Bryozoans as taphonomic engineers,with examples from the Upper Ordovician (Katian) of Midwestern North America

A combination of encrusting calcitic bryozoans and early seafloor dissolution of aragonitic shells recorded in the Cincinnatian Series of the upper Midwest of North America allowed the preservation of abundant moulds of mollusc fossils bioimmured beneath the attachment surfaces of the bryozoans. We here call this preservational process ‘bryoimmuration’, defined as a bryozoan‐mediated subset of bioimmuration. The bryozoans moulded very fine details of the mollusc shells, usually with more accuracy than inorganic sediment moulds. Most of the bryozoans are heterotrypid trepostomes with robust low‐Mg calcite skeletons. The molluscs are primarily bivalves, gastropods, nautiloids and monoplacophorans with their originally aragonitic shells now dissolved. Many of the encrusting bryozoans are so thin and broad that they give the illusion of calcitic mollusc shells clinging to the moulds. Some molluscs in the Cincinnatian, especially monoplacophorans and epifaunal bivalves, would be poorly known if they had not been bryoimmured. Unlike internal and external moulds in sediment, bryoimmured fossils could be transported and thus record aragonitic faunas in taphonomic assemblages (e.g. storm beds) in which they would otherwise be rare or absent. In addition, bryoimmurations of aragonitic shells often reveal the ecological succession of encrustation on the shells by exposing the earliest encrusters and borings that were later overgrown. Bryoimmuration was common during the Late Ordovician because the calcite sea at the time quickly dissolved aragonitic shells on the seafloor before final burial, and large calcitic bryozoans very commonly used molluscs as substrates. Bryoimmuration is an important taphonomic process for preserving aragonitic faunas, and it reveals critical information about sclerobiont palaeoecology. Several Cincinnatian mollusc holotypes are bryoimmured specimens. Bryozoans involved in bryoimmuration enhance the preservation of aragonitic fauna and thus act as taphonomic engineers.     

Linking taphonomy to community-level abundance: Insights into compositional fidelity of the Upper Triassic shell concentrations (Eastern Alps)

Although actualistic live/dead comparisons lead to robust estimates of fidelity of modern death assemblages, quantitative evaluation of fidelity of fossil assemblage remains uncertain. In this paper, effects of storm reworking on compositional fidelity of the Upper Triassic shell concentrations (Eastern Alps, Austria) are evaluated. An exploratory approach is based on comparison of reworked and non-reworked assemblages in ordination analyses. Non-reworked assemblages of one or more communities provide a baseline for evaluation of fidelity of reworked assemblages. In siliciclastic-rich intervals of the Kössen Formation, shell concentrations are represented by (1) packstones with small, shallow infaunal bivalves, (2) floatstones and pavements with large semi-infaunal bivalves, and (3) bioclastic marlstones. In carbonate-rich intervals, bioclastic floatstones with bivalves and brachiopods occur. Analyzing all shell concentrations, eight sample groups sharing similar species composition are discriminated. Limited effect of storm reworking on composition of shell concentrations is indicated by (1) a general persistence of six sample groups when only non-reworked assemblages are analyzed, (2) similarity in composition between reworked and non-reworked assemblages within sample groups, and (3) compositional segregation between non-reworked assemblages of distinctive sample groups, mostly without any reworked assemblages of intermediate composition.Depth-related variations in dead-shell production, shell destruction and body size governed preservation and distribution of the shell concentrations along onshore-offshore gradient in the Kössen Basin. First, at times when environmental conditions were unfavorable for shell producers, coupled with high background shell destruction rates, limestone beds formed during storm events were shell-poor. Second, less common shell concentrations in upper than in lower parts of siliciclastic intervals can be related to higher environmental stress in shallower habitats. Third, the difference between shell concentrations dominated by small and large bivalves is driven by between-habitat differences in body size and is not due to a differential sorting of small and large shells. Combining community analysis based on species abundances with taphonomic analysis can thus be helpful in tracking fidelity of fossil assemblages.     

Evidence for Palaeozoic orthoconic cephalopods with bimineralic shells

Based on the aragonite composition of extant and exceptionally preserved fossil cephalopods going back to the early Palaeozoic, it is commonly assumed that all externally shelled cephalopods had an aragonitic shell wall. We demonstrate herein that at least two taxa of Siluro‐Devonian orthoconic nautiloids (Dawsonoceras, Spyroceras) had an original bimineralic shell, which developed convergently with gastropods and bivalves.     

Facies and fauna of the Pennsylvanian Buckhorn Asphalt Quarry deposit: a review and new data on an important Palaeozoic fossil <Emphasis Type="Italic">Lagerstätte</Emphasis> with aragonite preservation

The Pennsylvanian Buckhorn Asphalt Quarry contains the best-preserved Palaeozoic mollusc fauna in the world. Early impregnation of mixed siliciclastic–carbonate rocks (mudstones, pack to grainstones, shell beds, and conglomerates) with hydrocarbons prevented aragonite destruction (“Impregnation Fossil Lagerstätte”). The exceptional preservation comprises shell microstructures, microornaments and early ontogenetic shells. Most gastropods had planktotrophic larval development indicating a high primary production although the remains of phytoplankton are very rare in this and other Late Palaeozoic deposits. Deposition occurred close to a shallow-water coastal area. Mass flow processes (density currents) triggered by storms were involved in the transport mechanisms of some units. Shells of benthic molluscs yield the most diverse known Palaeozoic microboring assemblage, indicating at least partly euphotic conditions. The invertebrate fauna comprises about 160 species and is dominated by molluscs, which is unusual for a Palaeozoic deposit, suggesting that aragonite dissolution produces a major bias in the fossil record. However, most mollusc genera in the Buckhorn deposit are also known from other Pennsylvanian occurrences as recrystallised shells. This shows that preservation bias via preferential aragonite dissolution may be overestimated.     

Aragonite shells are more ancient than calcite ones in bivalves: new evidence based on omics

Two calcium carbonate crystal polymorphs, aragonite and calcite, are the main inorganic components of mollusk shells. Some fossil evidences suggest that aragonite shell is more ancient than calcite shell for the Bivalvia. But, the molecular biology evidence for the above deduction is absent. In this study, we searched for homologs of bivalve aragonite-related and calcite-related shell proteins in the oyster genome, and found that no homologs of calcite-related shell protein but some homologs of aragonite-related shell proteins in the oyster genome. We explained the results as the new evidence to support that aragonite shells are more ancient than calcite shells in bivalves combined the published biogeological and seawater chemistry data.     

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.     

Tafonomía de bivalvos holocenos en la costa del estuario de Bahía Blanca, Argentina

In the Bahía Blanca Estuary (Argentina), Holocene deposits are fossiliferous sand ridges originated by the last transgressive-regressive marine event. By means of ternary taphograms applied on valves of Brachidontes rodriguezi a taphonomic analysis was carried out in two ridges, both in the external and internal estuary area. Six taphonomic attributes were considered. The disarticulation and the fragmentation are higher in the external ridge, whereas the bioerosion and incrustation, both barely manifested, are similar in the two environmental settings. In the outer deposit, as much the abrasions as the dissolution of the shells are more intense. Genetically, it is considered that these shell concentrations were originated by multiepisodic storm events. Based on the taphonomic analysis, it arises that in the generation of the internal ridge waves of moderate energy with different sorting degrees prevailed, whereas in the formation of the external deposits waves of greater energy with smaller sorting capacity prevailed.     

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.     

Sequence stratigraphic significance of sedimentary cycles and shell concentrations in the Upper Jurassic-Lower Cretaceous of Kachchh, western India

Upper Jurassic and Lower Cretaceous siliciclastic shallow water sediments of the Kachchh Basin, western India, form strongly asymmetric coarsening-upward cycles, which are interpreted as recording changes in relative sea level (deepening-shallowing cycles). These cycles correspond to depositional sequences, in which deposits of the lowstand systems tract are not present, the sequence boundary coinciding with the transgressive surface. Shell concentrations are found in the transgressive lags at the base of the transgressive systems tract (TST), in the maximum flooding zone (MFZ), and at or close to the top of the highstand systems tract. They belong to six assemblages, five of them dominated by large bivalves such as Seebachia, Herzogina, Gryphaea, Gervillella, Megacucullaea, Pisotrigonia and Indotrigonia, the sixth by the coral Amphiastraea. Three types of shell concentrations can be distinguished that differ from each other in a number of ecological and taphonomic features, such as species diversity, preservation quality, orientation in cross-section, percentage of disarticulation, and degree of biogenic alteration. Characteristic features of concentrations at the base of the TSTs are moderate time-averaging, sorting, a preferred convex-up orientation, and nearly total disarticulation of shells. They are suggestive of an environment in which reworking and local transport were frequent events. Similar features are shown by concentrations near the tops of the HSTs, except that there shells were largely concentrated in lenses and in pavements rather than in beds as in the transgressive lags. Associated sedimentary structures indicate deposition above fair weather wave base in a high-energy environment. Concentrations occurring in the MFZ, in contrast, are autochthonous and highly time-averaged, having accumulated during times of low rates of sedimentation below storm wave base. This is supported by their high preservation quality (comparatively high percentage of articulated shells, shells of infaunal organisms commonly preserved in life position), biogenic alteration being the most important taphonomic agent. The dominant elements of these shell concentrations, i.e. Seebachia, Megacuccullaea, and Indotrigonia in the Upper Jurassic, and Pisotrigonia in the Lower Cretaceous, are endemic to the Ethiopean faunal province and belong to lineages that rapidly evolved during this time period.     

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.     

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.     

Dissolution Dominating Calcification Process in Polar Pteropods Close to the Point of Aragonite Undersaturation

Thecosome pteropods are abundant upper-ocean zooplankton that build aragonite shells. Ocean acidification results in the lowering of aragonite saturation levels in the surface layers, and several incubation studies have shown that rates of calcification in these organisms decrease as a result. This study provides a weight-specific net calcification rate function for thecosome pteropods that includes both rates of dissolution and calcification over a range of plausible future aragonite saturation states (Ω_ar). We measured gross dissolution in the pteropod Limacina helicina antarctica in the Scotia Sea (Southern Ocean) by incubating living specimens across a range of aragonite saturation states for a maximum of 14 days. Specimens started dissolving almost immediately upon exposure to undersaturated conditions (Ω_ar∼0.8), losing 1.4% of shell mass per day. The observed rate of gross dissolution was different from that predicted by rate law kinetics of aragonite dissolution, in being higher at Ω_ar levels slightly above 1 and lower at Ω_ar levels of between 1 and 0.8. This indicates that shell mass is affected by even transitional levels of saturation, but there is, nevertheless, some partial means of protection for shells when in undersaturated conditions. A function for gross dissolution against Ω_ar derived from the present observations was compared to a function for gross calcification derived by a different study, and showed that dissolution became the dominating process even at Ω_ar levels close to 1, with net shell growth ceasing at an Ω_ar of 1.03. Gross dissolution increasingly dominated net change in shell mass as saturation levels decreased below 1. As well as influencing their viability, such dissolution of pteropod shells in the surface layers will result in slower sinking velocities and decreased carbon and carbonate fluxes to the deep ocean.     

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.     

Mollusc larval shell formation: amorphous calcium carbonate is a precursor phase for aragonite

The larval shells of the marine bivalves Mercenaria mercenaria and Crassostrea gigas are investigated by polarized light microscopy, infrared spectroscopy, Raman imaging spectroscopy, and scanning electron microscopy. Both species contain similar shell ultrastructures. We show that larval shells contain amorphous calcium carbonate (ACC), in addition to aragonite. The aragonite is much less crystalline than non-biogenic aragonite. We further show that the initially deposited mineral phase is predominantly ACC that subsequently partially transforms into aragonite. The postset juvenile shell, as well as the adult shell of Mercenaria also contains aragonite that is less crystalline than non-biogenic aragonite. We conclude that ACC fulfills an important function in mollusc larval shell formation. It is conceivable that ACC may also be involved in adult shell formation.     

Not all aragonitic molluscs are missing: taphonomy and significance of a unique shelly lagerstätte from the Jurassic of SW Britain

The Blue Lias Formation at Lyme Regis (Dorset, UK) includes an exceptional pavement of abundant large ammonites that accumulated during a period of profound sedimentary condensation. Ammonites were originally composed of aragonite, an unstable polymorph of calcium carbonate, and such fossils are typically prone to dissolution; the occurrence of a rich association of aragonitic shells in a condensed bed is highly unusual. Aragonite dissolution occurs when pore‐water pH is reduced by the oxidization of hydrogen sulphide close to the sediment‐water interface. Evidence suggests that, in this case, the oxygen concentrations in the overlying water column were low during deposition. This inhibited the oxidation of sulphides and the associated lowering of pH, allowing aragonite to survive long enough for the shell to be neomorphosed to calcite. The loss of aragonite impacts upon estimates of past biodiversity and carbonate accumulation rates. The preservational model presented here implies that diagenetic loss of aragonite will be greatest in those areas where dysoxic‐anoxic sediment lies beneath an oxic waterbody but least where the sediment and overlying water are oxygen depleted. Unfortunately, this implies that preservational bias through aragonite loss will be greatest in those biotopes which are typically most diverse and least where biodiversity is lowest due to oxygen restriction.     

The interplay between event and background sedimentation and the origin of fossil‐rich carbonate concretions: a case study in Permian rocks of the Paraná Basin,Brazil

The Permian Serra Alta Formation was generated under transgressive conditions within a large, calm epeiric sea. A monotonous succession of ‘barren’, massive mudstones deposited under oxygen‐deficient conditions (mainly below storm wave base) is the main lithofacies of this unit. Fossils are generally rare and diluted in the matrix, but certain intervals contain shell‐rich concentrations with well‐preserved, closed articulated bivalves, mixed with shells and comminuted debris with variable quality of preservation, all encased in carbonate concretions. Two main scenarios may account for the origin of these bivalve‐rich concretions (i.e. unique events in sea‐water chemistry or unique burial‐starvation couplets). Sedimentological and taphonomic information indicates that the final deposition of the original shell‐rich mudstone intervals was probably tied to episodic influx of fine‐grained sediments in distal settings. Moderate bioturbation is also recorded suggesting low rates of sedimentation prior to early diagenesis. Hence, the fossil concentrations in concretions were formed due to the interplay of event and background sedimentation. These are internally simple concentrations with complex depositional histories. The concretion‐bearing beds are not randomly distributed in the Serra Alta Formation. Rather, they are found in the sparsely fossiliferous offshore deposits of the basal to intermediate portions of the unit. Thus, the concretionary mudstone beds and associated deposits are preserved in particular intervals and can be tracked for kilometres. This indicates that the conditions essential for concretion development existed only at particular stratigraphical intervals. Finally, our study strongly corroborates the idea that concretions are critical sources of sedimentological, taphonomic and stratigraphical information.     

Aragonitic dendritic prismatic shell microstructure in Thracia (Bivalvia,Anomalodesmata)

The shells of most anomalodesmatan bivalves are composed of an outer aragonitic layer of either granular or columnar prismatic microstructure, and an inner layer of nacre. The Thraciidae is one of the few anomalodesmatan families whose members lack nacreous layers. In particular, shells of members of the genus Thracia are exceptional in their possession of a very distinctive but previously unreported microstructure, which we term herein “dendritic prisms.” Dendritic prisms consist of slender fibers of aragonite which radiate perpendicular to, and which stack along, the axis of the prism. Here we used scanning and transmission electron microscopical investigation of the periostracum, mantle, and shells of three species of Thracia to reconstruct the mode of shell calcification and to unravel the crystallography of the dendritic units. The periostracum is composed of an outer dark layer and an inner translucent layer. During the free periostracum phase the dark layer grows at the expense of the translucent layer, but at the position of the shell edge, the translucent layer mineralizes with the units typical of the dendritic prismatic layer. Within each unit, the c‐axis is oriented along the prismatic axis, whereas the a‐axis of aragonite runs parallel to the long axis of the fibers. The six‐rayed alignment of the latter implies that prisms are formed by {110} polycyclically twinned crystals. We conclude that, despite its distinctive appearance, the dendritic prismatic layer of the shell of Thracia spp. is homologous to the outer granular prismatic or prismatic layer of other anomalodesmatans, while the nacreous layer present in most anomalodesmatans has been suppressed.     

Shell microstructure and mineralogy of the Littorinidae: ecological and evolutionary significance

An examination of the shell microstructure and mineralogy of species from 30 of the 32 genera and subgenera of the gastropod family Littorinidae shows that most species have a shell consisting of layers of aragonitic crossed-lamellar structure, with minor variations in some taxa. However, Pellilitorina, Risellopsis and most species of Littorina have partly or entirely calcitic shells. In Pellilitorina the shell is made entirely of calcitic crossed-foliated structure, while in the other two genera there is only an outer calcitic layer of irregular-prismatic structure. A cladistic analysis shows that the calcitic layers have been independently evolved in at least three clades. The calcite is found only in the outermost layers of the shell and in species inhabiting cooler waters of both northern and southern hemispheres. Calcium carbonate is more soluble in cold than warm water and, of the two polymorphs, calcite is about 35% less soluble than aragonite. We suggest that calcitic shell layers are an adaptation of high latitude littorinids to resist shell dissolution.     

Shell mineralogical trends in epifaunal Mesozoic bivalves and their relationship to seawater chemistry and atmospheric carbon dioxide concentration

The Late Triassic-Early Jurassic change from aragonite- to calcite-facilitating conditions in the oceans, which was caused by a decrease of the Mg²⁺/Ca²⁺ ratio of seawater in combination with an increase of the partial pressure of carbon dioxide, also affected the shell mineralogy of epifaunal bivalves. In the “calcite sea” of the Jurassic and Cretaceous, the most diverse and abundant families of epifaunal bivalves had largely calcitic shells. Some of them, such as the Inoceramidae, acquired this shell mineralogy earlier in Earth's history but did not significantly diversify until the onset of “calcite sea” conditions. Others, however, replaced aragonite by calcite in their shell at the beginning of the Jurassic, as shown for the Ostreidae, Gryphaeidae, Pectinidae, Plicatulidae, and Buchiidae. In these families, replacement of aragonite by calcite took place in the middle and inner layer of the shell and was not associated with changes in morphology and life habit. It is therefore proposed that lower metabolic costs rather than higher resistance against dissolution or advantageous physical properties triggered the calcite expansion in their shells. This explanation fits well the observation that clades of thin-shelled bivalves were less affected by the change of seawater chemistry. Thick-shelled clades, by contrast, may suffer a severe decline in diversity until they adapt their shell mineralogy, as demonstrated by the Hippuritoida: The diversity of the Megalodontoidea, which failed to adapt their shell mineralogy to “calcite sea” conditions, dramatically decreased at the end of the Triassic, whereas their descendents became dominant carbonate producers during the Late Mesozoic after they acquired a calcitic outer shell layer in the Late Jurassic. These examples indicate that changes in the seawater chemistry and in the partial pressure of carbon dioxide are factors that influence the diversity of carbonate-secreting animals, and, as in the case of the decline of the Megalodontoidea, may contribute to mass extinctions.     

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.