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.     

Alternative biofacies model for dysaerobic communities

An abundant calcareous fauna has been discovered in the oxygen minimum zone (OMZ; < 0.5 ml/l O₂) off central California at oxygen concentrations considerably less than those predicted by previous ecological models. Analysis of box core samples and bottom photographs has revealed a distinct depth zonation of echinoderms. Asteroids and ophiuroids are most abundant along the upper and lower edges of the OMZ, respectively, whereas echinoids are found near the core of this zone where oxygen levels are as low as 0.3 ml/l O₂. As these heart urchins are very abundant (14/m²), they represent a potentially significant component of the fossil record of OMZ's. Locally these urchins are capable of disturbing over 90% of near surface sediment. The core of the OMZ is inhabited by hermit crabs that actively transport and recycle gastropod shells. This 'biotransport' results in an accumulation of potentially preservable, non-endemic. hard-bodied organisms which may lead to misinterpretation of paleo-oxygenation conditions. We propose an alternative to the Rhoads & Morse (1971, Lethaia 4 ) biofacies model for open-ocean, dysaerobic environments which consists of: (1) a zone devoid of maeroinvertebrates. characterized by laminated sediments (<0.1 ml/l O₂); (2) a zone dominated by small (1–2 mm) soft-bodied infauca which exhibits moderate disturbance of laminae due to bioturbation (0.1–0.3 ml/l O₂); and (3) a zone inhabited by an abundant calcareous fauna characterized by highly bioturbated sediments (>0.3 ml/l O₂).     

Compostional variations and patterns of condont reworking in Late Devonian and early carboniferous calciturbidites (Moravia, Czech Republic)

Summary Compositional variations and grain-size properties of both carbonate constituents and conodonts as an alternative component group were used for interpreting the processes governing the deposition of upper Famennian and middle Tournaisian calciturbidites in Moravia, Czech Republic. Both the composition and grain-size properties of conodont element associations showed to be markedly dependant on facies type of their host sediment. Upper Devonian calciturbidite successions deposited on flanks of wide, Moravian-Silesian carbonate platform are composed mainly of echinoderm-and peloid-rich wacke/packstones and intraclastic float/rudstones (fine-grained calciturbidites, “normal” calciturbidites with Tab Bouma sequences, debris-flow breccias) with abundance of shelf-and shelf margin conodont taxa and epipelagic and “mesopelagic” conodonts. Upper Devonian calciturbidites deposited on slopes of volcanic sea-mounts are composed of echinoderm-and peloid-rich wacke/packstones and float/rudstones with increased proportion of intraclasts and volcanigenic lithoclasts (fine-grained calciturbidites, normal calciturbidites), yeilding abundant conodont associations with higher proportion of “mesopelagic” taxa compared to the platform-flank examples. Middle Tournaisian calciturbidite succession composed of crinoid-, peloid-, intraclast-and lithoclast-rich lime mudstones, wacke/packstones and float/rudstones (normal calciturbidites and debris-flow breccias) yielded conodont element associations rich in shelt-and shelf-margin taxa, “mesopelagic” conodonts and reworked Middle-and Upper Devonian conodonts. In general, the ratio of shelf-and shelf margin conodont taxa to “mesopelagic” taxa is distinctly lower in finegrained calciturbidites than it is in normal calciturbidites and debris-flow breccias. Grain-size properties (mean grain size and sorting) and percentage of fragmented conodont elements, too, are markedly dependant on the facies type: in fine-grained calciturbidites the values of mean grain-size and fragmentation are low and the sorting is good to very good whereas in normal calciturbidites and debris-flow breccias the values of mean grain-size and fragmentation are distinctly higher and the sorting is poorer. The interdependence of facies type and composition and grain-size properties of conodont element associations in gravity-flow deposits is explained as resultant from hydrodynamic sorting during turbidity current flow and final deposition of the bed. Compositional variations observed in our sections may thus be attributed to facies variability (coarsening-and thickening-upward trends) rather than to sea-level fluctuations (highstand shedding of carbonate platforms). On the other hand, significant enrichment in reworked conodont taxa in middle Tournaisian normal calciturbidites compared to scarcity and/or absence of such conodonts in essentially identical facies of upper Famennian age indicate sea-level to be the major control governing such compositional variations, with low relative sea-level stand in middle Tournaisian and high relative sea-level stand in upper Famennian. Thorough analysis of conodont evolution, palaeoecology and taphonomy, with emphasis on understanding the processes of deposition of their host rock, are recommended for any biostratigraphic and biofacies study to be done in carbonate sediments deposited under strong hydrodynamic regimes, such as calciturbidites, temperstites, debris-flow deposits, shelf-edge oolitic sands, tidal-channel facies etc.     

Trilobite biofacies and sequence stratigraphy: an example from the Upper Ordovician of Oklahoma

There have been surprisingly few empirical investigations of the fundamental principle that the architecture of depositional sequences exerts considerable control on observed patterns of faunal distribution and replacement. In this paper, we examine trilobite associations in two sequences of the Upper Ordovician (Sandbian) Bromide Formation of southern Oklahoma. Cluster analysis and ordination of genus abundance data identified five lithofacies‐related biofacies that are also differentiated by diversity patterns. Biofacies of the transgressive system tract (TST) of successive sequences are more similar to each other than they are to biofacies in the highstand systems tract (HST) of the same sequence. This similarity likely records dominance of large, robust convex sclerites in taphonomically degraded samples from condensed, strongly winnowed grainstone and rudstone. Horizons with articulated exoskeletons of isoteline trilobites preserved by obrution deposits occur most commonly in the early HST and record behavioural aggregations. Grainstone and rudstone of the later HST are less winnowed than those of the TST and show less fragmentation and sorting of sclerites. These changes in taphonomic conditions preserve ecological patterns more clearly. In most biofacies, rarefied alpha diversity (samples) and gamma diversity (biofacies) of middle‐ and outer‐ramp HST deposits are greater than in the TSTs, and biofacies replace each other down ramp. Diversity patterns do not agree with model predictions and other data sets that indicate low beta and high alpha diversity in the TST, likely because of taphonomic degradation. Vertical replacement of biofacies is expressed by the appearance of peritidal facies in which trilobites are rare. Biofacies shifts also characterize sequence boundaries and are most profound in the inner‐ramp successions characterized by sharp facies offsets. Comparison with bathymetrically similar deposits in the Taconic foreland basin showed similar diversity trends along environmental gradients, with some differences in shallow‐water settings attributed to taphonomic differences.     

Late Devonian - Early Carboniferous vertebrate microremains from the Carnic Alps, northern Italy

Vertebrate microremains from the Late Devonian-Early Carboniferous of the Carnic Alps are predominantly chondrichthyan, with minor placoderm and actinopterygian remains. The faunas are sparse and, with very few exceptions, occur only in conodont-rich pelagic limestones (Pramosio Limestone) representative of the palmatolepid-bispathodid conodont biofacies. Phoebodont and jalodont chondrichthyans, also reflecting open-ocean environments, predominated during the Famennian, and eventually symmoriids seem to predominate during the Early Carboniferous. The presence of Siamodus in this assemblage gives a new locality for this genus known from few regions in the world and allows confirming its stratigraphical range (limpidus Zone) and its relation to deep-water environments. The Late Devonian vertebrate faunas are tropical and cosmopolitan, having much in common with coeval taxa from the North-Gondwanan margins and Asian terranes. Composition of the vertebrate faunas is consistent with the Carnic Alps terrane having occupied a position intermediate between Gondwana and Laurussia, as hypothesized by various authors, but because of sparsity of the taxa represented and the pronounced cosmopolitan nature of both the conodont and vertebrate faunas, the data are not compelling.     

Sedimentary and biofacies records in calciturbidites at the Devonian-Carboniferous boundary in Moravia (Moravian-Silesian Zone, Middle Europe)

Summary At the Devonian/Carboniferous boundary, major climatic and oceanographic changes influenced sedimentation on carbonate platforms and in peri-platfrom asreas. Three deep-water carbonate successions in Moravia, which were selected to represent different paleotectonic settings, have been studied with the aim of testing the influence of eustatic, climatic and tectonic controls on sedimentation and conodont paleoecology and taphonomy. On the slopes of the wide carbonate platforms of the Moravian Karst Development (Lesní lom and Grygov sections), an exemplary highstand shedding systems developed in the upper Famennian (expansa Zone), marked by a pronounced thickness of their respective calciturbidite successions and an abundance of shallow-water skeletal grains.Palamatolepis— andBispathodus-dominated conodont assemblages contain an admixture ofPolygnathus representing a transported, near-shore component. The eustatic sea-level fall in the praesulcata Zone and the lowstand conditions at the D/C boundary resulted in a decline of carbonate platform production and condensed deposition or nondeposition. In the Lesní lom section, a condensed sequence of turrbiditic calcarenites and shales (Middle praesulcata—lowermost sulcata Zone) was followed by lime mud calciturbidites (sulcata and duplicata Zones). In the conodont assemblages, the first event in the Lower praesulcata Zone was associated with the reduction of ‘mesopelagic’Palmatopic and a bloom of epipelagicPolygnathus communis. The second event in the Middle praesulcata Zone corresponds to the onset of polygnathidprotogranthodid biofacies, indicating a carbonate slope environment. In the Grygov section, a pronounced thickening and upward-coarsening succession of tubiditic calcilutites through calcarenites and intraclast breccias, with poor palmatolepid-bispathodid connodont assemblages (expansa Zone), indicates a progradation of the calciturbidite system associated with sea-level highstand. After a break in sedimentation, covering the interval from the Lower praseulcata to the base of Lower crenulata Zone, thick-bedded, fine-grained calciturbidites were deposited in the Lower crenulata Zone, and are associated with poor, mixed assemblages where siphonodellids and polygnathids predominate. At the isosticha-Upper crenulata/Lower typicus boundary, coasre grained, turbiditic calcarenites and breccias rich in clastic quartz grains and mixed conodont assemblages with reworked Frasnian and Famennian conodonts indicate a deep erosion of the source area, presumably due totectonic uplift (relative lowstand). In the Jesenec section, on the flanks of the volcanic seamount (the Drahany Development), a deep-water Upper Famennian condensed succession of calciturbidites and presumably winnowed pelagic limestones is marked by conodont assemblages of palmatolepid-bispathodid biofacies. More proximal calciturbidites with mixed deep-water and shallowwater conodonts prograde at the top of the Upper Famennian succession (Middle to Upper expansa Zone). A striking hiatus, covering the interval from the Early preaesulcata to the base of Lower crenulata Zone, resulted from extreme condensation and submarine bottom current erosion due to sea-level lowstand in the late Famennian and early Tournaisian. The renewed middle Tournaisian calciturbidite sedimentation with strong evidence of erosion at the source area indicates global eustatic rise and tectonic uplift of the Drahany Development seamounts (relative lowstand). The earlier occurrence of the uplift in the Jesenec area, relative to the Grygov section, shows the advance of tectonic processes over time in the Moravian-Silesian basin (orogenic polarity) as a consequence of Variscan orogenic movements.     

Pleistocene marine depositionalenvironments from Mauritius island,Indian Ocean

The Pleistocene elevated marine carbonate complexes on Mauritius island correspond to three types of depositional zones: (1) reef-crest zone, (2) backreef zone, (3) littoral zone.On reef crests, the primary frame-builders are scleractinian corals (Acroporidae, Faviidae, Poritidae) and crustose coralline algae. The secondary builders are encrusting Foraminifers (Homotrematidae) chiefly, Molluscs (Vermetidae), Bryozoans (Cheilostomata) and Serpulids. Associated internal sediments are coarse to fine sand-sized grainstones or packstones and wakestones. In decreasing order of abundance, skeletal elements include fragments of corals, branched and crustose red algae, Pelecypods, Gastropods, benthonic Foraminifera (Amphisteginidae, Peneroplidae, Calcarinidae, Nummulitidae, Alveolinidae and Homotrematidae), Echinoderms and Alcyonarians; the most abundant mud-sized grains are identified as tunicate spicules. Primary marine cements are needle, palisade, micrite and pelletal ones.The backreef zone is characterized by coarse tomedium grainstones, packstones and wakestones. In addition to pellets the most common components are skeletal in nature; they broadly show the same mode of occurrence as the reef-crest unit. Primary cements locally occur in packed fibrous or micrite form.In littoral areas, well-sorted grainstones to poorlysorted mudstones can be described. The distribution of the various categories of biogenic particles is dependent upon the extent of reef tracts; ancient beaches from narrow reefs are characterized by coralgal facies while consolidated muddy banks in large complexes display high amount of molluscan detritus. Primary cementation appears mainly to be the result of the precipitation of fibrous or micrite calcite.The morphology, biological associations andsediments of these Pleistocene limestones are homologous with those of the adjacent modern environments.     

Emsian (Early Devonian) Yujiang Event in South China

The Yujiang Event is reported here as a newly recognized event that occurred during the deposition of the Early Devonian Yukiang Formation in Liujing, Guangxi, and its equivalents in South China. The first episode of the Yujiang Event is characterized by the extinction of biostrome that is constructed mainly by colonies of rugose coral, tabulatimorph coral and bryozoan as well as the extinction of evolved biota including some brachiopods, bivalves, etc. The age of the first episode of the Yujiang Event dated by conodonts is at the beginning or the lower part of the Polygnathus nothoperbonus Zone. The main episode of Yujiang Event is evidenced by the extinction of the Rostrospirifer tonkinensis Fauna (sensu lato) at the top of Yukiang Formation and by the sudden occurrences of the dolostones and dolomitic limestone of the overlying Moding Formation. At the basal part of the Moding Formation, the new finding of conodont Polygnathus nothoperbonus and dacryconarids Nowakia (N.) barrandei indicates that this episode of Yujiang Event may occur in the upper part of the Polygnathus nothoperbonus Zone. Overlying the Yukiang Formation and its equivalent beds, various lithic deposits correspondingly yielding different biota can be seen widely in many localities and sections in South China and northern Vietnam. Therefore, the Yujiang Event can be recognized not only as a biotic extinction event, but also as a geological event.     

Microhermal nodules of <Emphasis Type="Italic">Renalcis</Emphasis>-like calcimicrobes from Oxfordian limestones of the Sierra Madre Oriental (Novillo Formation,Mexico)

Microhermal nodules very similar to those from the Oxfordian Smackover Formation are volumetrically important constituents of the Novillo Formation exposed in eastern Central Mexico. The nodules occur within a micritic limestone succession (Novillo Limestone). Coalescence of adjacent nodules leads to a delicate self-supported frame. Microhermal nodules consist of both microbial carbonate and clustered digitate and branching chambered microfossils. The latter occur in the form of crust-like agglomerations or bushy growth habit in small growth-framework cavities created by accretionary microbialites. Main growth patterns are branching upward, downward facing and pendant. Like Renalcis, the microfossils apparently resulted from the activity of calcimicrobes. Terebella, encrusting foraminifers, and Tubiphytes form part of the microhermal biota and occur as subordinate faunal components. The association developed on the floor of a calm, shallow-marine lagoon with restricted water interchange and reduced oxygenation at the sediment–water interface.