Pliocene coastal palaeomorphology and ostracod faunas of the Bass Strait hinterland,southeast Australia

Early to late Pliocene sedimentary strata present across the northern Bass Strait hinterland, southeastern Australia yield extensive fossil proxy data relevant to the interpretation of high sea level coastal palaeomorphology. Within the Pliocene Whalers Bluff Formation exposed in coastal cliffs near the township of Portland, Victoria, marine microfossil faunas delineate two broad cycles of deposition. Both these sedimentary cycles are bound below by unconformity surfaces. Within the lower sedimentary cycle, a basal stress-tolerant (low diversity) marginal marine microfossil fauna devoid of ostracods and suggestive of bottom-water hypoxia, is succeeded by a diverse shallow marine ostracod fauna dominated by stenohaline species indicative of a sheltered (but open) oceanic embayment. This lower sedimentary cycle has an early Pliocene (Zanclean) age. Equivalent shallow marine (e.g. coastal embayment) deposits occur broadly across the coastal hinterland of southeastern Australia—reflecting the generally higher global sea levels of this time. The upper cycle in the cliff exposures at Portland is late Pliocene (Piacenzian) in age. Equivalent deposits across the Bass Strait hinterland are restricted to former incised river valley settings. Euryhaline estuarine/coastal lagoon Ostracoda are present throughout the upper cycle in the Portland cliffs. These are associated with a low diversity microfauna at the base of the upper cycle and a high diversity microfauna towards the top of the cycle. Early Pliocene coastal marine deposits can be distinguished from late Pliocene coastal marine deposits across the northern Bass Strait hinterland on the basis of the presence or absence of certain open marine (‘stenohaline’) ostracod species.     

Foraminifera diversity changes and paleoenvironmental analysis: the Lower Cretaceous shallow-water carbonates of San Lorenzello, Campanian Apennines, southern Italy

A high-resolution stratigraphic study, carried out on the carbonate platform strata of the San Lorenzello section (Matese Mountains, southern Italy), Valanginian–Hauterivian in age, has allowed to: recognise lithofacies and their associations; assign the lithofacies associations to specific environments and individuate early meteoric diagenetic modifications, recurring at specific horizons. In this frame the vertical variation of benthic foram diversity has been analysed. On the whole, foraminiferal genera diversity decreases from open to restricted marine environments. Moreover, a climatic control on carbonate sedimentation is suggested by a Milankovitch cyclicity organised in elementary cycles, bundles and superbundles as well as by diagenetic characteristics testifying that humid and arid conditions alternated during the Early Cretaceous times. The orbital cyclicity is also documented by foraminiferal diversity changes, even if some discrepancies between the lithofacies and the diversity locally occur. Therefore, the above diversity changes do not appear to provide sufficient information for the sequence-stratigraphic interpretation of shallow-water carbonates.     

Magnetostratigraphy susceptibility of the Upper Ordovician Kope Formation,northern Kentucky

As a test of the utility of magnetic susceptibility (MS) measurements for correlation among lithified marine sequences, samples were collected from well-studied shale and limestone outcrops of the Upper Ordovician Kope Formation, northern Kentucky. The bulk (initial) low-field MS of these samples is compared among two litho- and biostratigraphically correlated sections containing the same beds. The results of this comparison demonstrated an excellent correlation among equivalent beds. Thermomagnetic susceptibility analysis supports previous XRD work indicating that illite is primary and is the main paramagnetic mineral responsible for the MS variations observed in this study. A composite of MS results for a sequence of 31 named shale–limestone couplets is used to build a composite section of Kope sequences for the area. These MS variations permit division of the various sections into a set of 28 MS cycles within the Kope Formation, covering ∼ 50 m of section. These cycles suggest a systematic control by climate on the influx of detrital material that formed these sedimentary sequences, and support other work indicating Kope climate cyclicity.     

Integrated analyses of litho- and biofacies in a Pliocene cyclothemic, alluvial to shallow marine succession (Tuscany, Italy)

Lithofacies analysis of the upper part of the Pliocene succession of the Valdelsa basin (central Italy) unravelled a number of depositional environments, ranging from alluvial plain to coastal, to marine. Strata are arranged in a hierarchy of elementary and composite unconformity-bounded units. A palaeoecological study of macro- (molluscs) and microfossils (pollen, dinocysts, foraminifera) allowed to finely reconstruct sub-environments within fine-grained terrestrial, coastal and marine deposits and thence to track the spatial and temporal change of physical conditions. The stacking pattern of sedimentary units highlights the lateral switching of onshore-offshore gradients and documents relative sea-level changes. These units are interpreted in a sequence stratigraphic framework. Elementary depositional sequences are arranged to form six composite depositional sequences, in turn encased within two major synthems. This hierarchy of unconformity-bounded sedimentary units suggests that sea-level variation has occurred at different time-frequencies. Glacio-eustasy and active tectonism are discussed as the main forcing factors regulating the different scales of sedimentary cyclicity.     

The sea as deathtrap: comment on a paper by miller and wiens

Miller & Wiens (2017) claim that low marine as compared with terrestrial diversity results from more frequent extinctions and insufficient time for diversification in marine clades. Their data on marine amniotes are unrepresentative of marine diversity, their analysis of clade dynamics is flawed, and they ignore previously proposed explanations for the diversity difference.     

A global perspective for biodiversity history with ancient environmental DNA

The past centuries have seen tremendous turnovers in species distributions and biodiversity due to anthropogenic impacts on a global scale. The processes are ongoing and mostly not well documented. Long‐term records of biotic change can be recovered from sedimentary deposits, but traditional analyses were restricted to organisms that leave behind visible traces and molecular genetic tools were mostly employed on samples that promised good DNA preservation. In this issue of Molecular Ecology, Shaw, Weyrich, Hallegraeff and Cooper (2019) and Gomez Cabrera et al. (2019) present two studies on marine sedimentary records from warm environments, in which they successfully analyze ancient environmental DNA (aeDNA) on a decadal and centennial scale. Notably, the studies were conducted on novel samples with nonoptimal preservation conditions for ancient DNA ‐ historical collections of ship ballast tank sediments from Australia and two coral reef cores spanning up to 750 years (Figure 1) ‐ but yielded a high diversity of taxa. This highlights that aeDNA is a promising tool to globally study biodiversity history on scales of decades to centuries ‐ the timeframe most relevant to human society in the context of both current climate change and direct anthropogenic modifications of the environment.     

Mesozoic marine tetrapod diversity: mass extinctions and temporal heterogeneity in geological megabiases affecting vertebrates

The fossil record is our only direct means for evaluating shifts in biodiversity through Earth''s history. However, analyses of fossil marine invertebrates have demonstrated that geological megabiases profoundly influence fossil preservation and discovery, obscuring true diversity signals. Comparable studies of vertebrate palaeodiversity patterns remain in their infancy. A new species-level dataset of Mesozoic marine tetrapod occurrences was compared with a proxy for temporal variation in the volume and facies diversity of fossiliferous rock (number of marine fossiliferous formations: FMF). A strong correlation between taxic diversity and FMF is present during the Cretaceous. Weak or no correlation of Jurassic data suggests a qualitatively different sampling regime resulting from five apparent peaks in Triassic–Jurassic diversity. These correspond to a small number of European formations that have been the subject of intensive collecting, and represent ‘Lagerstätten effects’. Consideration of sampling biases allows re-evaluation of proposed mass extinction events. Marine tetrapod diversity declined during the Carnian or Norian. However, the proposed end-Triassic extinction event cannot be recognized with confidence. Some evidence supports an extinction event near the Jurassic/Cretaceous boundary, but the proposed end-Cenomanian extinction is probably an artefact of poor sampling. Marine tetrapod diversity underwent a long-term decline prior to the Cretaceous–Palaeogene extinction.     

THE SHAPE OF THE PHANEROZOIC MARINE PALAEODIVERSITY CURVE: HOW MUCH CAN BE PREDICTED FROM THE SEDIMENTARY ROCK RECORD OF WESTERN EUROPE?

Abstract:  Palaeodiversity curves are constructed from counts of fossils collected at outcrop and thus potentially biased by variation in the rock record, specifically by the amount of sedimentary rock representative of different time intervals that has been preserved at outcrop. To investigate how much of a problem this poses we have compiled a high-resolution record of marine rock outcrop area in Western Europe for the Phanerozoic and use this to generate a model that predicts the sampled diversity curve. We find that we can predict with high accuracy the variance of the marine genus diversity curve (itself dominated by European taxa) from rock outcrop data and a three-step model of diversity that tracks supercontinent fragmentation, coalescence and fragmentation. The size and position of two of the five major mass extinction spikes are largely predicted by rock outcrop data. We conclude that the long-term trends in taxonomic diversity and the end-Cretaceous extinction are not the result of rock area bias, but cannot rule out that rock outcrop area bias explains many of the short-term rises and falls in sampled diversity that palaeontologists have previously sought to explain biologically.     

Demographic history and the low genetic diversity in Dipteryx alata (Fabaceae) from Brazilian Neotropical savannas

Genetic effects of habitat fragmentation may be undetectable because they are generally a recent event in evolutionary time or because of confounding effects such as historical bottlenecks and historical changes in species'' distribution. To assess the effects of demographic history on the genetic diversity and population structure in the Neotropical tree Dipteryx alata (Fabaceae), we used coalescence analyses coupled with ecological niche modeling to hindcast its distribution over the last 21 000 years. Twenty-five populations (644 individuals) were sampled and all individuals were genotyped using eight microsatellite loci. All populations presented low allelic richness and genetic diversity. The estimated effective population size was small in all populations and gene flow was negligible among most. We also found a significant signal of demographic reduction in most cases. Genetic differentiation among populations was significantly correlated with geographical distance. Allelic richness showed a spatial cline pattern in relation to the species'' paleodistribution 21 kyr BP (thousand years before present), as expected under a range expansion model. Our results show strong evidences that genetic diversity in D. alata is the outcome of the historical changes in species distribution during the late Pleistocene. Because of this historically low effective population size and the low genetic diversity, recent fragmentation of the Cerrado biome may increase population differentiation, causing population decline and compromising long-term persistence.     

A Late Triassic lake system in East Greenland: facies, depositional cycles and palaeoclimate

The Upper Triassic Fleming Fjord Formation of the Jameson Land Basin in East Greenland contains a well-exposed succession, 200–300 m thick, of lake deposits. The Malmros Klint Member, 100–130 m thick, is composed of cyclically bedded intraformational conglomerates, red siltstones and fine-grained sandstones and disrupted dolomitic sediments (paleosols). The cyclicity is composite with cycles having mean thicknesses of (25), 5.9 and 1.6 m. The overlying Carlsberg Fjord beds of the Ørsted Dal Member, 80–115 m thick, are composed of structureless red mudstones rhythmically broken by thin greyish siltstones. This unit also has a composite cyclicity with cycles having mean thicknesses of 5.0 and 1.0 m. The uppermost Tait Bjerg Beds of the Ørsted Dal Member, 50–65 m thick, can be divided into two units. A lower unit is composed of cyclically bedded intraformational conglomerates or thin sandstones, red mudstones, greenish mudstones and yellowish marlstones. An upper unit is composed of relatively simple cycles of grey mudstones and yellowish marlstones. Recognized cycles have mean thicknesses of 5.6 and 1.6 m. The lake deposits contain evidence of seasonal, orbital and long-term climatic change. Seasonal change is documented by numerous desiccation surfaces especially in the Malmros Klint Member and Carlsberg Fjord beds, orbital change is suggested by the composite cyclicity, and long-term climatic change is indicated by the systematic upwards change in sedimentary characteristics of the lake deposits. The sedimentary features of the Malmros Klint Member suggest lacustrine deposition in a dry climate that fluctuated between desert and steppe conditions, the Carlsberg Fjord beds probably record lacustrine lake deposition in a rather constant dry (steppe) climate, while the Tait Bjerg Beds record lake sedimentation in a climate that fluctuated between dry (steppe) and warm moist temperate. In the Tait Bjerg Beds the upward change in cycle characteristics indicates a shift towards more humid conditions. Climatic deductions from sedimentary facies are in good agreement with climate maps of Laurasia, as simulated by numerical climate models. Palaeomagnetic data indicate a northward drift of East Greenland of about 10° from ca. 25°N to ca. 35°N in the Middle to Late Triassic. The Fleming Fjord Formation which represents ca. 5 m.y. of the Late Triassic interval was deposited during latitudinal drift of 1–2°. It is possible that the observed long-term upward shift in climatic indicators within the formation can be ascribed to plate drift, but southward shift of climatic belts could also have been of importance.     

Equilibrium Responses Reflected in a Large Conichnus (Upper Cretaceous Eutaw Formation,Alabama, USA)

A vertically extensive (1.4 m) specimen of Conichnus in the Cretaceous Eutaw Formation (Alabama) exhibits marginal and internal fabrics that reflect equilibrium behavioral responses to changes in sedimentation rate and erosion events associated with tidal sandwave dynamics. Distinct fabrics can be linked to: (1) initial substrate penetration by the tracemaker, which followed stoss-side erosion of sandwave foresets; (2) gradual retrusive movement in response to slow sediment aggradation (<1 cm/day) in sandwave bottomsets; (3) intermediate to high rates of retrusive migration in response to moderate to rapid sediment aggradation (～1 cm/day) in sandwave toes; (4) tracemaker escape associated with extremely rapid (>5 cm/day) foreset deposition; and (5) protrusive equilibrium movement in response to stoss-side erosion of foresets. Morphological observations in the context of tidal cyclicity suggest that the trace, presumably produced by an anemone, was a semipermanent domicile occupied for at least two to three months. Relationships established between burrow fabric and sedimentary conditions in the Eutaw specimen may aid in the interpretation of conichnian burrows in other high-energy, shallow marine sequences.     

The oldest marine vertebrate fossil from the volcanic island of Iceland: a partial right whale skull from the high latitude Pliocene Tjörnes Formation

Extant baleen whales (Cetacea, Mysticeti) are a disparate and species‐rich group, but little is known about their fossil record in the northernmost Atlantic Ocean, a region that supports considerable extant cetacean diversity. Iceland's geographical setting, dividing North Atlantic and Arctic waters, renders it ideally situated to shed light on cetacean evolution in this region. However, as a volcanic island, Iceland exhibits very little marine sedimentary exposure, and fossil whales from Iceland older than the late Pleistocene are virtually unknown. Here, we present the first fossil whale found in situ from the Pliocene Tjörnes Formation (c. 4.5 Ma), Iceland's only substantial marine sedimentary outcrop. The specimen is diagnosed as a partial skull from a large right whale (Mysticeti, Balaenidae). This discovery highlights the Tjörnes Formation as a potentially productive fossil vertebrate locality. Additionally, this find indicates that right whales (Eubalaena) and bowhead whales (Balaena) were sympatric, with broadly overlapping latitudinal ranges in the Pliocene, in contrast to the modern latitudinal separation of their living counterparts.     

Environmental DNA (eDNA) metabarcoding reveals strong discrimination among diverse marine habitats connected by water movement

While in recent years environmental DNA (eDNA) metabarcoding surveys have shown great promise as an alternative monitoring method, the integration into existing marine monitoring programs may be confounded by the dispersal of the eDNA signal. Currents and tidal influences could transport eDNA over great distances, inducing false‐positive species detection, leading to inaccurate biodiversity assessments and, ultimately, mismanagement of marine environments. In this study, we determined the ability of eDNA metabarcoding surveys to distinguish localized signals obtained from four marine habitats within a small spatial scale (<5 km) subject to significant tidal and along‐shore water flow. Our eDNA metabarcoding survey detected 86 genera, within 77 families and across 11 phyla using three established metabarcoding assays targeting fish (16S rRNA gene), crustacean (16S rRNA gene) and eukaryotic (cytochrome oxidase subunit 1) diversity. Ordination and cluster analyses for both taxonomic and OTU data sets show distinct eDNA signals between the sampled habitats, suggesting dispersal of eDNA among habitats was limited. Individual taxa with strong habitat preferences displayed localized eDNA signals in accordance with their respective habitat, whereas taxa known to be less habitat‐specific generated more ubiquitous signals. Our data add to evidence that eDNA metabarcoding surveys in marine environments detect a broad range of taxa that are spatially discrete. Our work also highlights that refinement of assay choice is essential to realize the full potential of eDNA metabarcoding surveys in marine biodiversity monitoring programs.     

Reduced Global Genetic Differentiation of Exploited Marine Fish Species

Knowledge on genetic structure is key to understand species connectivity patterns and to define the spatiotemporal scales over which conservation management plans should be designed and implemented. The distribution of genetic diversity (within and among populations) greatly influences species ability to cope and adapt to environmental changes, ultimately determining their long-term resilience to ecological disturbances. Yet, the drivers shaping connectivity and structure in marine fish populations remain elusive, as are the effects of fishing activities on genetic subdivision. To investigate these questions, we conducted a meta-analysis and compiled genetic differentiation data (F_ST/Φ_ST estimates) for more than 170 fish species from over 200 published studies globally distributed. We modeled the effects of multiple life-history traits, distance metrics, and methodological factors on observed population differentiation indices and specifically tested whether any signal arising from different exposure to fishing exploitation could be detected. Although the myriad of variables shaping genetic structure makes it challenging to isolate the influence of single drivers, results showed a significant correlation between commercial importance and genetic structure, with widespread lower population differentiation in commercially exploited species. Moreover, models indicate that variables commonly used as proxy for connectivity, such as larval pelagic duration, might be insufficient, and suggest that deep-sea species may disperse further. Overall, these results contribute to the growing body of knowledge on marine genetic connectivity and suggest a potential effect of commercial fisheries on the homogenization of genetic diversity, highlighting the need for additional research focused on dispersal ecology to ensure long-term sustainability of exploited marine species.     

Summer temperature dependency of larch budmoth outbreaks revealed by Alpine tree-ring isotope chronologies

Larch budmoth (LBM, Zeiraphera diniana Gn.) outbreaks cause discernable physical alteration of cell growth in tree rings of host subalpine larch (Larix decidua Mill.) in the European Alps. However, it is not clear if these outbreaks also impact isotopic signatures in tree-ring cellulose, thereby masking climatic signals. We compared LBM outbreak events in stable carbon and oxygen isotope chronologies of larch and their corresponding tree-ring widths from two high-elevation sites (1800–2200 m a.s.l.) in the Swiss Alps for the period AD 1900–2004 against isotope data obtained from non-host spruce (Picea abies). At each site, two age classes of tree individuals (150–250 and 450–550 years old) were sampled. Inclusion of the latter age class enabled one chronology to be extended back to AD 1650, and a comparison with long-term monthly resolved temperature data. Within the constraints of this local study, we found that: (1) isotopic ratios in tree rings of larch provide a strong and consistent climatic signal of temperature; (2) at all sites the isotope signatures were not disturbed by LBM outbreaks, as shown, for example, by exceptionally high significant correlations between non-host spruce and host larch chronologies; (3) below-average July to August temperatures and LBM defoliation events have been coupled for more than three centuries. Dampening of Alps-wide LBM cyclicity since the 1980s and the coincidence of recently absent cool summers in the European Alps reinforce the assumption of a strong coherence between summer temperatures and LBM defoliation events. Our results demonstrate that stable isotopes in tree-ring cellulose of larch are an excellent climate proxy enabling the analysis of climate-driven changes of LBM cycles in the long term.     

Can annual cyclicity of protozoan communities reflect water quality status in coastal ecosystems?

With many advantages, many ecological parameters of protozoan communities have been successfully used as a useful bioindicator for bioassessment of water quality in Chinese marine waters. However, as regard the response of the annual cyclicity of protozoan communities to seasonal environmental stress, a further investigation was needed. In this study, the cyclicity of annual variations in community patterns of biofilm-dwelling protozoa was studied based on an annual dataset. Samples were monthly collected, using glass slide method, at four stations, within a pollution gradient, in coastal waters of the Yellow Sea, northern China during a 1-year period. The cyclicity patterns of the microbiota represented a significant spatial variation among four stations. The low value of cyclicity coefficients occurred in heavily polluted area, while the high values were in less stressed areas. Correlation analysis showed that the cyclicity measure was significantly related to environmental variables ammonia, transparency and dissolved oxygen. Thus, it is suggested that the annual cyclicity of protozoan communities may be used as a potential bioindicator of bioassessment in marine ecosystems.     

Biochemical studies of isolated glial (muller) cells from the turtle retina

A method has been developed for the preparation of large numbers of glial (Muller) cells from the turtle retina. After proteolytic dissociation of the retina, Muller cells were separated from retinal neurons by velocity sedimentation at unit gravity. Fractions containing >90 percent morphologically identifiable Muller cells were prepared by this procedure. Fractions containing only Muller cells were obtained by drawing selected cells individually into a micropipette under visual observation. Biochemical analyses of isolated Muller cells showed that (a) these cells did not synthesize and accumulate acetylcholine, γ-aminobutyric acid, or catecholamines when incubated with appropriate radioactive precursors; (b) the specific activities of choline acetyltransferase (EC 2.3.1.6), glutamate decarboxylase (EC 4.1.1.15), and tyrosine hydroxylase (EC 1.14.16.2) in these cells were less than 2 percent of those found in the retina; (c) Muller cells, however, contained high activities of transmitter degrading enzymes-acetylcholinesterase (EC 3.1.1.7) and γ-aminobutyrate- transamine (EC 2.6.1.19); and (d) the cells also possessed high levels of two presumably glial-specific-enzymes-glutamine synthetase (EC 6.3.1.2) and carbonic anhydrase (EC 4.2.1.1). These results, together with other findings, suggest that Muller cells are not capable of neurotransmitter syntheses but possess the enzymes necessary for two important roles in the retina: (a) the inactivation of certain transmitters after synaptic transmission by uptake and degradation, and (b) the maintenance of acid-base balance and the provision of a stable microenvironment in the retina by the removal of metabolic products such as carbon dioxide and ammonia.     

High Diversity in Cretaceous Ichthyosaurs from Europe Prior to Their Extinction

Background

Ichthyosaurs are reptiles that inhabited the marine realm during most of the Mesozoic. Their Cretaceous representatives have traditionally been considered as the last survivors of a group declining since the Jurassic. Recently, however, an unexpected diversity has been described in Upper Jurassic–Lower Cretaceous deposits, but is widely spread across time and space, giving small clues on the adaptive potential and ecosystem control of the last ichthyosaurs. The famous but little studied English Gault Formation and ‘greensands’ deposits (the Upper Greensand Formation and the Cambridge Greensand Member of the Lower Chalk Formation) offer an unprecedented opportunity to investigate this topic, containing thousands of ichthyosaur remains spanning the Early–Late Cretaceous boundary.

Methodology/Principal Findings

To assess the diversity of the ichthyosaur assemblage from these sedimentary bodies, we recognized morphotypes within each type of bones. We grouped these morphotypes together, when possible, by using articulated specimens from the same formations and from new localities in the Vocontian Basin (France); a revised taxonomic scheme is proposed. We recognize the following taxa in the ‘greensands’: the platypterygiines ‘Platypterygius’ sp. and Sisteronia seeleyi gen. et sp. nov., indeterminate ophthalmosaurines and the rare incertae sedis Cetarthrosaurus walkeri. The taxonomic diversity of late Albian ichthyosaurs now matches that of older, well-known intervals such as the Toarcian or the Tithonian. Contrasting tooth shapes and wear patterns suggest that these ichthyosaurs colonized three distinct feeding guilds, despite the presence of numerous plesiosaur taxa.

Conclusion/Significance

Western Europe was a diversity hot-spot for ichthyosaurs a few million years prior to their final extinction. By contrast, the low diversity in Australia and U.S.A. suggests strong geographical disparities in the diversity pattern of Albian–early Cenomanian ichthyosaurs. This provides a whole new context to investigate the extinction of these successful marine reptiles, at the end of the Cenomanian.     

Orbital signatures in a late Miocene dinoflagellate record from Crete (Greece)

A high-resolution palynological study of the cyclically bedded Faneromeni section (upper Tortonian-lower Messinian) on Crete (Greece) is presented. This study aims to recognize orbitally-driven variations in the palynological record and to validate the age model based on the astronomical calibration of the sedimentary cycles. Four palynology-based environmental proxies were utilised using interpretations of fossil dinoflagellate associations based on modern ecological characteristics. Cross-spectral analysis between the proxy records and astronomical target curve, the 65°N summer insolation, yielded in most cases significant spectral power and coherence in the precession and/or obliquity frequency bands. Precession-controlled variations in the proxy records are related to lithology and indicate that maxima in continental input and minima in sea surface salinity coincide with sapropel formation. The influence of obliquity is most clearly reflected in the index of continental versus marine palynomorphs (S-D). The absence of a distinct time lag relative to obliquity indicates that the 41-kyr component in continental input is controlled by oscillations in regional Mediterranean climate rather than by glacial cyclicity. Phase relations in the different astronomical frequency bands of the spectrum, as compared with the Mediterranean Pliocene, essentially confirm the validity of the Miocene astronomical time scale. Finally, a major non-cyclic change in the palynological assemblage at 6.68 Ma indicates enhanced salinity and decreased river discharge. This shift coincides with a significant drop in sedimentation rate informally termed the “Early Messinian Sediment starvation Event”.