origin. Most of the molecular data published so far are restricted to two genera, the bivalve Pinctada and the gastropod Haliotis. The shell protein content of these two genera are clearly different, suggesting independent origins or considerable genetic drift from a common ancestor. In order to describe putatively conserved mollusc shell proteins, here we have investigated the SMP set of a new bivalve model belonging to another genera, the edible mussel Mytilus, using an up-to-date proteomic approach based on the interrogation of more than 70,000 EST sequences, recently available from NCBI public databases. We describe nine novel SMPs, among which three are completely novel, four are homologues of Pinctada SMPs and two are very likely homologues of Haliotis SMPs. This latter result constitutes the first report of conserved SMPs between bivalves and gastropods. More generally, our data suggest that mollusc SMP set may follow a mosaic pattern within the different mollusc models (Mytilus, Pinctada, Haliotis). We discuss the function of such proteins in calcifying matrices, the molecular evolution of SMP genes and the origin of mollusc nacro-prismatic SMPs.     

Pyritized preservation of chancelloriids from the Cambrian Stage 3 of South China and implications for biomineralization

The enigmatic Cambrian animal chancelloriids were discovered in a wide range of taphonomic settings; however, preservation of biomineralized sclerite microstructure was solely known from secondarily phosphatized skeletal remains. Here, we investigate a uniquely pyritized chancelloriid from the lower Cambrian Guojiaba Formation in southern Shaanxi Province, China, using a combination of advanced analytic techniques. Results of the energy dispersive spectroscopy (EDS), X-Ray Fluorescence (XRF), and Raman spectrum show that the sclerites and scleritomes are preserved as pyritized internal moulds with a calcitic outer layer. The outer layer enveloping the internal moulds likely represents the recrystallized counterpart of the original biomineralized sclerite wall. Distinctive fibrous microstructures are discovered in the sclerites, which echo the features seen in the phosphatized fossils of chancelloriids. The typical microstructure, along with the recrystallized calcite, corroborate the interpretation that chancelloriid sclerites were originally constructed by fibrous aragonite. The stability of the microstructure and mineral composition in both carbonate and siliciclastic backgrounds indicate that chancelloriids were adapted to exploit aragonitic fibres to build their skeletons regardless of the change of their living environments.     

Mineralogy of Arctic bryozoan skeletons in a global context

Bryozoans are major carbonate producers in some ancient and Recent benthic environments, including parts of the Arctic Ocean. Seventy-six species of bryozoans from within the Arctic Circle have been studied using XRD to determine their carbonate mineralogies and the Mg content of the calcite. The majority of species were found to be calcitic, only four having bimineralic skeletons that combined calcite and aragonite, and none being entirely aragonitic. In almost all species, the calcite was of the low- (<4 mol% MgCO₃) or intermediate-Mg (4–11.99 mol% MgCO₃) varieties. Previous regional studies of bryozoan biomineralogy have found higher proportions of bimineralic and/or aragonitic species in New Zealand and the Mediterranean, with a greater number of calcitic species employing intermediate- and high-Mg calcite. The Antarctic bryozoan fauna, however, has a similar mineralogical composition to the Arctic. The lesser solubility of low-Mg calcite compared to both Mg calcite and aragonite in cold polar waters is most likely responsible for this latitudinal pattern. However, it is unknown to what extent environmental factors drive the pattern directly through eliciting an ecophenotypic response from the bryozoans concerned or the pattern reflects genetic adaptations by particular bryozoan clades.     

Selective solution of macroinvertebrate calcareous hard parts: a laboratory study

Ten suites of 16 common types of invertebrate hard parts were placed in acid baths for 24 hours to determine relative rates and common styles of dissolution. Skeletal mineralogies included aragonite and both high-magnesium and low-magnesium calcite. Hard parts included barnacle cxoskelctons, cchinoid tests, gastropod opercula and gastropod and bivalve shells. Calcitic barnacle plates dissolved most rapidly, aragonitic and high magnesium calcitic hard parts showed intermediate rates, and the calcitic shells of the oyster dissolved at the lowest rate. The surface area to weight ratio of the hard parts correlated ( r =0.650) significantly with the hard part's rate of dissolution. Skeletal remains with a high surface area to weight ratio dissolved faster than those with a low surface area to weight ratio. Skeletal porosity and mineralogy appeared to be responsible for additional variation in the rate of dissolution. The effect of the surface area to weight ratio is sufficient to overcome the effect of mineralogy. Dense, compact aragonitic hard parts can persist longer than porous, thin calcitic remains. Typical features associated with skeletal degradation include development of chalky textures, thinning of distal margins, surface etching and formation of holes in bivalve muscle scars. Such features may aid in the recognition of partial dissolution of skeletal remains in the rock record. □ Taphonomy, paleoecology, fossil-diagenesis.     

Taphonomic differentiation of Oxfordian ammonites from the Cracow Upland, Poland

Taphonomic analysis of Lower and Middle Oxfordian ammonites from the Cracow Upland, southern Poland (localities at Pod???e, Zalas, M?ynka) revealed differences in ammonite preservation. The studied ammonites, usually termed as external and internal moulds, show a more complex state of preservation. In the Middle Oxfordian glauconitic marls, ammonites are preserved as internal moulds with neomorphic calcite shells showing relics of the original internal structure. In the Middle Oxfordian platy peloidal limestones, ammonites are preserved mostly as external moulds, without septal suture, however under microscope might show relics of internal whorls and septa and/or subtle differences in sediment filling phragmocone chambers. In sponge–microbial bioherms and biostromes, ammonite internal moulds have shells, which in contrast to ammonites from glauconitic marls are not strictly neomorphic ones, but originated by shell dissolution and subsequent filling of moldic porosity by calcite cement. In sponge–microbial nodular limestones, the ammonites are strongly deformed and the outer wall is usually removed by dissolution under pressure. Other important taphonomic differences include the rate of compaction (highest in platy limestones), sedimentary infillings, microborings, encrustations and preservation of siphuncular tubes. The majority of the ammonites appear to be phragmocones; aptychi in all facies are rare. Siphuncular tubes are fossilized exclusively in oppeliids, only in specimens from glauconitic marls and platy limestones, although their other taphonomic attributes are different. Tubes seem to have fossilized due to microbially mediated phosphatization that could be favoured by a set of parameters which operated rather at the scale of ammonoid carcasses: closed, poorly oxygenated conditions, and reduced pH. Taphonomic processes were controlled by the sedimentary environment (fragmentation, sedimentary filling, phosphatization of siphuncular tubes), as well as by early and late diagenesis (neomorphic transformation, dissolution, cementation, compaction) influenced by lithology.     

Effect of end-Triassic CO<Subscript>2</Subscript> maximum on carbonate sedimentation and marine mass extinction

Correlation of stratigraphic sections from different continents suggests a worldwide interruption of carbonate sedimentation at the Triassic–Jurassic boundary, which coincided with one of the most catastrophic mass extinctions in the Phanerozoic. Both events are linked by a vulcanogenic maximum of carbon dioxide, which led to a temporary undersaturation of sea water with respect to aragonite and calcite and a corresponding suppression of carbonate sedimentation including non-preservation of calcareous skeletons. Besides the frequently cited climatic effect of enhanced carbon dioxide, lowering the saturation state of sea water with respect to calcium carbonate was an additional driving force of the end-Triassic mass extinction, which chiefly affected organisms with thick aragonitic or high-magnesium calcitic skeletons. Replacement of aragonite by calcite, as found in the shells of epifaunal bivalves, was an evolutionary response to this condition.     

Carbonate mineralogy of a tropical bryozoan biota and its vulnerability to ocean acidification

Decreasing pH levels in the world’s oceans are widely recognized as a threat to marine life. Bryozoans are among several phyla that produce calcium carbonate skeletons potentially affected by ocean acidification (OA). Depending on species, bryozoan skeletons can consist of calcite, aragonite or have a bimineralic combination of these two minerals. Aragonite is generally more soluble in seawater than calcite, making aragonitic species more vulnerable to OA. Here, for the first time we use Raman spectroscopy to determine the mineral composition of a tropical bryozoan biota. Compared with bryozoan biotas from higher latitudes in which calcite predominates, aragonite was found to occur in a much higher proportion of the 22 cheilostome bryozoan species collected from the shorelines of Penang and Langkawi in Malaysia, where 46% of species are calcitic, 41% aragonitic and 13% bimineralic. All but one of the aragonitic or bimineralic species belong to the ascophorans, whereas calcitic skeletons characterized most of the anascans, many of which are primitive ‘weedy’ malacostegines. These results suggest a relatively high vulnerability of tropical bryozoan faunas to OA, with the weedier taxa likely to be least impacted.     

Taphonomy and ichnology: tools for interpreting a maximum flooding interval in the Berriasian of Tlemcen Domain (Western Tellian Atlas,Algeria)

During the Middle-Late Berriasian, a long-term climatic and eustatic change occurred, documented in the literature. However, data from the northern Gondwana paleomargin are scarce. This research analyzes the Lamoricière Clay Formation at the Ouled Mimoun section, focusing on fossil assemblages and using taphonomic and ichnological aspects to interpret a transgressive–regressive cycle. The section starts with mudstones and oolitic grainstones representing shallow-water environments in the top part of the Ouled Mimoun Marly Limestones Formation (Upper Tithonian p.p. to lowermost Berriasian). The base of the Lamoricière Clay Formation is characterized by a high clay content but was still deposited in shallow water, as indicated by the record of the ostracod Asciocythere, dasyclad green algae, and the sponge Cladocoropsis. The subsequent record of fossil-rich calcareous beds at the beginning of the Upper Berriasian (Boissieri Zone) with ammonoids and calpionellids is congruent with an increase in water depth. The sedimentation rate in the Late Berriasian was reduced, as indicated by the increment of fossil remains and trace fossils. Ammonoid moulds show taphonomic features pointing to long-lasting exposure on the sea floor prior to burial with corrasion and encrustation by sessile organisms such as serpulids, thecideidinids, and bryozoans. During calm periods, crustaceans and worms intensely burrowed the sea floor. The record of Thalassinoides and Rhizocorallium indicates bottom conditions ranging from soft to firm. The low sedimentation rate and sediment by-passing probably favored early lithification. The increasing carbonate content as well as decreasing sedimentation rate is compatible with the maximum distance to emerged areas during maximum flooding. High-energy events, probably related to storms, favored the exhumation and extreme corrasion of ammonite moulds and trace fossils. In the resulting substrate, limonitic films developed and encrusting organisms proliferated (serpulids, bryozoans, and thecideidinids), colonizing both the bottom surface (hardground) and exhumed moulds of ammonoids and Thalassinoides. The uppermost 0.7 m of the section represents the return to shallow conditions, with increasing sedimentation rate and terrigenous detrital content, along with the disappearance of hemipelagic forms (ammonoids); hence it is interpreted as having developed at the beginning of a regressive context.     

Palaeocology of Hard Substrate Faunas from the Cretaceous Qahlah Formation of the Oman Mountains

Skeletal encrusters and carbonate hardgrounds are rare in siliciclastic sands and gravels because of high levels of abrasion and sediment movement. An exception to this is the Maastrichtian Qahlah Formation of the Oman Mountains, a sequence of coarse siliciclastic sediments deposited on a shallow marine shelf above wavebase and at an equatorial palaeolatitude. This unit contains intercalated carbonate hardgrounds and other hard substrates which were encrusted and bored. The hard substrates, comprising carbonate and silicate clasts, calcareous bioclasts (mollusc shells and coral fragments) and wood, supported a diverse encrusting and boring fauna dominated in biomass by the oyster Acutostrea . There are twelve bryozoan species and at least two serpulid worm species, most living cryptically. Other encrusters on exposed surfaces include the agglutinated foraminiferan Placopsilina and several species of colonial corals. Borings in the carbonate clasts and shells are predominantly those of bivalves ( Gastrochaenolites ), with subsidiary clionid sponge ( Entobia ) and acrothoracican barnacle ( Rogerella ) borings. The woodgrounds are thoroughly bored by teredinid bivalves ( Teredolites ). Of the common substrate types, carbonate hardground clasts support the greatest number of taxa, followed by chert clasts, with limestone rockground pebbles being depauperate. Clast composition and relative stability probably explain these differences. Individual clasts probably had variable and typically long colonisation histories. Detailed palaeoecological interpretation is constrained by taphonomic loss, time-averaging and clast transportation and reorientation. Evidence from the Qahlah Formation shows that tropical rocky-shore biotas in the Cretaceous were not impoverished as previously believed.     

Large gryphaeid oysters as habitats for numerous sclerobionts: a case study from the northern Red Sea

The shell of a living specimen of the Indo-Pacific gryphaeid giant oyster Hyotissa hyotis was colonized by numerous encrusting, boring, nestling and baffling taxa which show characteristic distribution patterns. On the upper valve, sponge-induced bioerosion predominates. On the lower valve intergrowth of chamid bivalves and thick encrusting associations—consisting mostly of squamariacean and corallinacean red algae, acervulinid foraminifera, and scleractinian corals—provides numerous microhabitats for nestling arcid and mytilid bivalves as well as for encrusting bryozoans and serpulids. Such differences between exposed and cryptic surfaces are typical for many marine hard substrata and result from the long-term stable position of the oyster on the seafloor. The cryptic habitats support a species assemblage of crustose algae and foraminifera that, on exposed surfaces, would occur in much deeper water.     

Signs of predation in the Middle Jurassic of south-central Poland: evidence from echinoderm taphonomy

Distinct faunal aggregates are described from the Middle Jurassic (uppermost Bajocian/lowermost Bathonian and Middle Bathonian) clay deposits of Częstochowa area, south-central Poland. These aggregates are composed of molluscs (scaphopods, gastropods, bivalves, ammonites and belemnites), articulate brachiopods and echinoderms (asteroids, crinoids and echinoids). A large percentage of the fossils, especially bivalves, are fragmented, but some fossils are complete. Although most of the fossils are crushed and fragmented, they are still identifiable to at least the genus level. Thorough statistical analysis of taphonomic features indicates that the preservation of asteroid marginal plates is distinct from the ossicles derived from the host clays. The high frequency of bite marks and the good state of preservation suggest that the accumulations are the products of predation activities and most probably are the effect of regurgitation. Taking into account the rich and diverse fauna, the predator was a bottom-feeding generalist. Possible predators include palaeospinacid sharks, a tooth of which was collected from the same bedding surface, but not associated with regurgitated remains. Although the bite marks on the asteroid ossicles point to sharks as potential producers of regurgitates, other vertebrates, like durophagous pycnodontiform fish, cannot be excluded.     

Shell microstructure in the Permian nonmarine bivalve Palaeomutela Amalitzky: Revision of the generic diagnosis

The microstructure of aragonitic and calcitic shells of the genus Palaeomutela Amalitzky, 1891 is examined. The aragonitic shell consists of three main layers, each is distinguished by certain crossed lamellar microstructure: comarginal, radial, and complex. As aragonite is recrystallized into pelitic calcite, microstructural shell features are preserved. Many species of Palaeomutela from localities of different age display the same microstructural pattern, which is possible to regard as a character of generic rank.     

Morphology and genesis of nodular phosphates in the Cenomanian Glauconitic Marl of south-east England

Phosphatic nodules are abundant in the Glauconitic Marl (Cretaceous, Cenomanian) of south-east England, particularly where the sequence is condensed. Some of the nodules are derived from the underlying Upper Greensand, and are phosphatic fossil fragments, fossil moulds, and calcareous concretions. Concretions in particular show signs of a complex history of multiple phases of boring, encrustation, phosphatisation, and glauconitisation. Phosphate and glauconite are both replacements after fine-grained carbonate sediment and cement. The majority of the phosphates are whole and fragmentary moulds of fossils. The origin of theses nodules involved: (1) infilling of shells, (2) burial, (3) prefossilisation-cementation of fossil infillings. probably by high magnesian calcite, (4) dissolution of aragonitic shell material, (5) disinterment and exposure of moulds on the sea floor, followed by (6) phosphatisation. boring. and enerustation by various organisms, and sometimes glauconitisation. Many nodules bear evidence of several cycles of cementation, exposure, mineralisation, boring and enerustation.
The closest Recent analogues to the Glauconitic Marl phosphates appear to be the phosphatic crust and nodules forming today off the coast of southern California. The features described and processes inferred from the Glauconitic Marl occurrences appear to have been widespread in nodular phosphatic facies.     

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     

Bryozoans and microbial communities of cool-temperate to subtropical latitudes—paleoecological implications

The environmental distribution of encrusting bryozoans settling on disarticulated and living bivalve shells has been recorded from five stations in Japan and New Zealand. Some insight into the observed distribution patterns emerges from information on the interaction of bryozoans with microbial mats. Advancing existing classifications, we have subdivided the encrusting bryozoan morphotypes into seven different growth types that largely reflect the biological potentials of bryozoans in competition for space on substrate surfaces. The frequency distribution of these types (s-/c-/m-/z-laminae, runners, spots, bryostromatolites) reveals the influence of microbial mats as a control factor of bryozoan substrate coverage. Microbial mats in turn are correlated with latitudinal gradients in Japan and New Zealand from cool-temperate to subtropical and tropical waters. Unlike erect bryozoans, laminar ones are probably underrated as facies fossils. Accordingly, laminar bryozoan growth types are reconsidered as a tool for paleoecological interpretation of marine hard substrate communities.     

Shell disintegration and taphonomic loss in rudist biostromes

Radiolitid biostromes in the Upper Cretaceous of Austria and Italy record a marked taphonomic loss controlled mainly by the composition of the biocoenosis, by the density of rudist colonization, by the style of radiolitid shell disintegration and by early diagenetic processes. Radiolitid shells consisted of a calcitic ostracum and an originally aragonitic hypostracum. The attached valve of most radiolitids was built of (1) an outermost ostracal layer of delicate calcite lamellae, (2) a thick layer of ‘boxwork ostracum’ built of radial funnel plates and cell walls, (3) a thin, inner ‘ostracal layer 3’ of thick-walled boxwork, and (4) the hypostracum that formed the innermost shell layer. The attached valve disintegrated by spalling of radial funnel plates of layer 2, and by selective removal of the boxwork ostracum. In the free valve, the ostracum consisted of two layers: (a) an inner, lid-shaped layer of dense calcite, and (b) an outer layer composed of calcite lamellae. The free valve disintegrated by spalling into ostracal and hypostracal portions, by spalling of the ostracum into layers a and b, and by disintegration of layer b into packages of calcite lamellae and individual lamellae. The specific style of disintegration of the radiolitids was aided or induced by discontinuities in shell structure. Lamellar fragments from the ostracum of the upper valve and from the radial funnel plates of the lower valve locally are abundant in free-valve-funnel-plate floatstones that comprise the matrix of or occur in lenses within radiolitid biostromes. In biostromes with an open parautochthonous fabric, selective removal of the boxwork ostracum of the attached valve occurred by mechanical spalling and, most probably, by early diagenetic dissolution. Complete removal of the boxwork ostracum yielded thin, relict shells composed of the ‘ostracal layer 3’ and the hypostracum. During early diagenesis, the hypostracum was replaced by blocky calcite spar, or was dissolved and became filled by internal sediments. The combination of both selective removal of boxwork ostracum and early diagenetic dissolution of aragonite locally resulted in the formation of ghost biostromes that entirely or largely consist of faint relics of radiolitids. The syndepositional formation of radiolitid shell relics and the presence of radiolitid ghost biostromes produced by bios-tratinomic and early diagenetic processes show that rudist biostromes can undergo marked taphonomic loss during fossilization. The presence of ghost biostromes with a burrowed, open parautochthonous rudist fabric indicates that the final preservation of a rudist biostrome was directly influenced by the characteristics of the biocoenosis, including unpreserved burrowing taxa. Rudist biostromes may be of markedly different taphonomy as a result of the taxonomic composition of the entire assemblage and the density of colonization by the rudists.