Mineralogical and geochemical data from five lake sediment cores of Lake Donggi Cona

Sediments of Lake Donggi Cona on the northeastern Tibetan Plateau were studied to infer changes in the lacustrine depositional environment, related to climatic and non-climatic changes during the last 19 kyr. The lake today fills a 30 X 8 km big and 95 m deep tectonic basin, associated with the Kunlun Fault. The study was conducted on a sediment-core transect through the lake basin, in order to gain a complete picture of spatiotemporal environmental change. The recovered sediments are partly finely laminated and are composed of calcareous muds with variable amounts of carbonate micrite, organic matter, detrital silt and clay. On the basis of sedimentological, geochemical, and mineralogical data up to five lithological units (LU) can be distinguished that document distinct stages in the development of the lake system. The onset of the lowermost LU with lacustrine muds above basal sands indicates that lake level was at least 39 m below the present level and started to rise after 19 ka, possibly in response to regional deglaciation. At this time, the lacustrine environment was characterized by detrital sediment influx and the deposition of siliciclastic sediment. In two sediment cores, upward grain-size coarsening documents a lake-level fall after 13 cal ka BP, possibly associated with the late-glacial Younger Dryas stadial. From 11.5 to 4.3 cal ka BP, grainsize fining in sediment cores from the profundal coring sites and the onset of lacustrine deposition at a litoral core site (2m water depth) in a recent marginal bay of Donggi Cona document lake-level rise during the early tomid-Holocene to at least modern level. In addition, high biological productivity and pronounced precipitation of carbonate micrites are consistent with warm and moist climate conditions related to an enhanced influence of summer monsoon. At 4.3 cal ka BP the lake system shifted from an aragonite- to a calcite-dominated system, indicating a change towards a fully open hydrological lake system. The younger clay-rich sediments are moreover non-laminated and lack any diagenetic sulphides, pointing to fully ventilated conditions, and the prevailing absence of lake stratification. This turning point in lake history could imply either a threshold response to insolation-forced climate cooling or a response to a non-climatic trigger, such as an erosional event or a tectonic pulse that induced a strong earthquake, which is difficult to decide from our data base.

In-situ Iron isotope measurements and mineral compositions from ODP Hole 206-1256D and IODP Hole 335-U1256D

In-situ Fe isotope measurements have been carried out to estimate the impact of the hydrothermal metamorphic overprint on the Fe isotopic composition of Fe-Ti-oxides and Fe-sulfides of the different lithologies of the drilled rocks from IODP Hole 1256D (eastern equatorial Pacific; 15 Ma crust formed at the East Pacific Rise). Most igneous rocks normally have a very restricted range in their 56Fe/54Fe ratio. In contrast, Fe isotope compositions of hot fluids (> 300 °C) from mid-ocean-ridge spreading centers define a narrow range that is shifted to lower delta 56Fe values by 0.2 per mil - 0.5 per mil as compared to igneous rocks. Therefore, it is expected that mineral phases that contain large amounts of Fe are especially affected by the interaction with a fluid that fractionates Fe isotopes during exsolution/precipitation of those minerals. We have used a femtosecond UV-Laser ablation system to determine mineral 56Fe/54Fe ratios of selected samples with a precision of < 0.1 per mil (2 sigma level) at micrometer-scale. We have found significant variations of the delta 56Fe (IRMM-014) values in the minerals between different samples as well as within samples and mineral grains. The overall observed scale of delta 56Fe (magnetite) in 1256D rocks ranges from - 0.12 to + 0.64 per mil, and of delta 56Fe (ilmenite) from - 0.77 to + 0.01 per mil. Pyrite in the lowermost sheeted dike section is clearly distinguishable from the other investigated lithological units, having positive delta 56Fe values between + 0.29 and + 0.56 per mil, whereas pyrite in the other samples has generally negative delta 56Fe values from - 1.10 to - 0.59 permil. One key observation is that the temperature dependent inter-mineral fractionations of Fe isotopes between magnetite and ilmenite are systematically shifted towards higher values when compared to theoretically expected values, while synthesized, well equilibrated magnetite-ilmenite pairs are compatible with the theoretical predictions. Theoretical considerations including beta-factors of different aqueous Fe-chlorides and Rayleigh-type fractionations in the presence of a hydrous, chlorine-bearing fluid can explain this observation. The disagreement between observed and theoretical equilibrium fractionation, the fact that magnetite, in contrast to ilmenite shows a slight downhole trend in the delta 56Fe values, and the observation of small scale heterogeneities within single mineral grains imply that a general re-equilibration of the magnetite-ilmenite pairs is overprinted by kinetic fractionation effects, caused by the interaction of magnetite/ilmenite with hydrothermal fluids penetrating the upper oceanic crust during cooling, or incomplete re-equilibration at low temperatures. Furthermore, the observation of significant small-scale variations in the 56Fe/54Fe ratios of single minerals in this study highlights the importance of high spatial-resolution-analyses of stable isotope ratios for further investigations.

(Appendix A) Geochemical composition of snow and eolian mineral dust from Berkner Island ice sheet, Antarctica

The Sr and Nd isotopic composition of dust extracted from recent snow layers at the top of Berkner Island ice sheet (located within the Filchner-Ronne Ice Shelf at the southern end of the Weddell Sea) enables us, for the first time, to document dust provenance in Antarctica outside the East Antarctic Plateau (EAP) where all previous studies based on isotopic fingerprinting were carried out. Berkner dust displays an overall crust-like isotopic signature, characterized by more radiogenic 87Sr/86Sr and much less radiogenic 143Nd/144Nd compared to dust deposited on the EAP during glacial periods. Differences with EAP interglacial dust are not as marked but still significant, indicating that present-day Berkner dust provenance is distinct, at least to some extent, from that of the dust reaching the EAP. The fourteen snow-pit sub-seasonal samples that were obtained span a two-year period (2002-2003) and their dust Sr and Nd isotopic composition reveals that multiple sources are at play over a yearly time period. Southern South America, Patagonia in particular, likely accounts for part of the observed spring/summer dust deposition maxima, when isotopic composition is shifted towards 'younger' isotopic signatures. In the spring, possible additional inputs from Australian sources would also be supported by the data. Most of the year, however, the measured isotopic signatures would be best explained by a sustained background supply from putative local sources in East Antarctica, which carry old-crust-like isotopic fingerprints. Whether the restricted East Antarctic ice-free areas produce sufficient eolian material has yet to be substantiated however. The fact that large (> 5 µm) particles represent a significant fraction of the samples throughout the entire time-series supports scenarios that involve contributions from proximal sources, either in Patagonia and/or Antarctica (possibly including snow-free areas in the Antarctic Peninsula and other areas as well). This also indicates that additional dust transport, which does not reach the EAP, must occur at low-tropospheric levels to this coastal sector of Antarctica.

Plio-Pleistocene age-depth points and occurrence of hiatuses in ODP sites of the Southern Ocean

Changes in Atlantic deep water circulation were reconstructed by comparing the benthic foraminiferal delta13C record at ODP Site 1090 in the South Atlantic with similar records from the North Atlantic (Sites 982, 607, 925, 929) and deep Pacific (Site 849) oceans. Important deep water circulation changes occurred in the early Pleistocene at 1.55 Myr and during the Mid-Pleistocene Transition at 0.9 Myr. At 1.55 Myr, glacial delta13C values in the Southern Ocean became significantly lower than those in the deep Pacific, establishing a pattern that persisted throughout the late Pleistocene. We propose that the lowering of delta13C values of Southern Component Water (SCW) at this time resulted from expansion of sea ice and reduced ventilation of deep water during glacial periods after marine isotope stage 52. Accompanying this change in Southern Ocean deep water circulation was enhanced interhemispheric coupling between the North and South Atlantic after 1.55 Myr. At ~0.9 Myr, the magnitude of glacial-to-interglacial variabilityin delta13C increased and shifted to a longer frequency (100 kyr) along with oceanic delta18O (ice volume). Calculation of percent Northern Component Water (NCW) using Site 1090 as the SCW end member yielded 20-30% less reduction of NCW during glacial periods of the late Pleistocene. Also, a trend toward reduced glacial suppression of NCW during the past 400 kyr is not evident. The apparent decoupling of ice volume and deep water circulation reported previously maybe an artifact of using a Pacific, rather than a Southern Ocean, carbon isotopic record to calculate past mixing ratios of NCW and SCW.

CaCO3, foraminifer fragmentation, benthic carbon isotopes, coarse lithic counts and radiogenic isotopes (Sr, Nd, Pb) of the Pliocene and earliest Pleistocene terrigenous component (3.3-2.4 Ma) of IODP Site 306-U1313

We present Plio-Pleistocene records of sediment color, %CaCO3, foraminifer fragmentation, benthic carbon isotopes (d13C) and radiogenic isotopes (Sr, Nd, Pb) of the terrigenous component from IODP Site U1313, a reoccupation of benchmark subtropical North Atlantic Ocean DSDP Site 607. We show that (inter)glacial cycles in sediment color and %CaCO3 pre-date major northern hemisphere glaciation and are unambiguously and consistently correlated to benthic oxygen isotopes back to 3.3 million years ago (Ma) and intermittently so probably back to the Miocene/Pliocene boundary. We show these lithological cycles to be driven by enhanced glacial fluxes of terrigenous material (eolian dust), not carbonate dissolution (the classic interpretation). Our radiogenic isotope data indicate a North American source for this dust (~3.3-2.4 Ma) in keeping with the interpreted source of terrestrial plant wax-derived biomarkers deposited at Site U1313. Yet our data indicate a mid latitude provenance regardless of (inter)glacial state, a finding that is inconsistent with the biomarker-inferred importance of glaciogenic mechanisms of dust production and transport. Moreover, we find that the relation between the biomarker and lithogenic components of dust accumulation is distinctly non-linear. Both records show a jump in glacial rates of accumulation from Marine Isotope Stage, MIS, G6 (2.72 Ma) onwards but the amplitude of this signal is about 3-8 times greater for biomarkers than for dust and particularly extreme during MIS 100 (2.52 Ma). We conclude that North America shifted abruptly to a distinctly more arid glacial regime from MIS G6, but major shifts in glacial North American vegetation biomes and regional wind fields (exacerbated by the growth of a large Laurentide Ice Sheet during MIS 100) likely explain amplification of this signal in the biomarker records. Our findings are consistent with wetter-than-modern reconstructions of North American continental climate under the warm high CO2 conditions of the Early Pliocene but contrast with most model predictions for the response of the hydrological cycle to anthropogenic warming over the coming 50 years (poleward expansion of the subtropical dry zones).

Pigments at time series station DYNAPROC

Phytoplankton taxonomic pigments and primary production were measured at the JGOFS-France time-series station DYFAMED in the northwestern Mediterranean Sea during May 1995 to investigate changes in phytoplankton composition and the biogeochemical implications (DYNAPROC experiment). The study period covered the transitional situation from late spring bloom to pre-oligotrophic. The late spring bloom situation, occurring at the beginning of the study, revealed high chlorophyll a concentrations (maximum 3 mg/m**3 at 30 m) and high primary production (maximum 497 mg C/m**2/ 14 h). At the end of the experiment, the trophic regime shifted towards pre-oligotrophic and was characterized by lower chlorophyll a concentrations (<1 mg/m**3), although primary production still remained high (659 mg C/m**2/ 14 h). At termination of the spring bloom, the phytoplankton community was composed of chromophyte nanoflagellates (38±4%), diatoms (29±2%), cryptophytes (12±1%) and cyanobacteria (8±1%). During the transition to the pre-oligotrophic period, the contribution of small cells increased (e.g. cyanobacteria 18±2%, green flagellates 5±1%). Vertical profiles of pigments revealed a partition of the phytoplankton groups: cyanobacteria were most abundant in the surface layer, nanoflagellates containing 19'-HF+19'BF at the depth of chlorophyll maximum, whereas diatoms were located below the chlorophyll maximum. At termination of the spring bloom, a wind event induced vertical transport of nutrients into the euphotic layer. Phytoplankton groups responded differently to the event: initially, diatom concentrations increased (for 24 h) then rapidly decreased. In contrast, all others groups decreased just after the event. The long-term effect was a decrease of biomass of dominant groups (diatoms and chromophyte nanoflagellates), which accelerated the community succession and hence contributed to the oligotrophic transition.

Global cooling during the Eocene Oligocene Transition

About 34 million years ago, Earth's climate shifted from a relatively ice-free world to one with glacial conditions on Antarctica characterized by substantial ice sheets. How Earth's temperature changed during this climate transition remains poorly understood, and evidence for Northern Hemisphere polar ice is controversial. Here, we report proxy records of sea surface temperatures from multiple ocean localities and show that the high-latitude temperature decrease was substantial and heterogeneous. High-latitude (45 degrees to 70 degrees in both hemispheres) temperatures before the climate transition were ~20°C and cooled an average of ~5°C. Our results, combined with ocean and ice-sheet model simulations and benthic oxygen isotope records, indicate that Northern Hemisphere glaciation was not required to accommodate the magnitude of continental ice growth during this time.

X-ray fluorescence (XRF) Ca intensity of sediment cores MD04-2760 and MD04-2788

Accelerator mass spectrometry (AMS) radiocarbon dating of ostracod and gastropod shells from the southwestern Black Sea cores combined with tephrochronology provides the basis for studying reservoir age changes in the lateglacial Black Sea. The comparison of our data with records from the northwestern Black Sea shows that an apparent reservoir age of ~1450 14C yr found in the glacial is characteristic of a homogenized water column. This apparent reservoir age is most likely due to the hardwater effect. Though data indicate that a reservoir age of ~1450 14C yr may have persisted until the Bølling-Allerød warm period, a comparison with the GISP2 ice-core record suggests a gradual reduction of the reservoir age to ~1000 14C yr, which might have been caused by dilution effects of inflowing meltwater. During the Bølling-Allerød warm period, soil development and increased vegetation cover in the catchment area of the Black Sea could have hampered erosion of carbonate bedrock, and hence diminished contamination by "old" carbon brought to the Black Sea basin by rivers. A further reduction of the reservoir age most probably occurred contemporary to the precipitation of inorganic carbonates triggered by increased phytoplankton activity, and was confined to the upper water column. Intensified deep water formation subsequently enhanced the mixing/convection and renewal of intermediate water. During the Younger Dryas, the age of the upper water column was close to 0 yr, while the intermediate water was ~900 14C yr older. The first inflow of saline Mediterranean water, at ~8300 14C yr BP, shifted the surface water age towards the recent value of ~400 14C yr.

Plant distribution and climate data across western North America

Using an extensive network of occurrence records for 293 plant species collected over the past 40 years across a climatically diverse geographic section of western North America, we find that plant species distributions were just as likely to shift upwards (i.e., towards higher elevations) as downward (i.e., towards lower elevations) - despite consistent warming across the study area. Although there was no clear directional response to climate warming across the entire study area, there was significant region-to region- variation in responses (i.e. from as many as 73% to as few as32% of species shifting upward or downward). To understand the factors that might be controlling region-specific distributional shifts, we explored the relationship between the direction of change in distribution limits and the nature of recent climate change. We found that the direction of distribution limit shifts was explained by an interaction between the rate of change in local summer temperatures and seasonal precipitation. Specifically, species shifted upward at their upper elevational limit when snowfall declined at slower rates and minimum temperatures increased. By contrast, species shifted upwards at their lower elevation limit when maximum temperatures increased or both temperature and precipitation decreased. Our results suggest that future species' elevational distribution shifts will be complex, depending on the interaction between seasonal temperature and precipitation change.

Revised composite depth scale, Fe content and orbital tuning of Middle Eocene Climate Optimum samples from ODP Site 207-1260

A high-resolution stratigraphy is essential toward deciphering climate variability in detail and understanding causality arguments of events in earth history. Because the highly dynamic middle to late Eocene provides a suitable testing ground for carbon cycle models for a waning warm world, an accurate time scale is needed to decode climate-driving mechanisms. Here we present new results from ODP Site 1260 (Leg 207) which covers a unique expanded middle Eocene section (magnetochrons C18r to C20r, late Lutetian to early Bartonian) of the tropical western Atlantic including the chron C19r transient hyperthermal event and the Middle Eocene Climate Optimum (MECO). To establish a detailed cyclostratigraphy we acquired a distinctive iron intensity records by XRF scanning Site 1260 cores. We revise the shipboard composite section, establish a cyclostratigraphy and use the exceptional eccentricity modulated precession cycles for orbital tuning. The new astrochronology revises the age of magnetic polarity chrons C19n to C20n, validates the position of very long eccentricity minima at 40.2 and 43.0 Ma in the orbital solutions, and extends the Astronomically Tuned Geological Time Scale back to 44 Ma. For the first time the new data provide clear evidence for an orbital pacing of the chron C19r event and a likely involvement of the very long eccentricity cycle contributing to the evolution of the MECO.

Benthic foraminiferal d18O and Mg/Ca studies of sediment cores off tropical NW Africa

Benthic foraminiferal d18O and Mg/Ca of sediment cores off tropical NW Africa are used to study the properties of Atlantic central waters during the Last Glacial Maximum (LGM) and Heinrich Stadial 1 (HS1). We combined our core top data with published results to develop a new Mg/Ca-temperature calibration for Planulina ariminensis, which shows a Mg/Ca-temperature sensitivity of 0.19 mmol/mol per °C. Estimates of the LGM and HS1 thermocline temperatures are comparable to the present-day values between 200 and 400 m water depth, but were 1.2-1.5°C warmer at 550-570 m depth. The HS1 thermocline waters (200-570 m depth) did not show any warming relative to the LGM. This is in contrast to previous climate model studies, which concluded that tropical Atlantic thermocline waters warmed significantly when Atlantic meridional overturning circulation was reduced. However, our results suggest that thermocline temperatures of the northeastern tropical Atlantic show no pronounced sensitivity to changes in the thermohaline circulation during glacial periods. In contrast, we find a significant increase in thermocline-water salinity during the LGM (200-550 m depth) and HS1 (200-400 m depth) with respect to the present-day, which we relate to changes in the wind-driven circulation. We infer that the LGM thermocline (200-550 m depth) and the HS1 upper thermocline (200-400 m depth) in the northeastern tropical Atlantic was ventilated by surface waters from the North Atlantic rather than the southern-sourced waters. This suggests that the frontal zone between the modern South Atlantic and North Atlantic Central Waters was probably shifted southward during the LGM and HS1.

Haemolymph inorganic ion composition, physiology and behaviour dependent on temperature and ambient magnesium concentration in early life history stages of Paralomis granulosa (Decapoda, Anomura, Lithodidae)

Marine brachyuran and anomuran crustaceans are completely absent from the extremely cold (-1.8 °C) Antarctic continental shelf, but caridean shrimps are abundant. This has at least partly been attributed to low capacities for magnesium excretion in brachyuran and anomuran lithodid crabs ([Mg2+]HL = 20-50 mmol/L) compared to caridean shrimp species ([Mg2+]HL = 5-12 mmol/L). Magnesium has an anaesthetizing effect and reduces cold tolerance and activity of adult brachyuran crabs. We investigated whether the capacity for magnesium regulation is a factor that influences temperature-dependent activity of early ontogenetic stages of the Sub-Antarctic lithodid crab Paralomis granulosa. Ion composition (Na+, Mg2+, Ca2+, Cl-, [SO4]2-) was measured in haemolymph withdrawn from larval stages, the first and second juvenile instars (crabs I and II) and adult males and females. Magnesium excretion improved during ontogeny, but haemolymph sulphate concentration was lowest in the zoeal stages. Neither haemolymph magnesium concentrations nor Ca2+:Mg2+ ratios paralleled activity levels of the life stages. Long-term (3 week) cold exposure of crab I to 1 °C caused a significant rise of haemolymph sulphate concentration and a decrease in magnesium and calcium concentrations compared to control temperature (9 °C). Spontaneous swimming activity of the zoeal stages was determined at 1, 4 and 9 °C in natural sea water (NSW, [Mg2+] = 51 mmol/L) and in sea water enriched with magnesium (NSW + Mg2+, [Mg2+] = 97 mmol/L). It declined significantly with temperature but only insignificantly with increased magnesium concentration. Spontaneous velocities were low, reflecting the demersal life style of the zoeae. Heart rate, scaphognathite beat rate and forced swimming activity (maxilliped beat rate, zoea I) or antennule beat rate (crab I) were investigated in response to acute temperature change (9, 6, 3, 1, -1 °C) in NSW or NSW + Mg2+. High magnesium concentration reduced heart rates in both stages. The temperature-frequency curve of the maxilliped beat (maximum: 9.6 beats/s at 6.6 °C in NSW) of zoea I was depressed and shifted towards warmer temperatures by 2 °C in NSW + Mg2+, but antennule beat rate of crab I was not affected. Magnesium may therefore influence cold tolerance of highly active larvae, but it remains questionable whether the slow-moving lithodid crabs with demersal larvae would benefit from an enhanced magnesium excretion in nature.

Geochemistry of foraminifera from sediment cores of the central California margin

Under present climate conditions, convection at high latitudes of the North Pacific is restricted to shallower depths than in the North Atlantic. To what extent this asymmetry between the two ocean basins was maintained over the past 20 kyr is poorly known because there are few unambiguous proxy records of ventilation from the North Pacific. We present new data for two sediment cores from the California margin at 800 and 1600 m depth to argue that the depth of ventilation shifted repeatedly in the northeast Pacific over the course of deglaciation. The evidence includes benthic foraminiferal Cd/Ca, 18O/16O, and 13C/12C data as well as radiocarbon age differences between benthic and planktonic foraminifera. A number of features in the shallower of the two cores, including an interval of laminated sediments, are consistent with changes in ventilation over the past 20 kyr suggested by alternations between laminated and bioturbated sediments in the Santa Barbara Basin and the Gulf of California [Keigwin and Jones, 1990 doi:10.1029/PA005i006p01009; Kennett and Ingram, 1995 doi:10.1038/377510a0; Behl and Kennett, 1996 doi:10.1038/379243a0]. Data from the deeper of the two California margin cores suggest that during times of reduced ventilation at 800 m, ventilation was enhanced at 1600 m depth, and vice versa. This pronounced depth dependence of ventilation needs to be taken into account when exploring potential teleconnections between the North Pacific and the North Atlantic.

Mid-Pleistocene planktonic foraminiferal composition and IRD content in ODP Sites 162-980 and 162-984

We investigated surface and deep ocean variability in the subpolar North Atlantic from 1000 to 500 thousand years ago (ka) based on two Ocean Drilling Program (ODP) sites, Feni drift site 980 (55°29'N, 14°42'W) and Bjorn drift site 984 (61°25'N, 24°04'W). Benthic foraminiferal stable isotope data, planktic foraminiferal faunas, ice-rafted debris data, and faunally based sea-surface temperature estimates help test the hypothesis that oceanographic changes in the North Atlantic region were associated with the onset of the 100-kyr world during the mid-Pleistocene revolution. Based on percentage of Neogloboquadrina pachyderma (s) records from both sites, surface waters during interglacials and glacials were cooler in the mid-Pleistocene than during marine isotope stages (MIS) 5 and 6. In particular, interglaciations at Bjorn drift site 984 were significantly cooler. Faunal evidence suggests that the interglacial Arctic front shifted from a position between the two sites to a position northwest of Bjorn drift site 984 after ca. 610 ka. As during the late Pleistocene, we find faunal evidence for lagging surface warmth at most of the glacial initiations during the mid-Pleistocene. Each initiation is associated with high benthic d13C values that are maintained into the succeeding glaciation, which we term "lagging NADW production." These findings indicate that lagging warmth and lagging NADW production are robust features of the regional climate system that persist in the middle to late Pleistocene.

Phenotypic evolution in a venerid bivalve species lineage from the late Middle Miocene Central Paratethys Sea: a multi-approach morphometric analysis

A morphometric analysis was performed for the late Middle Miocene bivalve species lineage of Polititapes tricuspis (Eichwald, 1829) (Veneridae: Tapetini). Specimens from various localities grouped into two stratigraphically successive biozones, i.e. the upper Ervilia Zone and the Sarmatimactra Zone, were investigated using a multi-method approach. A Generalized Procrustes Analysis was computed for fifteen landmarks, covering characteristics of the hinge, muscle scars, and pallial line. The shell outline was separately quantified by applying the Fast Fourier Transform, which redraws the outline by fitting in a combination of trigonometric curves. Shell size was calculated as centroid size from the landmark configuration. Shell thickness, as not covered by either analysis, was additionally measured at the centroid. The analyses showed significant phenotypic differentiation between specimens from the two biozones. The bivalves become distinctly larger and thicker over geological time and develop circular shells with stronger cardinal teeth and a deeper pallial sinus. Data on the paleoenvironmental changes in the late Middle Miocene Central Paratethys Sea suggest the phenotypic shifts to be functional adaptations. The typical habitats for Polititapes changed to extensive, very shallow shores exposed to high wave action and tidal activity. Caused by the growing need for higher mechanical stability, the bivalves produced larger and thicker shells with stronger cardinal teeth. The latter are additionally shifted towards the hinge center to compensate for the lacking lateral teeth and improve stability. The deepening pallial sinus is related to a deeper burrowing habit, which is considered to impede being washed out in the new high-energy settings.