An investigation into the seasonality of the Pliocene southern North Sea basin: a sclerochronological approach

The Pliocene world c. 5.3 Ma to c. 2.58 Ma exhibited a relatively stable climate with a warmer global mean surface temperature than present-day by ~2 °C to 3 °C, and palaeoclimate analysis from this interval is used to understand climate drivers in ‘warmer world’. Previous oxygen isotope thermometry investigations of Pliocene southern North Sea Basin (SNSB) Aequipecten opercularis from the Coralline Crag Formation in Suffolk, UK repeatedly reveal evidence of a cold-temperate climate regime. Contrastingly, other biological proxies record a warm-temperate/sub-tropical regime. This investigation concentrated on oxygen, carbon and microgrowth increment widths (MIWS) of fossil shell material from Pliocene SNSB spanning an interval of~4.4 Ma to ~2.5 Ma. The study sites included shallow marine Pliocene formations from the western and eastern SNSB, the Ramsholt Member of the Coralline Crag Formation, Suffolk UK, and the Luchtbal Sands and Oorderen Sands Members of the Lillo Formation, Belgium, and the Oosterhout Formation in the Netherlands. Oxygen isotopic palaeotemperature results showed cooler summer temperatures than presently in the SNSB, which were reflective of a cool-temperate regime. There was no evidence of warm-temperate or sub-tropical summer palaeotemperatures in the Pliocene SNSB as suggested by other planktonic proxies. This investigation discussed the possible causal factors for the cooler – than- expected winter and summer palaeotemperatures in the ‘warmer’ Pliocene world as recorded by this proxy. Discrepancies between the cool summer benthic palaeotemperatures from the bivalves and the warmer sub-tropical or warm-temperate summer palaeotemperature estimations from planktonic biological proxies was rectified by the application of a theoretical summer stratification factor (SSF). However, rectifying the discrepancies between cooler (cold-temperate) benthic winter palaeotemperatures and the warmer winter palaeotemperatures from other proxies was difficult because stratification does not occur during the winter. Dormancy behaviours in the warm- temperate –sub-tropical organisms was proposed as a suitable mechanism to allow their coexistence with the cool-tolerant bivalves, which were able to grow and feed underneath the thermocline during the summer months. Therefore, the investigation showed how the Pliocene SNSB exhibited a greater seasonality than occurs presently in the SNSB. The driver for the cooler winter temperatures in the Pliocene SNSB was not identified. Localised explanations including continental wind effects, interannual variations in MOC strength, and increased storm activity in the winter bringing cooler water into the SNSB were all suggested as potential drivers. Global features of climate including interglacial/glacial cycles and orbital forcing effects were factors also proposed for the overall mixed palaeotemperature signal in the Pliocene SNSB.NER

Life history, environment and extinction of the scallop Carolinapecten eboreus (Conrad) in the Plio-Pleistocene of the U.S. eastern seaboard.

Plio-Pleistocene mass extinction of marine bivalves on the U.S. eastern seaboard has been attributed to declines in temperature and primary production. We investigate the relationship of growth rate in the scallop Carolinapecten eboreus to variation in these parameters to determine which contributed to its extinction. We use ontogenetic profiles of shell d18O to estimate growth rate and seasonal temperature, microgrowth-increment data to validate d18O-based figures for growth rate, and shell d13C to supplement assemblage evidence of production. Postlarval growth started in the spring/summer in individuals from the Middle Atlantic Coastal Plain but in the autumn/winter in some from the Gulf Coastal Plain. Growth rate typically declined with age and was usually higher in summer than winter. Many individuals died in winter but the largest forms typically died in spring, possibly on spawning for the first time. No individuals lived longer than two years and some grew exceedingly fast overall, up to 60% more rapidly than any other scallop species (, 145.7 mm in a year). Faster growth was generally achieved by secreting more rather than larger microgrowth increments. Some very fast-growing individuals lived in settings of high production and low temperature. No individuals grew slowly under high production whereas most if not all grew slowly under ‘average’ production and low temperature. In that the rapid growth evidently enabled by high production would have afforded protection from predators, Plio-Pleistocene decline in production was probably contributory to the extinction of C. eboreus. However, the negative impact of low temperature on growth under ‘average’ production suggests that temperature decline played some part.British Geological Survey (BUFI S157), NERC Isotope Goscience Facilities (IP-1351-1112), University of Derby (Research-Inspired Curriculum Fund

The use of seasonally resolved temperature data to identify the cause of marine climate change

On the continental shelf of the eastern USA, seasonal variation in water temperature is much lower south of Cape Hatteras than it is to the north as a result of the influence of warm currents, which raise winter temperature. High temperatures north of Cape Hatteras during the Pliocene have been attributed to greater northward penetration of warm currents in the absence of a feature analogous to Cape Hatteras. However, oxygen isotope thermometry using serial ontogenetic samples from scallops reveals a high seasonal temperature range at some horizons, suggesting that overall warming was the consequence of general climate change, with the absence of a ‘Cape Hatteras’ feature allowing greater southward penetration of cold currents, resulting in low winter temperatures at a southerly latitude. Evidence from other taxa indicates that at times seasonal variation in water temperature was quite low and that there was greater northward penetration of warm currents. This may relate to increases in vigour of the Gulf Stream. The study shows how seasonally resolved temperature data can assist identification of the driving forces of marine climate change.British Geological Survey (BUFI S157

Anti-predation strategy, growth rate and extinction amongst Pliocene scallops of the US eastern seaboard

Placopecten, Chesapecten and Carolinapecten are scallop (pectinid bivalve) genera occurring in the Pliocene of the US eastern seaboard. The first, present in the area today, is a smooth, streamlined form, adept at escaping predators by swimming (‘flight’ strategy). The other two, which are extinct, are plicate (‘ribbed’) forms. Plication facilitates a ‘resistance’ strategy towards predators which is benefited by large size and high shell thickness - maximally so if these states are achieved early in life. Oxygen isotope profiles show that early ontogenetic extensional growth in Pliocene Placopecten was at the same moderate rate as in modern Placopecten. By contrast, in Chesapecten it was as fast as in the fastest-growing modern scallop (c. 80 mm/annum), and accompanied by development of an unusually thick shell, while in Carolinapecten it was substantially faster still (<150 mm/annum). Rapid growth in Chesapecten and Carolinapecten was probably enabled by high primary productivity, for which there is evidence from sediment composition and the associated biota. The extinction of Chesapecten and Carolinapecten, and the survival of Placopecten, can be attributed to a decline in primary productivity which prevented a maximally effective ‘resistance’ strategy towards predators but had no deleterious impact on a ‘flight’ strategy.British Geological Survey (BUFI S157

Isotopic temperatures from the early and mid-pliocene of the US Middle Atlantic coastal plain, and their implications for the cause of regional marine climate change

Mean seasonal extreme temperatures on the seafloor calculated from the shell δ18O of the scallop Placopecten clintonius from the basal part of the early Pliocene Sunken Meadow Member (Yorktown Formation) in Virginia are very similar to those from the same horizon at the latitude of Cape Hatteras in North Carolina (~210 km to the south). The lowest and highest temperatures calculated from each shell (using δ18Oseawater = +0.7‰) give mean values for winter and summer of 8.4 ± 1.1 °C (± 1σ) and 18.2 ± 0.6 °C in Virginia, and 8.6 ± 0.4 °C and 16.5 ± 1.1 °C in North Carolina (respective median temperatures: 13.3 °C and 12.6 °C). Patterns of ontogenetic variation in δ18O, δ13C and microgrowth increment size indicate summer water-column stratification in both areas, with summer surface temperatures perhaps 6 °C higher than on the seafloor. The low winter paleotemperatures in both areas are most simply explained by the greater southward penetration of cool northern waters in the absence of a feature equivalent to Cape Hatteras. The same current configuration but a warmer general climate can account for the high benthic seasonal range (over 15.0 °C in some cases) but warmer median temperatures (15.7-21.3 °C) derived from existing δ18O data from scallops of the higher Yorktown Formation (using δ18Oseawater = +0.7‰ for the upper Sunken Meadow Member and δ18Oseawater = +1.1‰ for the mid-Pliocene Rushmere, Morgarts Beach and Moore House members). Existing δ18O data from the infaunal bivalve Mercenaria of the Rushmere Member yields a similarly high median temperature (21.6 °C) but a low seasonal range (9.2 °C), pointing to the periodic influence of warm currents, possibly at times when the Gulf Stream was exceptionally vigorous.British Geological Survey (BUFI S157 Studentship); UoD Research-Inspired Curriculum Fun

Sclerochronological evidence of pronounced seasonality from the late Pliocene of the southern North Sea basin and its implications

Oxygen isotope (δ18O) sclerochronology of benthic marine molluscs provides a means of reconstructing the seasonal range in seafloor temperature, subject to use of an appropriate equation relating shell δ18O to temperature and water δ18O, a reasonably accurate estimation of water δ18O, and due consideration of growth-rate effects. Taking these factors into account, δ18O data from late Pliocene bivalves of the southern North Sea basin (Belgium and the Netherlands) indicate a seasonal seafloor range a little smaller than now in the area. Microgrowth-increment data from Aequipecten opercularis, together with the species composition of the bivalve assemblage and aspects of preservation, suggest a setting below the summer thermocline for all but the latest material investigated. This implies a higher summer temperature at the surface than on the seafloor and consequently a greater seasonal range. A reasonable (3 ∘C) estimate of the difference between maximum seafloor and surface temperature under circumstances of summer stratification points to seasonal surface ranges in excess of the present value (12.4 ∘C nearby). Using a model-derived estimate of water δ18O (0.0 ‰), summer surface temperature was initially in the cool temperate range (20 ∘C) before reverting to cool temperate values (in conjunction with shallowing and a loss of summer stratification). This pattern is in agreement with biotic-assemblage evidence. Winter temperature was firmly in the cool temperate range (<10 ∘C) throughout, contrary to previous interpretations. Averaging of summer and winter surface temperatures for the MPWP provides a figure for annual sea surface temperature that is 2–3 ∘C higher than the present value (10.9 ∘C nearby) and in close agreement with a figure obtained by averaging alkenone and TEX86 temperatures for the MPWP from the Netherlands. These proxies, however, respectively, underestimate summer temperature and overestimate winter temperature, giving an incomplete picture of seasonality. A higher annual temperature than now is consistent with the notion of global warmth in the MPWP, but a low winter temperature in the southern North Sea basin suggests regional reduction in oceanic heat supply, contrasting with other interpretations of North Atlantic oceanography during the interval. Carbonate clumped isotope (Δ47) and biomineral unit thermometry offer means of checking the δ18O-based temperatures

Marine climate and hydrography of the Coralline Crag (early Pliocene, UK): isotopic evidence from 16 benthic invertebrate taxa.

The taxonomic composition of the biota of the Coralline Crag Formation (early Pliocene, eastern England) provides conflicting evidence of seawater temperature during deposition, some taxa indicating cool temperate conditions by analogy with modern representatives or relatives, others warm temperate to subtropical/tropical conditions. Previous isotopic (δ18O) evidence of seasonal seafloor temperatures from serial ontogenetic sampling of bivalve mollusk shells indicated cool temperate winter (< 10 °C) and/or summer (< 20 °C) conditions but was limited to nine profiles from two species, one ranging into and one occurring exclusively in cool temperate settings at present. We supplement these results with six further profiles from the species concerned and supply seven more from three other taxa (two supposedly indicative of warm waters) to provide an expanded and more balanced database. We also supply isotopic temperature estimates from 81 spot and whole-shell samples from these five taxa and 11 others, encompassing ‘warm’, ‘cool’ and ‘eurythermal’ forms by analogy with modern representatives or relatives. Preservation tests show no shell alteration. Subject to reasonable assumptions about water δ18O, the shell δ18O data either strongly indicate or are at least consistent with cool temperate seafloor conditions. The subtropical/tropical conditions suggested by the presence of the bryozoan Metrarabdotos did not exist. Microgrowth-increment and δ13C evidence indicate summer water-column stratification during deposition of the Ramsholt Member, unlike in the adjacent southern North Sea at present (well mixed due to shallow depth and strong tidal currents). Summer maximum surface temperature was probably about 5 °C above seafloor temperature and thus often slightly higher than now (17–19 °C rather than 16–17 °C), but only sometimes in the warm temperate range. Winter minimum surface temperature was below 10 °C and possibly the same as at present (6–7 °C). An expanded surface temperature range compared to now may reflect withdrawal of oceanic heat supply in conjunction with higher global temperature.British Geological Survey (BUFI S157), NERC Isotope Geoscience Facilities (IP-1155- 1109

Effects of antiplatelet therapy on stroke risk by brain imaging features of intracerebral haemorrhage and cerebral small vessel diseases: subgroup analyses of the RESTART randomised, open-label trial

Background Findings from the RESTART trial suggest that starting antiplatelet therapy might reduce the risk of recurrent symptomatic intracerebral haemorrhage compared with avoiding antiplatelet therapy. Brain imaging features of intracerebral haemorrhage and cerebral small vessel diseases (such as cerebral microbleeds) are associated with greater risks of recurrent intracerebral haemorrhage. We did subgroup analyses of the RESTART trial to explore whether these brain imaging features modify the effects of antiplatelet therapy

Effects of antiplatelet therapy after stroke due to intracerebral haemorrhage (RESTART): a randomised, open-label trial

Background: Antiplatelet therapy reduces the risk of major vascular events for people with occlusive vascular disease, although it might increase the risk of intracranial haemorrhage. Patients surviving the commonest subtype of intracranial haemorrhage, intracerebral haemorrhage, are at risk of both haemorrhagic and occlusive vascular events, but whether antiplatelet therapy can be used safely is unclear. We aimed to estimate the relative and absolute effects of antiplatelet therapy on recurrent intracerebral haemorrhage and whether this risk might exceed any reduction of occlusive vascular events. Methods: The REstart or STop Antithrombotics Randomised Trial (RESTART) was a prospective, randomised, open-label, blinded endpoint, parallel-group trial at 122 hospitals in the UK. We recruited adults (≥18 years) who were taking antithrombotic (antiplatelet or anticoagulant) therapy for the prevention of occlusive vascular disease when they developed intracerebral haemorrhage, discontinued antithrombotic therapy, and survived for 24 h. Computerised randomisation incorporating minimisation allocated participants (1:1) to start or avoid antiplatelet therapy. We followed participants for the primary outcome (recurrent symptomatic intracerebral haemorrhage) for up to 5 years. We analysed data from all randomised participants using Cox proportional hazards regression, adjusted for minimisation covariates. This trial is registered with ISRCTN (number ISRCTN71907627). Findings: Between May 22, 2013, and May 31, 2018, 537 participants were recruited a median of 76 days (IQR 29–146) after intracerebral haemorrhage onset: 268 were assigned to start and 269 (one withdrew) to avoid antiplatelet therapy. Participants were followed for a median of 2·0 years (IQR [1·0– 3·0]; completeness 99·3%). 12 (4%) of 268 participants allocated to antiplatelet therapy had recurrence of intracerebral haemorrhage compared with 23 (9%) of 268 participants allocated to avoid antiplatelet therapy (adjusted hazard ratio 0·51 [95% CI 0·25–1·03]; p=0·060). 18 (7%) participants allocated to antiplatelet therapy experienced major haemorrhagic events compared with 25 (9%) participants allocated to avoid antiplatelet therapy (0·71 [0·39–1·30]; p=0·27), and 39 [15%] participants allocated to antiplatelet therapy had major occlusive vascular events compared with 38 [14%] allocated to avoid antiplatelet therapy (1·02 [0·65–1·60]; p=0·92). Interpretation: These results exclude all but a very modest increase in the risk of recurrent intracerebral haemorrhage with antiplatelet therapy for patients on antithrombotic therapy for the prevention of occlusive vascular disease when they developed intracerebral haemorrhage. The risk of recurrent intracerebral haemorrhage is probably too small to exceed the established benefits of antiplatelet therapy for secondary prevention