Description: We present and examine a multi-sensor global compilation of mid-Holocene (MH) sea surface temperatures (SST), based on Mg/Ca and alkenone palaeothermometry and reconstructions obtained using planktonic foraminifera and organic-walled dinoflagellate cyst census counts. We assess the uncertainties originating from using different methodologies and evaluate the potential of MH SST reconstructions as a benchmark for climate-model simulations. The comparison between different analytical approaches (time frame, baseline climate) shows the choice of time window for the MH has a negligible effect on the reconstructed SST pattern, but the choice of baseline climate affects both the magnitude and spatial pattern of the reconstructed SSTs. Comparison of the SST reconstructions made using different sensors shows significant discrepancies at a regional scale, with uncertainties often exceeding the reconstructed SST anomaly. Apparent patterns in SST may largely be a reflection of the use of different sensors in different regions. Overall, the uncertainties associated with the SST reconstructions are generally larger than the MH anomalies. Thus, the SST data currently available cannot serve as a target for benchmarking model simulations.

Description: Warming and changes in ocean carbonate chemistry alter marine coastal ecosystems at an accelerating pace. The interaction between these stressors has been the subject of recent studies on reef organisms such as corals, bryozoa, molluscs, and crustose coralline algae. Here we investigated the combined effects of elevated sea surface temperatures and pCO2 on two species of photosymbiont-bearing coral reef Foraminifera: Heterostegina depressa (hosting diatoms) and Marginopora vertebralis (hosting dinoflagellates). The effects of single and combined stressors were studied by monitoring survivorship, growth, and physiological parameters, such as respiration, photochemistry (pulse amplitude modulation fluorometry and oxygen production), and chl a content. Specimens were exposed in flow-through aquaria for up to seven weeks to combinations of two pCO2 (~790 and ~490 µatm) and two temperature (28 and 31 °C) regimes. Elevated temperature had negative effects on the physiology of both species. Elevated pCO2 had negative effects on growth and apparent photosynthetic rate in H.depressa but a positive effect on effective quantum yield. With increasing pCO2, chl a content decreased in H. depressa and increased in M. vertebralis. The strongest stress responses were observed when the two stressors acted in combination. An interaction term was statistically significant in half of the measured parameters. Further exploration revealed that 75 % of these cases showed a synergistic (= larger than additive) interaction between the two stressors. These results indicate that negative physiological effects on photosymbiont-bearing coral reef Foraminifera are likely to be stronger under simultaneous acidification and temperature rise than what would be expected from the effect of each of the stressors individually.

Description: The eastern Mediterranean is a hotspot of biological invasions. Numerous species of Indo-pacific origin have colonized the Mediterranean in recent times, including tropical symbiont-bearing foraminifera. Among these is the species Pararotalia calcariformata. Unlike other invasive foraminifera, this species has been discovered only two decades ago and is restricted to the eastern Mediterranean coast. Combining ecological, genetic and physiological observations, we attempt to explain the recent invasion of this species in the Mediterranean Sea. Using morphological and genetic data, we confirm the species attribution to P. calcariformata McCulloch 1977 and identify its symbionts as a consortium of diatom species dominated by Minutocellus polymorphus. We document photosynthetic activity of its endosymbionts using Pulse Amplitude Modulated Fluorometry and test the effects of elevated temperatures on growth rates of asexual offspring. The culturing of asexual offspring for 120 days shows a 30-day period of rapid growth followed by a period of slower growth. A subsequent 48-day temperature sensitivity experiment indicates a similar developmental pathway and high growth rate at 28°C, whereas an almost complete inhibition of growth was observed at 20°C and 35°C. This indicates that the offspring of this species may have lower tolerance to cold temperatures than what would be expected for species native to the Mediterranean. We expand this hypothesis by applying a Species Distribution Model (SDM) based on modern occurrences in the Mediterranean using three environmental variables: irradiance, turbidity and yearly minimum temperature. The model reproduces the observed restricted distribution and indicates that the range of the species will drastically expand westwards under future global change scenarios. We conclude that P. calcariformata established a population in the Levant because of the recent warming in the region. In line with observations from other groups of organisms, our results indicate that continued warming of the eastern Mediterranean will facilitate the invasion of more tropical marine taxa into the Mediterranean, disturbing local biodiversity and ecosystem structure.

Description: Fossil shells of planktonic foraminifera serve as the prime source of information on past changes in surface ocean conditions. Because the population size of planktonic foraminifera species changes throughout the year, the signal preserved in fossil shells is biased towards the conditions when species production was at its maximum. The amplitude of the potential seasonal bias is a function of the magnitude of the seasonal cycle in production. Here we use a planktonic foraminifera model coupled to an ecosystem model to investigate to what degree seasonal variations in production of the species Neogloboquadrina pachyderma may affect paleoceanographic reconstructions during Heinrich Stadial 1 (~18-15 cal. ka B.P.) in the North Atlantic Ocean. The model implies that during Heinrich Stadial 1 the maximum seasonal production occurred later in the year compared to the Last Glacial Maximum (~21-19 cal. ka B.P.) and the pre-industrial era north of 30 ºN. A diagnosis of the model output indicates that this change reflects the sensitivity of the species to the seasonal cycle of sea-ice cover and food supply, which collectively lead to shifts in the modeled maximum production from the Last Glacial Maximum to Heinrich Stadial 1 by up to six months. Assuming equilibrium oxygen isotopic incorporation in the shells of N. pachyderma, the modeled changes in seasonality would result in an underestimation of the actual magnitude of the meltwater isotopic signal recorded by fossil assemblages of N. pachyderma wherever calcification is likely to take place.

Description: Invasive species allow an investigation of trait retention and adaptations after exposure to new habitats. Recent work on corals from the Gulf of Aqaba (GoA) shows that tolerance to high temperature persists thousands of years after invasion, without any apparent adaptive advantage. Here we test whether thermal tolerance retention also occurs in another symbiont-bearing calcifying organism. To this end, we investigate the thermal tolerance of the benthic foraminifera Amphistegina lobifera from the GoA (29° 30.14167 N 34° 55.085 E) and compare it to a recent "Lessepsian invader population" from the Eastern Mediterranean (EaM) (32° 37.386 N, 34°55.169 E). We first established that the studied populations are genetically homogenous but distinct from a population in Australia, and that they contain a similar consortium of diatom symbionts, confirming their recent common descent. Thereafter, we exposed specimens from GoA and EaM to elevated temperatures for three weeks and monitored survivorship, growth rates and photophysiology. Both populations exhibited a similar pattern of temperature tolerance. A consistent reduction of photosynthetic dark yields was observed at 34°C and reduced growth was observed at 32°C. The apparent tolerance to sustained exposure to high temperature cannot have a direct adaptive importance, as peak summer temperatures in both locations remain 〈32°C. Instead, it seems that in the studied foraminifera tolerance to high temperature is a conservative trait and the EaM population retained this trait since its recent invasion. Such pre-adaptation to higher temperatures confers A. lobifera a clear adaptive advantage in shallow and episodically high temperature environments in the Mediterranean under further warming.

Description: The transition from benthos to plankton requires multiple adaptations, yet so far it remains unclear how these are acquired in the course of the transition. To investigate this process, we analyzed the genetic diversity and distribution patterns of a group of foraminifera of the genus Bolivina with a tychopelagic mode of life (same species occurring both in benthos and plankton). We assembled a global sequence dataset for this group from single-cell DNA extractions and occurrences in metabarcodes from pelagic environmental samples. The pelagic sequences all cluster within a single monophyletic clade within Bolivina. This clade harbors three distinct genetic lineages, which are associated with incipient morphological differentiation. All lineages occur in plankton and benthos, but only one lineage shows no limit to offshore dispersal and has been shown to grow in the plankton. These observations indicate that the emergence of buoyancy regulation within the clade preceded the evolution of pelagic feeding and that the evolution of both traits was not channeled into a full transition into the plankton. We infer that in foraminifera, colonization of the planktonic niche may occur by sequential cooptation of independently acquired traits, with holoplanktonic species being recruited from tychopelagic ancestors.

Description: We present a new coccolithophore productivity reconstruction spanning the last 300 ka in core GeoB12613-1 retrieved from the western tropical Indian Ocean (IO), an area that mainly derives its warm and oligotrophic surface waters from the eastern IO. Application of a calibrated assemblage-based productivity index indicates a reduction in estimated primary productivity (EPP) from 300 ka to the present, with reconstructed EPP values ranging from 91 to 246 g C/m2/yr. Coccolithophore assemblages and coccolith fraction Sr/Ca indicate three main phases of productivity change, with major changes at 160 and 46 ka. The productivity and water-column stratification records show both dominant precession and obliquity periodicities, which appear to control the paleoproductivity in the study area over the last two glacial-interglacial cycles. Shallowing of the thermocline due to strengthening of the trade winds in response to insolation maxima resulted to peaks in EPP. Comparison with the eastern IO productivity and stratification coccolithophore data reveals good correspondence with our records, indicating a strong tropical Pacific influence in our study area. Both of these records show high productivity from 300 ka to 160 ka, interpreted to be due to stronger Walker Circulation while the declining productivity from 160 ka to the present day is a consequence of its weakening intensity.

Description: The chronology of deglacial meltwater pulses from the Laurentide Ice Sheet is well documented. However, the deglacial history of the North American-Arctic (north-eastern Laurentide and Innuitian) and western Greenland ice sheets draining into the Labrador Sea via Baffin Bay is less well constrained. Here we present new high-resolution, radiocarbon-dated records from the central Baffin Bay spanning ~17 to 10 kyr BP and documenting the full deglacial history of Baffin Bay. Sedimentological and geochemical data confirm the presence of two periods of enhanced detrital carbonate delivery, termed Baffin Bay Detrital Carbonate Events (BBDCs). These events are dated to ~14.2-13.7 kyr BP and ~12.7-11 kyr BP. They are synchronous across Baffin Bay and their mineralogical signature indicates a common source of detrital carbonate from northern Baffin Bay. The first event, BBDC 1, postdates Heinrich Event 1 and the second event, BBDC 0, predates the recently revised timing of Heinrich Event 0. The onset of the BBDC events appears not to be systematically linked to Greenland temperature change as they occur during both interstadial and stadial periods. This indicates that deglaciation of North American-Arctic and western Greenland ice sheets with the associated iceberg and meltwater discharge were decoupled from the dominant North Atlantic climate mode, where iceberg discharge events from the Laurentide Ice Sheet occurred during stadial periods.

Description: The modern state of the Atlantic meridional overturning circulation promotes a northerly maximum of tropical rainfall associated with the Intertropical Convergence Zone (ITCZ). For continental regions, abrupt millennial-scale meridional shifts of this rainbelt are well documented, but the behavior of its oceanic counterpart is unclear due the lack of a robust proxy and high temporal resolution records. Here we show that the Atlantic ITCZ leaves a distinct signature in planktonic foraminifera assemblages. We applied this proxy to investigate the history of the Atlantic ITCZ for the last 30,000 years based on two high temporal resolution records from the western Atlantic Ocean. Our reconstruction indicates that the shallowest mixed layer associated with the Atlantic ITCZ unambiguously shifted meridionally in response to changes in the strength of the Atlantic meridional overturning with a southward displacement during Heinrich Stadials 2-1 and the Younger Dryas. We conclude that the Atlantic ITCZ was located at ca. 1°S (ca. 5° to the south of its modern annual mean position) during Heinrich Stadial 1. This supports a previous hypothesis, which postulates a southern hemisphere position of the oceanic ITCZ during climatic states with substantially reduced or absent cross-equatorial oceanic meridional heat transport.

Description: The Sonne transit cruise SO226-3 DipFIP took place from March 4th (Wellington, New Zealand) to March 28th (Kaohsiung, Taiwan) in 2013. CTD data for 16 stations along the cruise track were recorded using the onboard a SEABIRD SBE 9 plus CTD down to depth of 800m. Obtained hydrographic data were binned to 1 m intervals with the available SBE software. Obvious outliers in the readings of the oxygen sensor close to the sea surface have been manually removed. Fluorospectrometer (bbe Moldaenke) pigment data measured for 24 depth intervals are available for 10 stations. Measurements were conducted in the shipboard laboratory on water samples from the CTD rosette. Data are averages of a least 30 readings per sample.