Taphonomy of Clypeaster fragments: preservation and taphofacies

The preservation of echinoid fragments of the common echinoid genus Clypeaster recovered from bulk sediment samples is used to explore the possibilities of recognizing taphonomic patterns and establishing taphofacies within a shallow water carbonate environment (Northern Bay of Safaga, Red Sea, Egypt). This is accomplished by the comparison and quantification of the taphonomic signatures of numerous fragments of roughly similar size. The main taphonomic processes observed are abrasion and destruction of plate surfaces, including tubercles and plate edges. Encrustation, especially by foraminifera, can be common and intense. Intraplate fracturing and surface features, such as scratch marks, were also quantified. Four different taphofacies with different characteristic signatures were separated using a cluster analysis of cumulative taphonomic grades. The distribution of taphofacies clearly reflects the topographic complexity of the study area and can be explained by ambient environmental parameters that reflect the general picture of wave and current intensities and sedimentation patterns. The potential use of taphofacies analysis as an independent measure of environmental heterogeneity in palaeoecological studies is confirmed in that taphofacies analysis can reveal parameters not immediately obvious when investigating the textural and biotic characteristics of a sediment.     

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.     

Taphofacies analysis of recent shelly cheniers (beach ridges), northeastern baja california,Mexico

Summary This report presents the results of taphofacies analyses of shelly cheniers (mollusk-dominated lag-concentrations) from the tidal flats of northeastern Baja California, Mexico. The three generations of moderm (formed during last 70 years), submodem (younger than 1,500 BP), and subfossil (5,000–2,000 BP) cheniers can be distinguished by their position relative to the shoreline, their topography, and the radiocarbon-age of their shells. The generations differ in the duration and complexity of their taphonomic history. Sixty-one samples from nine localities were collected to test the utility of the taphofacies approach for studying chenier-type shell deposits. The three chenier generations, although all dominated by the bivalve molluskMulinia coloradoensis, differ significantly in their taxonomic composition due to taphonomic and/or biologic factors. The taphofacies analysis included 4,334 specimens ofM. coloradoensis described by nine taphonomic variables. Univariate analysis of those variables indicated that the shells that accumulated in the cheniers are little-affected by biological processes (bioerosion, encrustation), and moderately affected by physical processes (fragmetation, cracking, peeling, edge preservation). Only the luster features of shells (external luster, internal luster, and internal features) vary substantially and consistently with chenier age —a result of subaerial weathering. Multivariate taphofacies analysis discriminates the three generations of cheniers even when the poorly preservable luster variables are excluded from the analysis. This suggests that taphofacies discrimination is possible for fossil cheniers. The shells collected from the chenier surface have substantially poorer preservation than shells from the subsurface, indicating that taphonomic degradation in the chenier plain environment is a surface phenomenon. Chenier plain shelly assemblages are taphonomically distinct from assemblages formed in other marine environments: they have a very low frequency of macroscopically recognizable bioerosion and encrustation. The existence of preservable taphonomic differences between the cheniers that differ in their age (i.e., duration of preburial history), suggests that fossil lag concentrations may be useful in detecting incompleteness gradients along stratigraphic boundaries. A ‘taphonomic clock’—a correlation between a ‘time-sincedeath’ and shell preservation—was found only for luster features, taphonomic attributes that are unlikely to be preserved in the fossil record.     

Taphonomic pathways and comparative biofacies and taphofacies in a Recent intertidal/shallow shelf environment

The development of taphonomic approaches to facies analysis requires a foundation in facies-based actualistic studies. Modern intertidal and shallow shelf environments at Provincetown Harbor. northern Cape Cod, Massachusetts (USA) provide an opportunity to compare pattcrns and controlling factors in molluscan biofacies and taphofacies distributions. Variation in faunal composition, ecologic variables, and taphonomic attributes of molluscan death assemblages produce distinct patterns of environmental zonation: (1) Faunal composition (biofacies) primarily tracks variation in substrate type among environments (sand, rock, peat, and Zostera marina beds). (2) Ecologic variables (equitability, infauna: epifauna ratio, gastropod:bivalve ratio, and predation on M. mercenaria) appear to reflect tidal exposure time. (3) Taphononic attributes (fragmentation, abrasionm, corrosion, bioerosion, and encrustation) of the common bivalve M. mercenaria track environmental energy, in particular its effects on the stability and reworking of hardparts at the sediment surface. Shells in different environments proceed along different taphonomic pathways - the order of acquisition of taphonomic features by hardparts. An encrustation/bioerosion-dominated pathway characterizes low energy environments; the upper intertidal and deeper subtidal. An abrasion-dominated pathway characterizes the high energy lower intertidal and shallow subtidal. Contrasting pathways produce distinct proportions of taphonomic attributes in time-averaged samples; proportions that delineate taphofacies. Integrated taxonomic, ecologic and taphonomic data provide a more complete picture of environmental processes than any approach alone. Taphonomic data not only furnish information not readily provided by other approaches, but free paleoecology from the constraints of taxonomic uniformitarianism. □Taphonomy, comparative taphonomy, taphofacies, biofacies, cluster analysis, multidimensional scaling, taphonomic pathways, Recent, actualism, intertidal, molluses.     

Depth as an overarching environmental variable modulating preservation potential and temporal resolution of shelly taphofacies

In the marine realm, the interpretation of taphofacies relies heavily on how oceanographic and sedimentary conditions affect the preservation state of fossils. Several taphonomic variables either covary with depth or are directly influenced by depth. Facies‐level factors rather than broad, basin‐scale parameters influence the taphonomic profile of mollusc death assemblages according to actualistic and experimental evidence. To determine the possible relation between depth and the taphonomic conditions of multiple species of bivalve remains, we used seven samples gathered over a comprehensive bathymetric gradient (from 7 to 150 m below mean sea level; topmost 10‐ to 20‐cm layer, roughly corresponding to the taphonomically active zone). We selected samples from predominantly muddy facies on the southern Brazilian shelf (SBS). The taphonomic damage profile (TDP) was measured using site samples based on a standard taphonomic analysis (categorical scoring system) of shells and fragments larger than 4 mm, to identify site damage patterns. Restricting the sedimentary grain size (samples from fine sediments) enabled the determination of the variation in damage with depth among the samples. Constrained analysis of proximities (CAP) revealed that up to 46% of the taphonomic variation observed was related to variation in depth (with approximately 28% unexplained by environmental factors). Part of the unexplained fraction was due to the effect of temporal mixing, which is predictable along large‐scale patches but is inversely linked to the TDP. Our results show that taphonomic analysis, considering large spatial scales in recent environments, can explain the variations present in shell beds that formed during distinct time periods of the evolution of a Quaternary sedimentary basin.     

Taphonomy of Eemian marine molluscs and acorn barnacles from eastern Arkhangelsk region, northern Russia

Marine Eemian deposits along the Pyoza river and its tributary Varchuska, Arkhangelsk region, constitute successions of muddy and sandy facies with rich macrobenthic fauna dominated by bivalves and barnacles. Taphonomic features formed by abrasion, disarticulation, dissolution, fragmentation, bioerosion and encrustation define taphofacies for a palaeoenvironmental model. Five bivalve taphofacies and three barnacle taphofacies could be distinguished. Both bivalves and barnacles are poorly preserved in foreshore/shoreface environments, as the shells were subjected to extensive transportation by currents. The shells were best preserved in offshore environments, where rapid episodic sedimentation enabled within-habitat preservation, in some cases even preservation in life position. Barnacles are absent from the most clay-rich offshore deposits, probably because of clogging of filters by turbidity and lack of suitable substrate. Such dissimilarities suggest that the number and distribution of taphofacies may depend on which fossil groups are used. Interspecific variability may exist within the individual taphofacies. The barnacles, for example, tend to be better preserved than the mussel Mytilus edulis, although both are fixosessile suspension feeders. This indicates that not only life habits but also intrinsic shell properties influence preservation. Thus, taphofacies analyses should combine data on taphonomic features, specific life habit and shell properties to determine overall preservation patterns. In that way, taphofacies analyses may form a powerful tool for palaeoenvironmental analyses of marine deposits.     

Modern benthic foraminifera on Banco Chinchorro,Quintana Roo,Mexico

Benthic foraminiferal assemblages may be used as environmental indicators on Banco Chinchorro, an isolated carbonate platform off the eastern coast of the Yucatan Peninsula. Seventy-seven species from 44 genera were identified in 14 surface sediment samples, which were collected along an E–W windward-to-leeward transect across the platform. A total of 15,493 foraminiferal tests (max. 1,200 tests per sample) were investigated and served as the basis of this study. As many taxa range throughout several platform zones, assemblages are better environmental indicators as compared to individual species. Four foraminiferal assemblages were identified using statistical methods including (1) a Homotrema assemblage, which occurs at the windward platform margin, (2) an Archaias-Homotrema assemblage which is found on the leeward margin and on platform interior coral patch reefs, (3) a Quinqueloculininae-Archaias-Rosalina assemblage of the western platform, and (4) an Archaias-Quinqueloculininae assemblage characteristic of the eastern platform interior. Environmental factors which influence variation in foraminiferal distribution and diversity on Banco Chinchorro include exposure to waves and currents and substrate (plant and algal growth). Sediment transport does not play a major role in Banco Chinchorro based on the observation that there are only limited amounts of taxa found outside their typical habitats, and, that mean grain-size and sorting of foraminiferal tests do not exhibit clear patterns. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Live and dead benthic foraminiferal assemblages from coastal environments of the Aegean Sea (Greece): Distribution and diversity

Benthic foraminiferal composition assemblages and their temporal changes, ecological indices and foraminiferal densities are used to compare three coastal environments with different physicogeographical features in the Aegean Sea (coastal environment of Avdira–Vistonikos Gulf and Kitros–Thermaikos Gulf and open lagoonal environment of Vravron–South Evoikos Gulf). Three main foraminiferal assemblages have been recognized: a) “Assemblage A”; high degree of similarity between living and dead foraminiferal species, dominated by Ammonia beccarii, Elphidium spp. and relatively abundant and diverse miliolids, b) “Assemblage B1”; intermediate degree of similarity between live and dead assemblages, characterized by highly-abundant and well-diversified foraminiferal assemblages including the algal symbiont bearing Peneroplis pertusus together with Ammonia tepida and several small epiphytic rotaliids and miliolids, and c) “Assemblage B2”; absence of living individuals, strongly dominated by the opportunistic species A. tepida. Our results suggest a good comparison between living and dead assemblages from different coastal environments in the Aegean Sea, however the prevailing environmental conditions (vegetation cover, hydrodynamics, fresh water influx) have a strong impact on the taphonomic processes.     

Ichnofacies: a mixture of taphofacies and biofacies

The environmental significance of individual ichnofacies has become a point of debate, and the Scoyenia ichnofacies is not clearly defined. Removal of the marine/non-marine boundary from ichnofacies definitions, as well as the bathymetry control, would render the Scoyenia ichnofacies unnecessary and would consolidate the definitions of the remaining ichnofacies. The constitution of the remainder is far from uniform. In some, the characteristic features are influenced by taphonomic bias more than by ecological factors, and thus these ichnofacies are in effect taphofacies (Skolithos, Cruziana and, especially, Zoophycos and Nereites). Others, on the other hand, have a primarily ecological definition and function as biofacies (Glossifungites, Trypanites, Teredolites and Psilonichnus), taphonomic bias playing a relatively minor role. Opportunistic occurrences of Skolithos in storm deposits do not fall within the Skolithos ichnofacies sensu stricto , and may be regarded as a separate Arenicolites ichnofacies, having an ecological character. □ Trace fossils, ichnofacies, salinity.     

A palaeoecological model for the interpretation of wild plant species

The interpretation of subfossil records of wild plant species with respect to both environmental conditions and past vegetation is complicated by the following: (1) production and dispersal of plant remains including diaspores, (2) the formation of the soil flora, (3) taphonomic processes and differential preservation that act on subfossil assemblages and (4) methods applied to produce subfossil records. Whereas the similarity between recent plant communities and seed banks is often weak, the relationship between past vegetation and subfossil assemblages is still more complicated. It is therefore unlikely that macrofossil assemblages derived from soil samples can be considered as pure samples representing particular palaeobiocoenoses. The assumption that plant communities, in the past, may have been in some way aberrant with respect to composition and that the ecological ranges of species varied during the Quaternary has to be rejected, if not based on well considered assumptions or evidence from pure samples. Only if a sufficient number of suitable studies is available, which enable evaluation between all kinds of plant communities and their respective seed floras, can progress be made with regard to the reconstruction of past vegetation and environmental conditions. As long as these data are not available, the ecological interpretation of particular subfossil assemblages isolated from soil samples has to be carefully evaluated within their particular context.     

Foraminifera,Environmental Bioindicators in the Highly Impacted Environments of the Mekong Delta

Foraminiferal assemblages has been studied in three regions of the Mekong Delta subjected to different environmental characteristics: the Tra Vinh province, in the middle part of the Delta, under high freshwater influence; the Ca Mau peninsula, to the south, under maximum marine influence; and the Can Gio region, in the northern part of the Delta, with intermediate conditions. Intensive Rice–Shrimp Farming System is developing in these regions. Foraminiferal assemblages show the prevalence of the Mekong’s water pressure. Marine influence is strong only at a few stations along the shoreline, mainly in the Can Gio region. The rarity of allochthonous coastal species, in all the samples except one, suggests that up-river transport and settlement of foraminifera and sediment is significant only in the major channels. On a local level, foraminiferal assemblages provide a variety of information on environmental conditions. In a tidal channel of the Tra Vinh province, they suggest that lower reaches are more impacted by pollution than upper reaches; in the Ca Mau peninsula, under marine influence, low density and dominance of agglutinated species indicate the impact of deforestation and extension of acid sulfate soils; in shrimp pools, foraminifera may be used as indicators of the degradation of local conditions, in relation with accumulation of organic matter. The distribution of foraminiferal assemblages described in this study should be used as part of a reference point for further observations with a view to assessing environmental impacts of increasing human activities and/or climatic change.     

Modern foraminiferal record of alternating open and restricted environmental conditions in the Santo André lagoon,SW Portugal

Benthic foraminifera from eleven stations sampled seasonally were analyzed in order to examine the biological response to rapid and intense environmental changes taking place in the Santo André coastal lagoon. Foraminiferal assemblages show a very low species diversity and a high dominance of three euryhaline species throughout the year. Under closed-inlet conditions, foraminiferal assemblages exhibit low abundance of foraminiferal tests, whereas under open-inlet conditions absolute abundance greatly increases due to sea water entrance. Present-day dramatic environmental changes are thought to be responsible for deformed foraminiferal tests that are commonly found. Comparison of modern assemblages with those obtained from the Holocene sedimentary record indicates persistent restrictive environmental conditions after sandy barrier formation, circa 5000 years ago.     

The effect of bioturbation by polychaetes (Opheliidae) on benthic foraminiferal assemblages and test preservation

Biological activity such as burrowing can alter benthic foraminiferal shell preservation and may also modify benthic foraminiferal assemblages by vertical mixing, inducing sediment homogenization. Here, we analyse benthic foraminiferal assemblages and taphonomy of upper Miocene marine deposits from Conil de la Frontera (Cádiz, south‐western Spain). The deposits consist of marls displaying a pervasive alternation of intensively bioturbated beds dominated by Macaronichnus segregatis traces (ichnofabric index 4–5) and non‐bioturbated beds. Benthic foraminiferal assemblages are dominated by Cibicidoides mundulus and Cibicides refulgens, indicating that the marls were deposited on an oligotrophic, well‐oxygenated upper slope. The impact of burrowing on the preservation of benthic foraminiferal tests was tested using Q‐mode cluster analysis, which found two well‐differentiated groups of samples, one including the non‐bioturbated beds and the other encompassing the bioturbated ones. Fragmentation and recrystallization account for the differentiation of these groups, both being higher in the bioturbated sediments. Aggressive chemical digestion by the Macaronichnus trace‐makers, assumed to be a polychaete worm of the family Opheliidae, etched the microfossil shells, making them more vulnerable to fragmentation. Intense bioturbation favoured the circulation of pore fluids, encouraging recrystallization. Pervasive burrowing resulted in significant vertical reworking of microfossils. As a consequence, benthic foraminiferal assemblages in the bioturbated beds were homogenized in the mixed layer; that is, the uppermost layer of the substrate totally burrowed. The alternation of bioturbated and non‐bioturbated beds reflects episodic transfer of food particles down slope from shallower parts of the shelf as well as from the continent due to storms under otherwise homogeneous oligotrophic marine conditions.     

Benthic foraminiferal bioevents in lower to upper Toarcian strata of Southern Beaujolais (SE France)

This paper documents changes in benthic foraminiferal assemblages compared with high resolution ammonite biozonation along the lower Toarcian to upper Toarcian marine succession of Southern Beaujolais in southeastern France. Eight ammonite and three benthic foraminiferal zones including five subzones are distinguished based on the occurrence of twelve foraminiferal events. Each benthic foraminiferal subzone is characterized by its taxonomic and morphogroup composition, which represents the paleoecological response of these taxa and morphotypes of benthic foraminifera in the Early Jurassic and early Middle Jurassic. Major changes in abundance and diversity occur at the end of the Toarcian Oceanic Anoxic Event (T-OAE) and near the Early-Middle Jurassic transition. The low-abundance foraminiferal assemblages recorded in the Serpentinus ammonite Zone are interpreted as reflecting adverse environmental conditions after the T-OAE. The later recovery and development of the foraminiferal assemblages is documented in the Bifrons up to the Aalensis zones and is attributed to improved bottom water oxygenation. Common occurrences of agglutinated foraminifera represented mostly by Trochammina pulchra Ziegler in the Dispensum Zone point to an influx of cooler water masses during the late Toarcian. The morphogroup analysis carried out on the foraminifera and their paleoecological interpretations shed light on the changes in the stratigraphic record at the end of the T-OAE up to the Toarcian/Aalenian boundary.     

Ammonoid taphonomy, palaeoenvironments and sequence stratigraphy at the Bajocian/Bathonian boundary on the Bas Auran area (Subalpine Basin, south-eastern France)

Bas Auran, in south‐eastern France, is the candidate area for Global Boundary Stratotype Section and Point (GSSP) of the base of the Bathonian Stage (Middle Jurassic). In the Bas Auran area, upper Bajocian and lower Bathonian deposits are made up of limestone beds alternating with marls that correspond to the ‘Marno‐calcaires àCancellophycus’, below the ‘Terres Noires’ Formation. Taphonomic analyses of the successive ammonoid fossil assemblages provide new and complementary data for biostratigraphic completeness, palaeoenvironmental setting and sequence stratigraphy. Lithologic differentiation between limestone and marl intervals resulted from alternating episodes of carbonate input and starvation. Both lithologic phases may contain evidence for sedimentary and taphonomic reworking, associated with scours, that reflects low rates of sedimentation and stratigraphic condensation. Three successive types of elementary cycles resulted from increasing rates of stratigraphic condensation, sedimentary condensation and substrate stabilization during early Bathonian. The occurrence of reelaborated ammonoids (i.e. exhumed and displaced before their final burial) implies that tractive current flows or winnowing affected the burial of concretionary internal moulds. In the lower Bathonian strata, the dominance of homogeneous concretionary internal moulds of phragmocones, completely filled with sediment, is indicative of low rates of sedimentation and sediment accumulation, respectively associated with low degrees of stratigraphic and sedimentary condensation. However, at the Bajocian/Bathonian transition, hemipelagic, bed‐scale limestone–marl alternations show a maximum value of biostratigraphic completeness and there is no evidence for taphonomic condensation in the ammonoid fossil assemblages. Taphonomic analyses of the successive ammonoid fossil assemblages and taphofacies confirm the development of a deepening phase associated with sedimentary starvation, which characterizes the last episode within the deepening half‐cycle of third and second order cycles, in the Bas Auran area of French Subalpine Basin during early Bathonian.     

Benthic foraminiferal assemblages between two major extinction events: the Paleocene El Kef section,Tunisia

The development of benthic foraminiferal assemblages from the Paleocene outcrops of the El Haria Formation near El Kef, Tunisia is discussed qualitatively and quantitatively. The aim of the study is to reconstruct the paleoenvironmental evolution between the K/Pg boundary interval and the late Paleocene event, and to compare this evolution with results from other sites along the southern Tethyan margin. Eighty-four samples, covering virtually the entire Paleocene, provide a dataset that allows detailed qualitative and multivariate analysis. The benthic foraminiferal faunas indicate a complex pattern of environmental changes during the Paleocene, marked by the succession of different benthic associations. Following the K/Pg boundary event, community restoration was characterized by the gradual build-up of faunal diversity. Decreasing dominance and the entry of taxa common to normal marine, outer neritic to upper bathyal environments indicate the completion of the ecosystem restoration in Zone Plb. A highly diverse benthic foraminiferal assemblage persisted throughout the remainder of the early Paleocene into the earliest late Paleocene. At the P3a-P3b zonal transition relative sea-level lowering is evidenced by the sudden disappearance or decreasing abundance of deeper-water taxa (e.g. Anomalinoides affinis, A. susanaensis, Gavelinella beccariiformis). Neritic deposition continued into Zone P4, when trophic levels at the seafloor increased as indicated by the entry and increasing dominance of species such as Anomalinoides cf. aegyptiacus, Bulimina midwayensis, and B. strobila, which we consider to be sensitive to eutrophication. The combined effect of shallowing and the subsequent eutrophication led to the establishment of assemblages similar to late Paleocene benthic foraminiferal assemblages from Egyptian sections, some of which record the latest Paleocene extinction event. These assemblages were interpreted to be indicative of a middle neritic, highly eutrophic environment. Enhanced vertical fluxes of organic matter along the southern Tethyan margin may have resulted from intensified upwelling. This eventually led to oxygen deficiency at the seafloor. It appears that oxygen-deficient, high-productivity shelves were a common feature of the southern Tethyan margin during the latest Paleocene.     

Effectiveness of benthic foraminiferal and coral assemblages as water quality indicators on inshore reefs of the Great Barrier Reef, Australia

Although the debate about coral reef decline focuses on global disturbances (e.g., increasing temperatures and acidification), local stressors (nutrient runoff and overfishing) continue to affect reef health and resilience. The effectiveness of foraminiferal and hard-coral assemblages as indicators of changes in water quality was assessed on 27 inshore reefs along the Great Barrier Reef. Environmental variables (i.e., several water quality and sediment parameters) and the composition of both benthic foraminiferal and hard-coral assemblages differed significantly between four regions (Whitsunday, Burdekin, Fitzroy, and the Wet Tropics). Grain size and organic carbon and nitrogen content of sediments, and a composite water column parameter (based on turbidity and concentrations of particulate matter) explained a significant amount of variation in the data (tested by redundancy analyses) in both assemblages. Heterotrophic species of foraminifera were dominant in sediments with high organic content and in localities with low light availability, whereas symbiont-bearing mixotrophic species were dominant elsewhere. A similar suite of parameters explained 89% of the variation in the FORAM index (a Caribbean coral reef health indicator) and 61% in foraminiferal species richness. Coral richness was not related to environmental setting. Coral assemblages varied in response to environmental variables, but were strongly shaped by acute disturbances (e.g., cyclones, Acanthaster planci outbreaks, and bleaching), thus different coral assemblages may be found at sites with the same environmental conditions. Disturbances also affect foraminiferal assemblages, but they appeared to recover more rapidly than corals. Foraminiferal assemblages are effective bioindicators of turbidity/light regimes and organic enrichment of sediments on coral reefs.     

Postmortem transport and resedimentation of foraminiferal tests: relations to cyclical changes of foraminiferal assemblages

Cyclical changes in microfossil (mainly foraminiferal) assemblages were analysed for sixteen boreholes from three stratigraphical levels in the Lower and Middle Miocene of the Central Paratethys. The following characteristics of assemblages were quantified and used for interpretation of cyclical changes in assemblages: (1) abundance of foraminifers, calcareous nannoplankton, sponge spicules and diatoms; (2) similarity, diversity and epifauna/infauna ratio of benthonic foraminiferal assemblages; (3) planktic/benthic ratio of foraminiferal assemblages. The palaeoecology (mainly palaeobathymetry) fluctuations were interpreted from the species composition of assemblages. Values of the mentioned quantitative characteristics as well as palaeoecological interpretations may be influenced by postmortem transport and resedimentation of foraminiferal tests. Therefore, prior to the interpretation of cyclical changes of quantitative characteristics, the studied sections were classified on the basis of the intensity of taphonomical changes in foraminiferal assemblages. Three different categories of sections were obtained. For every category, those quantitative characteristics of foraminiferal assemblages were chosen which reflect cyclical changes most efficiently: (1) cyclical changes of abundance of foraminifers, calcareous nannoplankton and sponge spicules, as well as the fluctuations in palaeobathymetry for sections dominated by indigenous foraminifers; (2) percentage of indigenous, suspension-transported, bedload-transported and reworked foraminiferal tests and changes in the abundance of indigenous tests for sections dominated by transported tests; (3) different modes of test preservation used for the identification of a cycle boundary for sections with only transported or reworked tests. The distinguished cycles were interpreted predominantly as manifestations of relative sea-level changes. If comparable data exist from other Paratethys regions, the determined cycles can be correlated with the other basins.     

Experimental dissolution of a fossil foraminiferal assemblage (Paleocene–Eocene Thermal Maximum,Dababiya, Egypt): Implications for paleoenvironmental reconstructions

Dissolution experiments were carried out on a foraminiferal assemblage from the Paleocene–Eocene Thermal Maximum (PETM) at Dababiya, Egypt, in order to: 1) reveal the effects of differential dissolution on the composition of the foraminiferal assemblage and 2) develop objective criteria for the evaluation of dissolution in foraminiferal assemblages used in early Paleogene paleoenvironmental reconstructions, particularly with respect to neritic Midway-type assemblages from the Paleocene/Eocene transition. Our results confirm two general observations on modern foraminifera: 1) planktic foraminifera are much more vulnerable to dissolution than benthic foraminifera, leading to depressed P/B ratios and 2) dissolution susceptibility differs between size fractions, with the smaller specimens dissolving more rapidly than the bigger ones, leading to a larger average size of the remaining assemblage. Within a size fraction, wall structure and thickness are considered to be the main factors controlling differential dissolution susceptibility. We propose a ranking scheme for taxa with respect to dissolution resistance. Among the benthic taxa, Lenticulina is most resistant, followed by the agglutinated Gaudryina cf. ellisorae and Alabamina midwayensis. Biserial and triserial hyaline taxa and the porcelaneous Spiroloculina sp. are most susceptible to dissolution, whereas rotaliines, such as Cibicidoides and Anomalinoides have an intermediate susceptibility. This implies that mild dissolution of a Midway-type benthic assemblage leads to a relative enrichment in Lenticulina, Gaudryina and rotaliines. Amongst planktic foraminifera, the muricate taxa Acarinina and Morozovella are most resistant, followed by the cancellate Subbotina. The smooth and generally small Globanomalina and Zeauvigerina are least resistant to dissolution. Our data enable to objectively evaluate various degrees of dissolution in benthic and planktic foraminiferal assemblages retrieved from the lower Paleogene Tethyan outcrops. In this way taphonomic artifacts can be readily distinguished from paleoenvironmental signals affecting the primary composition of the assemblages. More generally, we propose that the combined use of foraminiferal numbers, P/B ratio and relative abundances of non-calcareous agglutinated taxa and Lenticulina may provide a powerful proxy for assessing dissolution in hemipelagic assemblages from Cenozoic and upper Cretaceous continental margins. In order to achieve more robust pre-Quaternary paleoenvironmental reconstructions based on quantitative foraminiferal data, application of dissolution proxies, like proposed here, or in slightly modified form, should become a more widely used micropaleontologic procedure. Particularly continental margin studies dealing with major biotic events (e.g. PETM) or employing P/B ratios for sea-level reconstructions should benefit from such an approach.     

Autochthonous death assemblages from chemoautotrophic communities at petroleum seeps: Palaeoproduction,energy flow,and implications for the fossil record

Death assemblages produced by chemoautotrophic communities at cold seeps represent a type of autochthonous accumulation that is difficult to differentiate from other heterotrophic autochthonous communities using taphonomic characteristics. We test the hypothesis that cold‐seep assemblages can be discriminated by unique biological or community attributes rather than taphonomic attributes. To test this hypothesis, we compared several cold seeps on the Louisiana upper continental slope to heterotrophic sites on the Louisiana slope and to a putative seep site in the middle‐late Campanian Pierre Shale near Pueblo, Colorado. Seep assemblages are characterized by a unique tier and guild structure, size‐frequency composition, and animal density that together identify the palaeoenergetics structure of these communities and distinguish them from the other assemblages of the shelf and slope. All seep assemblages were dominated by primary consumers, whereas the heterotrophic assemblage was dominated by carnivores. Carnivore dominance seems to be typical of shelf (or euhaline) death assemblages. Seep assemblages, in contrast, retain the theoretically‐expected rarity of predaceous forms in fossil assemblages. Epifauna and semi‐infauna dominate the tier structure of the heterotrophic assemblage as is typical for continental shelf and slope assemblages. The infaunal tier was unusually well represented in most petroleum seep assemblages. Local enrichment of food resources and the dominance of shelled primary consumers explain the guild and tier structure of seep assemblages. Hindcasting of energy demand (palaeoingestion) and an estimate of sedimentation rate confirms that energy demand by the community exceeds the supply from planktonic rain in seep communities. Thus, seep assemblages can be recognized using biological attributes where taphonomic analysis is ambiguous.