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.     

Taphonomy and compositional fidelity of Quaternary fossil assemblages of terrestrial gastropods from carbonate-rich environments of the Canary Islands

Quaternary aeolian deposits of the Canary Islands contain well‐preserved terrestrial gastropods, providing a suitable setting for assessing the taphonomy and compositional fidelity of their fossil record over ~13 kyr. Nine beds (12, 513 shells) have been analysed in terms of multivariate taphonomic and palaeoecological variables, taxonomic composition, and the stratigraphic and palaeontological context. Shells are affected by carbonate coatings, colour loss and fragmentation. Shell preservation is size‐specific: juveniles are less fragmented and show colour preservation more commonly than adults. In palaeosols, the adult shell density correlates negatively with the proportion of fragmented adults, negatively with the proportion of juveniles, and positively with the proportion of adults with coatings. High bioturbation intensity in palaeosols is associated with low shell fragmentation and high proportion of shells with coatings. These relationships imply that high adult density in palaeosols was driven by an increase in shell production rate (related to a decrease in predation rates on adults and a decrease in juvenile mortality) and a decrease in shell destruction rate (related to an increase in durability enhanced by carbonate precipitation). In dunes, the relationships between taphonomic alteration, shell density and bioturbation are insignificant. However, dune assemblages are characterized by a lower frequency of shells with coatings and higher rates of colour loss, indicating lower shell durability in dunes than in palaeosols. Additionally, non‐random differences in the coating proportion among palaeosols imply substantial temporal variation in the rate of carbonate crust formation, reflecting long‐term changes in bioturbation intensity that covaries positively with shell preservation. Dunes and palaeosols do not differ in species abundances despite differences in the degree of shell alteration, suggesting that both weakly and strongly altered assemblages offer data with a high compositional fidelity. Carbonate‐rich terrestrial deposits originating in arid conditions can enhance the preservation of gastropods and result in fossil assemblages that are suitable for palaeoecological and palaeoenvironmental studies of terrestrial ecosystems.     

Factors controlling upper Jurassic ammonite assemblages in north-central Mexico

Lower Kimmeridgian to Lower Tithonian (Upper Jurassic) sections studied at Sierra de Palotes (Durango) and Sierra de Catorce (San Luis Potosí), Mexico, show low-energy deposits in which the composition of fossil macroinvertebrate assemblages, including megabenthos, reflects biostratinomic control. Monotonous siltstones provide continuous records of ammonite assemblages and reflect dominant deposition of shells in living areas; meanwhile, discontinues records were forced by episodic post-mortem transportation of shells, which was especially accentuated under storm influence. Rhythmic marly-silty limestones and marls illustrate a fossil record probably determined by minor transgressive-regressive pulses. The major changes in lithofacies are reflected by condensed silty and phosphatic mudstones deposited during significant floodings affecting areas under dominant terrigenous sedimentation. These changes determined more or less significant variations in the composition of fossil assemblages according to their relation to changing ecological conditions. However, shifting ecospaces exhibit no direct relationship to changes in lithofacies. Post-mortem transportation, operating in relation to both marine floodings and changes in the pattern of upper-water currents, was the main biostratinomic factor affecting the areal distribution of ammonite populations. Shell transportation and sedimentation rate controlled preservation and ultimately influenced diversity in recorded ammonite assemblages. The post-mortem behaviour (interpreted from shell structure and preservation), and therefore distribution, of ammonite shells points to shallow-water environments during the Kimmeridgian - Early Tithonian in areas (such as SE Durango and San Luís Potosí) close to the changing boundary between dominant carbonate and terrigenous sedimentation. No reworking affecting ammonite biostratigraphy has been identified in the sections studied.     

Microfacies and depositional environment of an Upper Triassic intra-platform carbonate basin: the Fatric Unit of the West Carpathians (Slovakia)

Facies associations of the Rhaetian Fatra Formation from the Veká Fatra Mts. (West Carpathians) were deposited in a storm-dominated, shallow, intra-platform basin with dominant carbonate deposition and variable onshore peritidal and subtidal deposits, with 21 microfacies types supported by a cluster analysis. The deposits are formed by bivalves, gastropods, brachiopods, echinoderms, corals, foraminifers and red algae, ooids, intraclasts and peloids. A typical feature is the considerable variation in horizontal direction. The relative abundance and state of preservation of components as well as the fabric and geometric criteria of deposits can be correlated with depth/water energy-related environmental gradients. Four facies associations corresponding to four types of depositional settings were distinguished: a) peritidal, b) shoreface, above fair-weather wave base (FWWB), c) shallow subtidal, above normal storm wave base and d) above maximum storm wave base. The depositional environment can be characterized as a mosaic of low-relief peritidal flats and islands, shoreface banks and bars, and shallow subtidal depressions. The distribution and preservation of components were mainly controlled by the position of base level (FWWB), storm activity and differences in carbonate production between settings. Poorly or moderately diverse level-bottom macrobenthic assemblages are dominated by molluscs and brachiopods. The main site of patch-reef/biostrome carbonate production was located below the fair-weather wave base. Patch-reef/biostrome assemblages are poorly diverse and dominated by the branched scleractinian coral Retiophyllia, forming locally dm-scale autochthonous aggregations or more commonly parautochthonous assemblages with evidence of storm-reworking and substantial bioerosion by microborings and boring bivalves.Facies types and assemblages are comparable in some aspects to those known from the Upper Triassic of the Eastern and Southern Alps (Hochalm member of the Kössen Formation or Calcare di Zu Formation), pointing to similar intra-platform depositional conditions. The absence of large-scale patch-reefs and poor diversity of level-bottom and patch-reef/biostrome assemblages with abundance of eurytopic taxa indicate high-stress/unstable ecological conditions and more restricted position of the Fatric intra-platform setting from the open ocean than the intra-platform habitats in the Eastern or Southern Alps.     

Taphonomy and taphofacies models of the Pliocene deposits of Vejer de la Frontera (Cádiz, SW Spain)

A quantitative taphonomic analysis of the fossil assemblages, together with a sedimentary study of the Pliocene deposits of the Vejer Basin (Cádiz, Spain), has been carried out. This multidisciplinary analysis has allowed us to establish with great precision the palaeoenvironmental conditions under which these materials were formed. The Pliocene deposits can be divided into three units, named from bottom to top: Unit 1, Unit 2 and Unit 3. Unit 1 is formed of medium-to-coarse-grained sands at the base, corresponding to subtidal bar deposits. At the top, the sands become coarser, and beach and littoral dunes were formed. Within these sedimentological contexts, the fossil remains exhibit a strong degree of taphonomic destruction due to high hydrodynamic energy. Unit 2, consisting of clays, corresponds to deposits formed in a protected coastal bay. A low-energy environment allowed the fossils to maintain a good state of conservation and even thin-shelled bivalves with a delicate ornamentation are preserved. Unit 3 comprises highly bioclastic sands and calcarenites-calcirudites. This unit displays a shallowing upwards trend, passing from deposits generated under shallow subtidal conditions in the lower part to materials deposited on beaches and coastal dunes in the upper part. Under these conditions, the fossil remains show a high degree of taphonomic destruction. In Units 1 and 3, the fossils are mainly accumulated in horizontal beds, with extensive lateral continuity and with erosive bottoms and graded tops. These accumulations of fossils are interpreted as tempestites. As compared with the fossils preserved in surrounding sediments, the taphonomic attributes of these bioclasts suggest a superposition of the background and event processes. During normal conditions, the high energy of the littoral environments produced a strong degree of taphonomic destruction. During storm events, a mixture of taphonomic attributes was produced: high fragmentation and disarticulation and little abrasion and edge rounding of the fragments. This mixture was a consequence of sudden burial after storms, which prevented long periods of exposure of the bioclasts in the taphonomically active zone. Three taphofacies models are proposed, based on taphonomic attributes of fossils contained in the different units: inner-shelf taphofacies, protected shallow-lagoon taphofacies and storm bed taphofacies.     

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.     

Development of rudist lithosomes in the Coniacian–Lower Campanian carbonate shelves of central-southern Italy: high-energy vs low-energy settings

The Late Cretaceous shallow-water depositional areas of southern Tethys were complexes of unprotected shelves occupied by foramol assemblages that produced loose, diagenetically stable bioclastic debris not involved in significant in situ cementation processes. Both storm- and wind-induced currents and waves exercised a strong control on the distribution of the shifting biogenic sediments which covered the open sea-floor, constituting large coalescing sheets of winnowed fine to coarse skeletal sands. Rudists spread over all shelf sectors, from more open and external areas to more internal ones, occupying different substrata and furnishing the bulk of the skeletal component by means of bioerosion processes. They colonised mobile sediments giving rise to complex bodies with peculiar characteristics related to environmental parameters of the different sectors of the shelf. On the basis of detailed sedimentological, taphonomic and palaeontological data, we recognised two main rudist-rich depositional settings (‘end members’) in the southern Italy Senonian rudist-bearing successions. In successions pertaining to hypothesised marginal shelf sectors, characterised by high-energy regime deposits, rudist lithosomes are metric in thickness and lateral extent and lens-like in morphology, rich in bioerosion-derived skeletal sand and silt. Rudists are highly diversified. Large elongated cylindro-conical hippuritids (mostly pertaining to the genera Hippurites and Vaccinites), thick-shelled radiolitids and plagioptychids largely dominate. Rudists clustered in life position are subordinate; they often form small bouquets. More commonly these organisms appear fallen but only barely reworked. The rudist-rich bodies laterally pass into clean bioclastic grainstone in which sedimentary structures, related to current and/or storm erosional action, are common. No evidence of significant original relief of the rudist bodies in respect of the neighbouring sediment can be recognised. The submarine erosion and/or the high-energy processes operating presumably inhibited the aggradation of the tidal sediments above the marginal ones. As a consequence the vertical facies organisation shows widespread subtidal cycles, as commonly recognised in open shelves with ramp-like morphologies. In successions pertaining to more internal and/or low-energy sectors, rudist-rich beds rhythmically alternate with finer-grained foraminiferal limestones. Small elevator radiolitids with oligospecific diversity are dominant, mostly concentrated in clumps. Rudists in growth position are abundant, although a large quantity of shells appear toppled with little reworking. They may form laterally continuous biostromal shell beds. Sedimentary structures such as cross-lamination and gradation are only occasionally present. The resulting facies are commonly arranged in peritidal/shallow subtidal cycles in which evidence of subaerial (up to pedogenic modifications on a large and small scale) and, less frequently, submarine exposure is common. Intermediate successions have been recognised, characterised by deposits of silty-sand plains, which present intercalations of graded, bioclastic, storm-related beds. Sedimentological characteristics seem to document more open conditions in which submarine erosion was intermittently prevalent. In these successions rudist species that are commonly found both in high-energy and low-energy assemblages coexist.     

Anthropogenic effect recorded in the live-dead compositional fidelity of land snail assemblages from San Salvador Island, Bahamas

Terrestrial malacofaunas that inhabit islands are vulnerable to human activities. Habitat destruction, introduction of exotic species, predators, etc. are distorting the composition and distribution of indigenous snail communities. Specifically, the taxonomic discordance between live and dead assemblages may be the consequence of anthropogenic disturbances rather than natural post-mortem processes. Live-dead fidelity may hence reflect the degree of human alteration in a given locality. This approach was used to study the relative abundance of live and dead land snails from pristine and urbanized localities around San Salvador Island (Bahamas). Thirteen species were encountered from 64 samples containing 5,343 specimens. Taxonomic composition was significantly different between areas of negligible anthropogenic disturbance and those impacted by humans. Although which species of land snails are endemic to San Salvador is unknown, all identified species are native to the Caribbean region and invasive species were not encountered. However, some synanthropic species, found exclusively in urban-developed areas, have not been recovered from the local fossil record, pointing to their relatively recent introduction possibly from nearby islands. Non-metric MDS, Jaccard-Chao index, and Spearman correlation analyses indicated that dead assemblages displayed a good compositional correspondence to live communities at pristine sites, whereas considerable disparity was sometimes observed at human-modified areas. Urbanized areas probably favored the proliferation of synanthropic species whereas dead assemblages may contain anthropophobic taxa that lived there prior to recent human modification. These findings are consistent with previous live-dead fidelity studies of marine molluscan and terrestrial small-mammal assemblages, and suggest that the taxonomic discordance between live and dead assemblages of land snails may be indicative of recent anthropogenic alteration.     

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.     

Two contemporaneous equilibrium communities in the Ordovician of Wisconsin

Two statistically different fossil assemblages were recognized in the Mifflin Submember of the Platteville Formation. These are believed to be ecologic communities because of their recurrence over a large area, their well-defined trophic structure, and their concurrence with lithofacies boundaries. The Hesperorthis—Eoleperditia community, which roughly coincides with the limestone facies of the Mifflin, has a diverse brachiopod fauna, abundant ostracodes, and several forms of trilobites and gastropods. The Hesperorthis-Sinuites community, coincident with the Miffin dolomite facies, contains abundant. but less diverse, brachiopods, and fewer trilobites and ostracodes. Gastropods and cephalopods are more abundant and diverse, and corals and bivalves are present. In both communities the most abundant genera were non-competitive in terms of trophic resources. and each trophic group was dominated by one or two genera. The two communities are interpreted to be equilibrium associations based on their vertical persistence through the section. Lithologic and faunal evidence indicates that these communities inhabited a quiet-water open-shelf environment which was occasionally disturbed by physical reworking (storm?) events. The distinction of two separate communities within an overall rather homogeneous environment suggests that the control was not physical energy or substrate type but a non-preservable factor such as food supply.     

Seafloor erosion and sea-level change: Early Jurassic Blue Lias Formation of central England

Minor bedforms within the mudstone-dominated Early Jurassic Hettangian Saltford Shale Member (Liasicus up to Angulata Chronozone) of the Blue Lias Formation in central England, indicate weak seafloor erosion in a mid to outer ramp setting. Distal storm flows below maximum storm wave base are proposed as the most likely generative mechanism for silty scour and gutter casts that enclose concentrations of well-preserved schlotheimiid ammonites and arthropod trace fossils. Within the upper part of the Saltford Shale (probably Angulata Chronozone), a discrete layer of reworked and bioencrusted limestone nodules signifies an episode of more persistent seafloor erosion. The immediately overlying strata, transitional to the Hettangian–Sinemurian Rugby Limestone Member, are relatively bioturbated and feature fossils of macrobenthos, as well as shell concentrations resembling relatively proximal storm beds. This suggests that the reworked nodule horizon marks sea-level fall, rather than stratigraphic condensation associated with sediment starvation. The biostratigraphic evidence raises the possibility that this erosional episode correlates with a mid-Angulata Chronozone hiatus documented from the Wessex Basin, southwest England. Equally however, it could be linked to contemporaneous movement on one or more nearby faults, affecting the southern part of the English East Midlands Shelf.     

Reworked carboniferous spores: An example from the lower jurassic of northeast Scotland

An association of contemporaneous Jurassic mega- and miospores with reworked Carboniferous mega- and miospores is described from the Lower Lias rocks of northeast Scotland. The assemblage is compared with other accounts of reworked spores in Jurassic sediments. It is suggested that such reworking was particularly common in the Lower Jurassic of northern Europe and that four situations exist in the literature concerning probable cases of reworking: (1) the spores are recognised as reworked; (2) the identity of the spores as Carboniferous species is recognised and the “range” of these species is extended into the Jurassic; (3) the reworked spores are not recognised as such, and are incorrectly assigned to Jurassic genera and species; (4) the spores are recognised as Carboniferous, but their origin is attributed to contamination rather than reworking. In conclusion, the contrasting preservational features exhibited by the reworking and contemporaneous megaspores are discussed and interpreted as the result of differing depositional and postdepositional histories.     

Shell Beds as paleoecological puzzles: A case study from the Upper Permian of the Paraná Basin, Brazil

Summary Microstratigraphic, sedimentological, and taphonomic features of the Ferraz Shell Bed, from the Upper Permian (Kazanian-Tatarian?) Corumbataí Formation of Rio Claro Region (the Paraná Basin, Brazil), indicate that the bed consists of four distinct microstratigraphic units. They include, from bottom to top, a lag concentration (Unit 1), a partly reworked storm deposit (Unit 2), a rapidly deposited sandstone unit with three thin horizons recording episodes of reworking (Unit 3), and a shell-rich horizon generated by reworking/winnowing that was subsequently buried by storm-induced obrution deposit (Unit 4). The bioclasts of the Ferraz Shell Bed represent exclusively bivalve mollusks.Pinzonellaillusa andTerraia aequilateralis are the dominant species. Taphonomic analysis indicates that mollusks are heavily time-averaged (except for some parts of Unit 3). Moreover, different species are time-averaged to a different degree (disharmonious time-averaging). The units differ statistically from one another in their taxonomic and ecological composition, in their taphonomic pattern, and in the size-frequency distributions of the two most common species. Other Permian shell beds of the Paraná Basin are simílar to the Ferraz Shell Bed in their faunal composition (they typically contain similar sets of 5 to 10 bivalve species) and in their taphonomic, sedimentologic, and microstratigraphic characteristics. However, rare shell beds that include 2–3 species only and are dominated by articulated shells preserved in life position also occur. Diversity levels in the Permian benthic associations of the Paraná Basin were very low, with the point diversity of 2–3 species and with the within-habitat and basin-wide (alpha and gamma) diversities of 10 species, at most. The Paraná Basin benthic communities may have thus been analogous to low-diversity bivalve-dominated associations of the present-day Baltic Sea. The ‘Ferraz-type’ shell beds of the Paraná Basin represent genetically complex and highly heterogeneous sources of paleontological data. They are cumulative records of spectra of benthic ecosystems time-averaged over long periods of time (10²–10⁴ years judging from actualistic research). Detailed biostratinomic reconstructions of shell beds can not only offer useful insights into their depositional histories, but may also allow paleoecologists to optimize their sampling designs, and consequently, refine paleoecological and paleoenvironmental interpretations.     

Phylogenetic implications and diagenetic stability of macromolecules from pleistocene and recent shells of Mercenaria mercenaria (mollusca,bivalvia)

The phylogeny and diagenesis of Pleistocene and Recent bivalves were studied immunologically by use of a conventional antiserum elicited against an EDTA‐soluble macromolecular extract from shells of the modern bivalve mollusc Mercenaria mercenaria. ELISA tests of the antiserum with shell fragments of a wide range of modern bivalves gave taxonomically significant results. The antiserum reacted with palaeoheterodonts and heterodonts but not with representatives of other bivalve subclasses. This phylogenetic reactivity was also apparent in tests with fossil shells, although the specificity and overall strength of the reaction were both reduced. Absorption of the antiserum with etched shell powders of various (palaeo)heterodonts yielded more specific antibody preparations.

Investigations of shell matrix diagenesis, using the anti‐Mercenaria serum, demonstrated that small amounts of original determinants could be detected even in fossils over one million years old. The reactivity of the serum with extracts of fossil Mercenaria decreased with sample age. The relationship between serum reactivity and the degree of amino acid racemization was almost linear. Clearly, the various determinants to which antibodies were elicited were being destroyed at different rates.     

Coralline red algal assemblage from the Middle Pliocene shallow-water temperate carbonates of the Monte Cetona (Northern Apennines,Italy)

During the Pliocene and Pleistocene, the Monte Cetona (Northern Apennines, central Italy) was part of an elongated island. The Middle Pliocene deposits around the Monte Cetona are represented by shallow-water marine carbonates rich in coralline red algae and bryozoans. These skeletal carbonates, characterising a coralline algal-dominated factory, were analysed in terms of microfacies, taxonomy, and growth-forms of coralline red algal assemblage. Three microfacies were distinguished on the basis of component distribution and fabric analysis: coralline algal rudstones, coralline algal floatstones, and bioclastic packstones. Skeletal components are commonly abraded, bioeroded, and encrusted. The shallow-water skeletal carbonates are strongly bioturbated and any primary sedimentary structure is obliterated. The distribution of the coralline growth-forms suggests a decreasing hydrodynamic gradient from the coralline algal rudstone, through the coralline algal floatstone to the bioclastic packstone microfacies. The coralline algal flora consists of eight species representing the subfamilies Lithophylloideae, Mastophoroideae and Melobesioideae. The assemblage is dominated by lithophylloids. Other biogenic components are bryozoans, barnacles, echinoderms, and benthic foraminifera. These coralline algal assemblages were deposited just above the fair-weather wave base and indicate a shallow-marine temperate water setting for the eastern Tyrrhenian Sea during the Mid Pliocene.     

Ammonite taphonomy and stratigraphy of the Bajocian at Maizet,south of Caen (Calvados,NW France)

A study of the ammonite assemblages from the “Oolithe ferrugineuse de Bayeux” Formation of Bajocian age is presented herein. The section at the locality of Maizet shows a high level of stratigraphic condensation, and taphonomic reworking is common within the sequence. All the ammonites being studied are classified herein as having been reworked, transported or displaced on the sea-floor prior to burial, and as such, are determined taphonomically as being resedimented or reelaborated fossil elements. Seven evidences of reelaboration within the sequence under investigation are detailed here. The palaeontological units, so-called taphorecords, characterized by distinctive taphonomic features, are used to directly or indirectly assign beds to biochronostratigraphic units. In addition, identification of taphorecord relationships regarding successive or contemporaneous deposition allows their registratic succession and order of depositional events to be inferred. The latter may be deduced on the basis of bed succession or by reference to stratigraphical intervals that now are only represented in the stratigraphic column by fossils. Deposition by tractional currents and winnowing is indicated by sharp bed-base and by reworked fossil elements. Biochronostratigraphic correlation with other sections of the inland Bajocian successions in the Calvados area (Bretteville, Feuguerolles) highlights a common depositional evolution that may be related to an eastward-deepening carbonate ramp.     

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.     

What's in a beach? Soil micromorphology of sediments from the Lower Paleolithic site of 'Ubeidiya, Israel

A micromorphological study of archaeological sediments from the early Pleistocene site of 'Ubeidiya (Jordan Valley, Israel) was conducted to provide microenvironmental detail for the hominin occupation contexts and investigate site formation issues. Previous research shows that the hominin groups occupied the marshes and pebbly beaches at the shores of a lake during a regressive period, but given that some portions of the lithic and faunal assemblages are abraded and others fresh, there remains a question of whether the archaeological assemblages are in situ or reworked, and if reworked, by what mechanisms and from where. The rates of sedimentation within the regressive cycle, by which we can learn about the frequency and duration of exposed surfaces amenable for hominin occupation is also unknown. Finally, the artificial nature of some of the pebbly layers has been questioned. The micromorphological analysis yielded the identification of twelve microfacies; the majority of these represent fluvially derived floodplain soils or distal mudflow deposits, and a minor number are sediments of lacustrine origin: mudflats and shallow subaqueous sediments. These represent the natural habitats of the 'Ubeidiya hominins and might serve as a reference to similar contexts of other early hominin sites. The sedimentary model proposed here entails the rapid deposition of fluvially derived low-energy sediments at and around the shoreline, followed by prolonged periods of exposure, during which surfaces stabilized within a relatively wet, marshy environment. This interpretation suggests that the abraded portions of the archaeological assemblages are a result of prolonged surface exposure rather than high-energy transport from a distant source or to wave reworking at the shoreline, and supports the consideration of these assemblages as archaeological palimpsests, with locally reworked fresh and abraded elements. No micromorphological evidence supporting anthropogenic agency in the formation of the pebbly layers was found. The entire regressive cycle entailed unvarying climatic conditions with seasonal fluctuations and episodic lacustrine incursions, and with a trend towards arid conditions in the end.     

Early Jurassic shell beds from marginal marine environments in southern Mendoza, Argentina

External estuarine facies in Late Sinemurian beds from the base of the Puesto Araya Formation in the Atuel river region, Mendoza Province, Argentina, contain concentrations of a new species of Cardinioides (a shallow burrowing suspension-feeding Pachycardiidae bivalve) and a low diversity benthic fauna. These bivalves appear in tidal inlet facies of a wave-dominated estuary (environment similar to lagoon-barrier island systems), forming bioclastic lags at the channel bases. The taphonomic attributes (such as shell articulation, degree of fragmentation and abrasion, bioclast size, orientation) change from North to South, from monospecific concentrations of nearly complete large Cardinioides shells with low degree of fragmentation and abrasion, through beds with smaller Cardinioides shells associated with a few bivalves from marine environments, to only isolated fragmented specimens associated with a fully marine biota including not only other bivalves but also brachiopods and ammonoids. This, together with different palaeoecologic, sedimentologic and stratigraphic attributes at the studied localities are evidence of southwards changing environmental conditions from marginal marine (brackish) to fully marine on the eastern margin of the Neuquén basin at that time. From detailed analysis of taphonomic features, size distribution of shells, and the associated fauna at the different localities, the new species is regarded as a brackish water endemic, possibly a low salinity euryhaline species, which inhabited well oxygenated waters. Salinity, turbulence levels and food supply were the main limiting factors in the distribution of the species.     

Patchiness of local species richness and its implication for large-scale diversity patterns: an example from the middle Miocene of the Paratethys

The best hope for understanding global diversity patterns is to compare local assemblages, which are mostly preserved in taphonomically-complex shell beds. The present study investigates the variability in faunal composition and diversity at the scale of a single outcrop. A total of 152 species (3315 shells) occurred in 25 samples from 5 tempestitic shell beds. Although sampling intensity was high, total species richness was not captured by far at the hierarchical levels present (outcrop, shell beds, samples) because the majority of species is rare. In contrast, sampling intensity was sufficient to cover the most abundant species, as indicated by stable evenness values. Four taxa dominate the assemblage, but their rank order differs strongly between individual shell beds and individual samples; significant differences between some shell beds are evident for faunal composition, and one shell bed differs from all others with respect to species accumulation curves. Within shell beds, rarefaction curves are generally characterized by strongly overlapping confidence intervals, but outliers occur in three of five shell beds. Patchiness is additionally indicated by a wide scatter of diversity indices in some shell beds and by a wide scatter of samples of one shell bed in an ordination on faunal composition. Most of the outcrop-scale variability in faunal composition and diversity can be related to differences between shell beds. This suggests that sampling a single shell bed of the outcrop is insufficient to characterize the local fauna and its diversity, even when sampling intensity (i.e. the number of samples and shells) within the shell bed was high. Similarly, a single sample from such a shell bed may not be sufficient to characterize its diversity, even when the number of counted shells was high. It is therefore confirmed that sampling strategy and sampling intensity are crucial to confidently characterize the shelly assemblages at such a small spatial scale and that dispersed sampling effort with many small replicate samples will characterize a local assemblage and its diversity better than a few large samples. Diversity comparisons of individual samples between localities must account for the high variability present at the smaller spatial scale, as observed in our study.