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Among Antarctic bryozoans, some species are able to develop calcitic bioconstructions promoting habitat complexity, but the processes leading to biomineral formation are mostly unknown. The present work investigated three Antarctic bryozoans, from morphological to skeletal features, including the organic matrix associated with the skeleton (SOM). Cellarinella nutti Rogick, 1956 and Reteporella frigida Waters, 1904 were collected in November 2018 from a shallow site (25 m) and Cellarinella njegovanae Rogick, 1956 from a deep site (110 m) at Terra Nova Bay (Ross Sea, Antarctica). Both Cellarinella species showed 5–6 “growth check lines” (gcl) on their laminae. The morphometrical characterization conducted on the growth bands (gb) and zooids, within the band across bands, revealed a variability in length with time (C. nutti: from 4099 µm for gb1 to 1449 µm for gb6; C. njegovanae: from 1974 µm for gb 3 to 7127 µm for gb2). Zooid length varied within gb, from the proximal to the distal part of the bands, but differences also occurred across bands. The shortest zooids (~625 µm) were found at the proximal part and the longest (~ 1190 µm) in the middle part of the gb in C. nutti, whereas in C. njegovanae the shortest zooids (~ 660 µm) were found in the distal part and the longest (~1190 µm) in the proximal part of the gb. Micro-CT analyses indicated the ratio of basal zooidal walls (RbwT gcl/gb) ranged from 3.0 to 4.9 in C. nutti and from 2.3 to 5.9 in C. njegovanae, whereas Reteporella frigida did not form any gcl on either side of the colony. Preliminary characterizations of the SOM for the three species evidenced a mixture of proteins and polysaccharides with properties similar to those of better-known biominerals, in terms of quantity and electrophoretic behavior. In addition, a “lectin fingerprint” has been established for the first time in bryozoans, displaying the presence of chitin or chitin-related saccharides. Understanding the complexity of the processes regulating skeleton formation is a key aspect in comprehending the adaptation of bioconstructional ecosystems and the survival of the associated biodiversity under the future ocean.

The finding of a partially preserved elk skeleton from the Bavarian Alps is reported. Remnants of an adult male were found, together with skeletal elements of juvenile moose calves, at the base of a talus cone in the pit cave Stiefelschacht, next to Lenggries (southern Germany). The adult's bones exhibited anthropogenic traces like cut marks and were radiocarbon-dated to the Late Iron Age. A projectile hole in the left shoulder blade and cut marks on the bones are indicative of hunting and meat usage. The elk remains were associated with several wild and domestic species such as ungulates and hare but were not, however, accompanied by archaeological artefacts. Other archaeological sites of the Late Iron Age are so far not known within a distance of less than 30 km to the Stiefelschacht. While the presence of elk during prehistoric times in the Alps has already been known before, the finds and the location are unique in that they are the first evidence of elk hunting during the Late Iron Age in the northern Alps.

Bivalve shells are accretionary biogenic carbonates that yield a record of the organisms’ life history, also with respect to the physiological response to the ambient environmental conditions. This is reflected by variations in the shells’ growth bands, their chemical composition, and morphological features, making them useful tools in paleobiology and paleoenvironmental reconstructions. Corbula gibba is an abundant bivalve species in the Adriatic Sea that is commonly used to resolve questions in age determinations, paleoenvironmental reconstructions and sequence stratigraphic studies [1,2]. Nevertheless, a detailed analysis of its response in growth bands and chemical signatures to environmental variations is currently missing. This study evaluates growth increments and chemical variations in Corbula gibba shells. The shells have been collected alive from the Western Adriatic Sea between Senigallia and Ancona to shed light on the reliability of the Corbula gibba as an archive for (paleo)environmental information. Mutvei's solution has been used [3] to stain the alternating dark and light bundles in the shell, enhancing the visualization of growth increments. High-resolution elemental profiles have been acquired across the outer shells layer using the non-destructive Micro X-ray fluorescence (µ-XRF) technique [4]. The results are compared to available climatic information for Ancona and indicate that Corbula gibba shells record detailed temporal changes in the environment such as tidal fluctuations caused by lunar cycle.[1] Tomašových, A., Gallmetzer, I., Haselmair, A., Kaufman, D., Vidović, J., & Zuschin, M. (2017). Stratigraphic unmixing reveals repeated hypoxia events over the past 500 yr in the northern Adriatic Sea. Geology, 45 (4), 363-366.[2] Tomašových, A., Gallmetzer, I., Haselmair, A., Kaufman, D., Kralj, M., & Cassin, D. et al. (2018). Tracing the effects of eutrophication on molluscan communities in sediment cores: outbreaks of an opportunistic species coincide with reduced bioturbation and high frequency of hypoxia in the Adriatic Sea. Paleobiology, 44 (4), 575-602.[3] Schöne, B., Dunca, E., Fiebig, J., & Pfeiffer, M. (2005). Mutvei's solution: An ideal agent for resolving microgrowth structures of biogenic carbonates. Palaeogeography, Palaeoclimatology, Palaeoecology, 228 (1-2), 149-166.[4] de Winter, N.J., Sinnesael, M., Makarona, C., Vansteenberge, S., Claeys, P., (2017). Trace element analyses of carbonates using portable and micro-X-ray fluorescence: performance and optimization of measurement parameters and strategies. Journal of Analytical Atomic Spectrometry, 32, 1211–1223.

Crustose coralline algae (CCA) occur from the tropics to the poles in photic benthic environments. Here, we report on some of the world’s southernmost and coldest CCA sites in Terra Nova, Ross Sea, Antarctica at 74°41′ S. The recently described red alga Tethysphytum antarticum is investigated for its skeletal architecture, its mineralogical and geochemical composition, as well as for its taxonomic classification. A phylogenetic analysis based on molecular genetics and the sequencing of the photosystem II protein D1 (psbA) gave a perfect match with T. antarcticum. Histological sections and micro-CT-scans provide new diagnostic details for the conceptacles (the reproductive organs of the alga). X-ray diffractometry and electron-microprobe measurements yielded a clear high-Mg calcite (~8 mol%) composition of the skeletal parts. Detailed back-scattered electron imaging of polished petrographic thin sections revealed a two-layered thallus (vegetative plant tissue), comprising an organic-rich irregularly calcified basal layer with rectangular cells, overlain by the main thallus. Elemental maps show relatively increased sulphur in the basal layer, clearly tied to organic cell walls. MgCO3 and SrCO3 were targeted with semiquantitative elemental mappings and in an ontogenetic quantitative spot transect. Compared with temperature (−1.95 °C to +1.08 °C), the MgCO3 (mol%) reflects this world’s coldest CCA site temperature with the lowest MgCO3 content of 7.9 ± 1.6 mol%. The along transect variability, however, shows with ~6 mol% a larger MgCO3 variability than expected for the 3 °C intra-annual temperature amplitude in Terra Nova Bay. This implies that in low amplitude settings the biomineralisation control on Mg/Ca ratios can outcompete its temperature sensitivity. Mark-recapture studies, next to the environmental logger station La Zecca are suggested, to perform a detailed growth rate and biomineralisation quantification.

The order Heterocorallia is an enigmatic, anatomically distinct group of calcifying Palaeozoic corals with unclear affinities to other, either extinct or modern cnidarian lineages. Here, we present results of microstructure and stable isotope studies on exceptionally well-preserved skeletons of two species of the heterocoral Oligophylloides from the Famennian (Upper Devonian) of the Anti-Atlas, Morocco and the Holy Cross Mountains, Poland. The investigations provide new information on the skeletogenesis and isotope fractionation effects in Palaeozoic heterocorals. In order to estimate the possible role of diagenetic alteration on the observed carbon and oxygen isotope ratios, we have conducted several preservation tests, including transmitted-light, scanning-electron, fluorescence and cathodoluminescence microscopy, Raman spectroscopy, and trace element analyses. The leastand most-altered skeleton portions identified based on this approach are typified by clearly different isotopic signatures. The least-altered parts of the specimens reveal carbon and oxygen isotope ratios close to those of contemporaneous marine-equilibrated calcites. The moderately-pronounced, positive correlations between the 813C and 818O values observed for these coralla may reflect relatively minor vital fractionation effects, obscured to some degree by diagenetic overprint. The results suggest that the isotope fractionation mechanisms of the heterocorals differed markedly from the significant kinetic fractionation characteristic of the modern scleractinian corals. Rather, the moderate, if any, extent of non-equilibrium fractionation during skeletogenesis in the Heterocorallia resembled the typical, environmental and metabolic CO2-dominated fractionation effects shown by most other marine calcifiers. These include the extinct rugose corals, modern hydrocorals and, in particular, modern octocorals, with which the heterocorals show notable architectural and structural similarities.

Determining secular changes in seawater temperature and Mg/Ca ratio is of high importance in paleooceanographic, paleoenvironmental and paleoecological studies. On the one hand, intra-shell variation in Mg/Ca in brachiopod shells is used to reconstruct historical changes in temperature and in seasonality. On the other hand, intra-shell analyses show a weak relationship between Mg/Ca and delta O-18, but a positive correlation between Mg/Ca and Sr/Ca, and an increase of Mg/Ca at growth lines, indicating that Mg/Ca does not simply track seawater temperature alone. Internal growth lines are rarely visible in brachiopod shells, and the relation between growth lines that mark growth slowdown or cessation and Mg/Ca fluctuation is thus not properly understood. Here, using four species of extant brachiopods from the NE Pacific and the SW Pacific, from environments with low seasonality in seawater temperature (

The present dataset was collected to evaluate the environmental stressors on a lacustrine basin in the Eastern Alps of glacial origin that has been affected in recent years by natural and anthropogenic events such as the construction of a hydroelectric power plant and a series of strong earthquakes during 1976-1977. We collected sediment cores in different sites from the lake margins to the depocenter and performed a multiproxy analysis of sediment sample to highlight lake stratigraphy and major changes occurring at a decadal scale (Polonia et al., [1]). The integrated analyses of sedimentological, geochemical, isotopic, mineralogical and micropaleontological analyses aimed at reconstructing changes in sediment composition and define the triggering mechanisms of altered environmental conditions. The dataset demonstrates that evaluating ex post the effects of artificial modification in a natural environment during relatively long time spans (decades) can provide important insights for managing and protection strategies in similar environments worldwide (C) 2021 The Authors. Published by Elsevier Inc.

The Lake of Cavazzo is a natural, lacustrine basin in the Friuli-Venezia Giulia region (NE Italy), which occupies a fluvio-glacial suspended valley of the Tagliamento river, at the southern front of the Alpine chain. The lake formed after the melting of major glacial tongues at the end of the Last Glacial Maximum, and has been affected in recent years by natural and anthropogenic events, with economic, cultural and environmental consequences. According to local witnesses, the lake environment has changed dramatically after the 1950s, when the basin was connected to a hydroelectric power plant collecting waters from a wider catchment area, as it served as its final discharge basin. In this work, based on a densely-spaced grid of high-resolution seismic reflection profiles complemented by sediment cores, we analysed the uppermost lake stratigraphy, highlighting major changes occurring at a decadal scale. Our main purpose is to verify whether and how the lake sediments record the transition from pristine/natural to artificial conditions, as well as the effects of multiple natural impacts including the 1976–77 Friuli earthquake sequence. The results of our analysis suggest varying environmental conditions of the lake after the 1950s indicated by changing sediment mineralogy, increased deposition of allochthonous clastic sediments, and recurrent episodes of anoxic conditions at the lake's floor, likely triggered by hyperpycnal flows derived from the hydroelectric power plant discharges. We also observe that the effects of the 1976–77 seismic sequence are recorded in the lake as resedimented levels, likely due to shaking, in situ deformation, landslides, and turbidity currents. Our results stress the importance of lacustrine environments as efficient recorders of anthropogenic and natural events.

The scleractinian cold-water corals (CWCs), including the species Madrepora oculata and especially Lophelia pertusa, have been studied extensively in an attempt to decipher environmental signals recorded during biomineralisation in order to extract environmental chronologies. However, understanding the mechanisms of carbonate precipitation is a prerequisite to interpret variations in geochemical signatures locked into the skeleton during coral growth; to date results are still inconclusive. Here a novel approach, comparing the calcification patterns within the coral microstructure of species L. pertusa and M. oculata and the geochemistry along the contact surfaces with calcified polychaete tubes is undertaken to provide additional information on the mechanisms of biomineralisation in colonial corals. The fact that no significant difference in microstructures, variations in growth rate, or geochemical composition between the corallite theca and the calcified polychaete tube was detectable leads to the conclusion that both have been deposited by the coral tissue in L. pertusa and M. oculata. Based on prior knowledge on the symbiotic relationship between CWCs and the polychaete Eunice norvegica, an involvement of mucus in the calcification of the parchment tubes had been suspected. However, we found only evidence for aragonite precipitated by coral tissue, without evidence for an involvement of mucus in the calcification.

Stable isotope compositions of oxygen (delta O-18) and carbon (delta C-1(3)) of the aragonite skeleton of the reef-building gastropod Dendropoma petreaum provide high-resolution records of the Mediterranean climate over the last millennium. In particular, the isotopic composition of vermetid cores collected from the west and east Mediterranean reveals that the different regions have had distinct thermal and primary production behaviors throughout the last millennium. The rate of warming in the recent Industrial Period is variable among the different regions of the Mediterranean Sea. The delta O-18-derived Sea Surface Temperature (SST) anomalies show that the Eastern Mediterranean surface temperature is increasing much more rapidly than in the Western Mediterranean. Additionally, the signals of the Little Ice Age and of the Medieval Climate Anomaly are more apparent in the western and in the central Mediterranean while they are almost absent in the eastern Mediterranean. We aim to reconcile the SST temporal and spatial pattern with the variability of the North Atlantic Oscillation (NAO) and the South Asian Monsoon (SAM) climate systems by analyzing coupled atmosphere-ocean models from the CMIP5/PMIP3 projects that simulate the global climate of the past 1000 years. We show that even though the NAO is more dominant in the western Mediterranean SST, its latitudinal movement, on a centennial time scale, is evident in the eastern Mediterranean SST signal. We also discuss the Atlantic water inflow role in the observed similarities of the surface productivity signals and the Suess effect that prevail the Industrial Period signal in all regions of the Mediterranean.

Ravine slopes at the recently discovered Nesseltalgraben site in southeastern Germany provide a unique last glacial sediment record for the Northern Calcareous Alps. The 21 m-long profile is dominated by fine-grained lacustrine-palustrine sediments overlain by several metres of glacifiuvial gravels and lodgement tills of the Last Glacial Maximum and underlain by a diamicton. The age model includes 29 radiocarbon analyses and one paleomagnetic anomaly (Laschamp event) together providing a modelled age range from c. 59 to 29.6 ka cal BP, i.e. Marine Isotope Stage (MIS) 3. Apart from a description of the lithofacies, X-ray-fluorescence (XRF) scanning and elemental analyses provide high-resolution records of the geochemical composition. Multivariate analyses of XRF data separate Ca from other major elements. Carbonate contents, represented by Ca and total inorganic carbon, reach maxima in repeatedly occurring calcareous silty to sandy layers and are related to glacigenic origin. These indicate repeated inner-alpine local glaciations during stadials of MIS 3. Scanning electron microscopy and XRF data confirm the detrital origin of these layers. In contrast, organic matter and elements more resistant to chemical weathering (Si, Ti, Zr) accumulated during interstadials and concurrently elevated Rb/Sr ratios indicate intensified weathering. The high-frequency proxy variations determined for Nesseltalgraben reflect interstadialstadial climate variability comparable with oxygen-isotope records from Greenland ice cores and Alpine speleothems. Thus, Nesseltalgraben is among the very few independently dated sediment records from continental Europe covering the entire MIS 3 and reflecting the full Dansgaard-Oeschger climate variability. (C) 2019 Elsevier Ltd. All rights reserved.

Cold-water coral ecosystems present common carbonate factories along the Atlantic continental margins, where they can form large reef structures. There is increasing knowledge on their ecology, molecular genetics, environmental controls and threats available. However, information on their carbo-nate production and accumulation is still very limited, even though this information is essential for their evaluation as carbonate sinks. The aim of this study is to provide high-resolution reef aggradation and carbonate accumulation rates for Norwegian cold-water coral reefs from various settings (sunds, inner shelf and shelf margin). Furthermore, it introduces a new approach for the evaluation of the cold-water coral preservation within cold-water coral deposits by computed tomography analysis. This approach allows the differentiation of various kinds of cold-water coral deposits by their macrofossil clast size and orientation signature. The obtained results suggest that preservation of cold-water coral frameworks in living position is favoured by high reef aggradation rates, while preservation of coral rubble prevails by moderate aggradation rates. A high degree of macrofossil fragmentation indicates condensed intervals or unconformities. The observed aggradation rates with up to 1500 cm kyr−1 exhibit the highest rates from cold-water coral reefs so far. Reef aggradation within the studied cores was restricted to the Early and Late Holocene. Available datings of Norwegian cold-water corals support this age pattern for other fjords while, on the shelf, cold-water coral ages are reported additionally from the early Middle Holocene. The obtained mean carbonate accumulation rates of up to 103 g cm−2 kyr−1 exceed previous estimates of cold-water coral reefs by a factor of two to three and by almost one order of magnitude to adjacent sedimentary environments (shelf, slope and deep sea). Only fjord basins locally exhibit carbonate accumulation rates in the range of the cold-water coral reefs. Furthermore, cold-water coral reef carbonate accumulation rates are in the range of tropical reef carbonate accumulation rates. These results clearly suggest the importance of cold-water coral reefs as local, maybe regional to global, carbonate sinks

Skeletons of cold-water corals are potential geochemical archives that record oceanographic parameters, such as bottom water temperature in the deep sea. In this study, the three solitary scleractinian species, Caryophyllia cyathus, Desmophyllum cristagalli and Stenocyathus vermiformis, were sampled from upper bathyal 500 m water depths in the southern Faial Channel (Azores archipelago). The stable carbon isotope (δ13C) and oxygen isotope (δ18O) compositions of the corals were analysed and related to their skeletal architecture. Background data, including data from high-resolution temperature logs and on the ambient seawater isotopic composition, were compared with δ18O-based temperature reconstructions to test the suitability of these cold-water corals as geochemical archives and to explore the role of vital effects on skeletal geochemistry. Although all of the corals experienced identical and near-stable salinities and temperatures, the stable isotopic signatures vary within and among species. Both δ13C and δ18O values fluctuate in relation to growth bandings and are therefore most likely dependent on calcification rates and associated kinetic vital effects. The studied specimens are 13C depleted with respect to equilibrium with ambient seawater, indicating significant uptake of respired CO2 by their skeletons. Skeletal δ18O values vary by up to 3.6‰ among the growth bands of one of the coral specimens, translating into an unrealistic reconstructed temperature amplitude of ~ 16 °C. This contrasts with the minute intra-annual amplitude of the stable temperature regime (annual mean 12.30 ± 0.25 °C). Nevertheless, δ13C vs. δ18O regression lines can be used to deduce ambient temperatures by applying the lines technique (Smith et al., 2000; Palaios 15, 25–32), which yields temperatures that coincide with the measured maximum temperature range in the case of D. cristagalli and for most C. cyathus samples, but deviate by 2.95 °C from the measured annual mean in the case of the S. vermiformis specimen.

A high-resolution record, covering 9.3–0.2 ka BP, from the sub-arctic Stjernsund (70°N) was studied for benthic foraminiferal faunas and stable isotopes, revealing three informally named main phases during the Holocene. The Early- to Mid-Holocene (9.3–5.0 ka BP) was characterized by the strong influence of the North Atlantic Current (NAC), which prevented the reflection of the Holocene Climatic Optimum (HCO) in the bottom-water temperature. During the Mid-Holocene Transition (5.0–2.5 ka BP), a turnover of benthic foraminiferal faunas occurred, Atlantic Water species decreased while Arctic-Polar species increased, and the oxygen isotope record showed larger fluctuations. Those variations correspond to a period of global climate change, to spatially more heterogeneous benthic foraminiferal faunas in the Nordic Seas region, and to regionally diverging terrestrial temperatures. The Cool Late Holocene (2.5–0.2 ka BP) was characterized by increased abundances of Arctic-Polar species and a steady cooling trend reflected in the oxygen isotopes. In this period, our record differs considerably from those on the SW Barents Sea shelf and locations farther south. Therefore, we argue that regional atmospheric cooling triggered the late Holocene cooling trend. Several cold episodes centred at ∼8.3, ∼7.8, ∼6.5, ∼4.9, ∼3.9 and ∼3.3 ka BP were identified from the benthic foraminiferal faunas and the δ18O record, which correlated with marine and atmospherically driven proxy records. This suggests that short-term cold events may result from reduced heat advection via the NAC or from colder air temperatures.

International audience; U-series age patterns obtained on reef framework-forming cold-water corals collected over a nearly 6000-km-long continental margin sector, extending from off Mauritania (17 degrees N; northwest Africa) to the southwestern Barents Sea (70 degrees N; northeastern Europe), reveal strong climate influences on the geographical distribution and sustained development of these ecosystems. Over the past three glacial-interglacial cycles, framework-forming cold-water corals (Lophelia pertusa and Madrepora oculata) seem to have predominantly populated reefs, canyons, and patches in the temperate East Atlantic and the Mediterranean Sea. Above 50 degrees N corals colonize reefs in the northern East Atlantic primarily during warm climate periods with the biogeographic limit advancing from similar to 50 degrees N to similar to 70 degrees N. We propose that north-south oscillations of the biogeographic limit of reef developments are paced by ice ages and may occur synchronously with north-south displacement of cold nutrient-rich intermediate waters and surface productivity related to changes of the polar front.

Uranium-series and radiocarbon ages are reported for deep-sea corals Madrepora oculata , Desmophyllum dianthus , Lophelia pertusa and Caryophyllia smithii from the Mediterranean Sea. U-series dating indicates that deep-sea corals have persisted in the Mediterranean for over 480,000 years, especially during cool interstadial periods. The most prolific period of growth however appears to have occurred within the Younger Dryas (YD) period from 12,900 to 11,700 years BP followed by a short (~ 330 years) phase of post-YD coral growth from 11,230 to 10,900 years BP. This indicates that deep-sea corals were prolific in the Mediterranean not only during the return to the more glacial-like conditions of the YD, but also following the rapid deglaciation and transition to warmer conditions that followed the end of the YD. Surprisingly, there is a paucity Last Glacial Maximum (LGM) coral ages, implying they were largely absent during this period when cold-water conditions were more prevalent. Radiocarbon ages show that the intermediate depth waters of the Mediterranean generally had Δ 14 C compositions similar to surface waters, indicating that these waters were extremely well ventilated. The only exception is a narrow period in the YD (12,500 ± 100 years BP) when several samples of Lophelia pertusa from the Ionian Sea had Δ 14 C values falling significantly below the marine curve. Using a refined approach, isolation ages (τ isol ) of 300 years to 500 years are estimated for these intermediate (800–1000 m) depth waters relative to surface marine waters, indicating a reduction or absence of deep-water formation in the Ionian and adjacent Adriatic Seas during the YD. Contrary to previous findings, we find no evidence for widespread intrusion of low Δ 14 C Atlantic waters into the Mediterranean. Prolific growth of deep-sea corals in the Mediterranean ended abruptly at ~ 10,900 years BP, with many of the coral-bearing mounds on the continental slopes being draped in a thin veneer of mud. Their demise is attributed to a number of factors, including the direct loss of habitat due to high sedimentation that accompanied glacial meltwater pulses, together with rising temperatures that would have finally pervaded the deeper water of the Mediterranean following the onset of Holocene warming.

The pure white fans of the stylasterid Errina dabneyi are a conspicuous feature on the upper bathyal slopes in Azorean waters and were documented and recovered alive with the aid of a submersible in the southern Faial Channel. Etched vacuum-epoxy-casts of the aragonite skeleton, studied by scanning electron microscopy, reveal the 3-dimensional internal architecture comprising coenosarc canal meshwork, dactylopores, gastropores and ampullae. Near the surface, the canals are narrow and interconnected in a regular 3-dimensional meshwork. Deeper inside the canals are less abundant, more irregular and wider. This pattern implies that the skeletal architecture is modified during growth, with more central canals being enlarged by dissolution, and other canals, gastropores and dactylopores in turn being filled with aragonite reprecipitates. The skeleton is primarily com- posed of irregular spherulitic aggregates and overprinting during growth is evident from ghost struc- tures in the form of successive semi-spherulitic infill of former canals. Due to differential dissolution and reprecipitation, this internal rebuild process inevitably involves an alteration of initial geochem- ical signatures such as stable isotope ratios (δ 18 O and δ 13 C), trace element signals and the distribution of radiogenic isotopes of carbon and uranium. This has to be taken into account when applying radio- metric dating techniques and when using stylasterids as a geochemical archive.

A conspicuous new deep-sea oyster, Neopycnodonte zibrowii sp. n., is described from the Azores Archipelago, where it thrives in 420 to >500 m water depth in high densities concealed underneath overhangs. The new species reaches a relatively large size, which may exceed 20 cm, and is characterised by a very unusual hinge line morphology, straight without a bulge of the resilium. It is compared to the extant Indo-Pacific Empressostrea kostini Huber and Lorenz, 2007 and to the cosmopolitan Neopycnodonte cochlear (Poli, 1791), which has a broadly sympatric distribution at shallower depths in the Azores and Bay of Biscay. Radiocarbon dating reveals that individuals reach an impressive lifespan of one to more than five centuries, placing them among the longest-lived molluscs known to date. They often grow on top of each other, forming stacks that resemble dish piles—an effective measure to optimise shell stability with minimal biomineralisation effort, but with the drawback of increased bioerosion ultimately leading to detachment. Three microstructure types are developed in N. zibrowii : (1) the cross-foliated, calcitic, dorsal to central endostracum and aragonitic ligostracum, (2) the porous vesicular structure of the calcitic ventral endostracum, and (3) the simple prismatic aragonitic myostracum. Foliated and vesicular shell portions show sub-millimetre-scale first-order increments delineated by conchiolin-rich growth breaks (interpreted as reproductive cyclicity), and less distinct second-order increments (interpreted as annual in nature). This pattern is clearly reflected by the elemental composition with the primary growth breaks lacking Ca and Sr but including Mg and S as organic matrix constituents. The second-order increments within the calcite are mirrored by moderately co-varying Mg/Ca and S/Ca fluctuations at stable Sr concentrations, reflecting varying proportions of organic matrix. Dorsal and central endostracum transects reveal a low inter-valve, but considerable inter-specimen variability with high Mg/Ca molar ratios and fluctuations (22.5±17.6 mmol/mol), low Sr/Ca values (0.2±0.1 mmol/mol), and a typical to high S/Ca content (6.9±2.2 mmol/mol), when compared to other calcitic bivalves. Unlike short-lived, shallow-water oysters, N. zibrowii thrives under very stable environmental conditions. Minimal temperature fluctuations and stable open marine salinity provide an optimal basis to recognise biological fractionation processes. Strong Mg/Ca fluctuations indicate a physiological control related to metabolism and biomineralisation, prohibiting the use of this ratio as a temperature or ocean chemistry proxy. Low Sr/Ca ratios indicate rather constant and low long-term accretion rates, while short-scale fluctuations may be attributed to short-term variations in growth rate and Mg incorporation. Oxygen isotopes yield a considerable spread of 1.8‰ with a mean of 2.0±0.3‰ δ 18 O V-PDB, and low correlation between different contemporaneous parts of the shell and between specimens. These values surprisingly exceed expected equilibrium conditions, calculated from in situ temperature data (annual mean 12.3±0.3 °C) and seawater isotopic composition (0.5±0.1‰ δ 18 O SMOW), by 0.5‰ on average. Such positively shifting vital effects, previously reported for limpets and barnacles, are often overlooked in high-temperature and high-amplitude settings and may be more common than is currently believed. Carbon isotopes range from 0.2 to 3.5‰ δ 13 C V-PDB (mean 1.8±0.7‰) and show an ontogenetic decrease, but may incorporate an environmental signal in adult portions, indicated by a strong correspondence of peaks between specimens. This signal is likely driven by a complex interplay of different factors, such as primary production, current-based food supply and metabolism.

This study investigates stable isotope signatures of five species of Silurian and Devonian deep-water, ahermatypic rugose corals, providing new insights into isotopic fractionation effects exhibited by Palaeozoic rugosans, and possible role of diagenetic processes in modifying their original isotopic signals. To minimize the influence of intraskeletal cements on the observed signatures, the analysed specimens included unusual species either devoid of large intraskeletal open spaces ('button corals': Microcyclus, Palaeocyclus), or typified by particularly thick corallite walls (Calceola). The corals were collected at four localities in the Holy Cross Mountains (Poland), Mader Basin (Morocco) and on Gotland (Sweden), representing distinct diagenetic histories and different styles of diagenetic alteration. To evaluate the resistance of the corallites to diagenesis, we applied various microscopic and trace element preservation tests. Distinct differences between isotopic compositions of the least-altered and most-altered skeleton portions emphasise a critical role of material selection for geochemical studies of Palaeozoic corals. The least-altered parts of the specimens show marine or near-marine stable isotope signals and lack positive correlation between δ13C and δ18O. In terms of isotopic fractionation mechanisms, Palaeozoic rugosans must have differed considerably from modern deep-water scleractinians, typified by significant depletion in both 18O and 13C, and pronounced δ13C-δ18O co-variance. The fractionation effects exhibited by rugosans seem similar rather to the minor isotopic effects typical of modern non-scleractinian corals (octocorals and hydrocorals). The results of the present study add to growing evidence for significant differences between Scleractinia and Rugosa, and agree with recent studies indicating that calcification mechanisms developed independently in these two groups of cnidarians. Consequently, particular caution is needed in using scleractinians as analogues in isotopic studies of extinct coral lineages. Answering some of the pertinent palaeoecological questions, such as that of the possibility of photosymbiosis in Palaeozoic corals, may not be possible based on stable isotope data.

In order to assess the effects of ocean acidification and warming on the Mediterranean mussel (Mytilus galloprovincialis), specimens were reared in aquarium tanks and exposed to elevated conditions of temperature (+3°C) and acidity (−0.3 pH units) for a period of 10 months. The whole system comprised a factorial experimental design with 4 treatments (3 aquaria per treatment): control, lowered pH, elevated temperature, and lowered pH/elevated temperature. Mortality was estimated on a weekly basis and every 2 months, various biometrical parameters and physiological processes were measured: somatic and shell growth, metabolic rates and body fluid acid-base parameters. Mussels were highly sensitive to warming, with 100% mortality observed under elevated temperature at the end of our experiment in October. Mortality rates increased drastically in summer, when water temperature exceeded 25°C. In contrast, our results suggest that survival of this species will not be affected by a pH decrease of ~0.3 in the Mediterranean Sea. Somatic and shell growth did not appear very sensitive to ocean acidification and warming during most of the experiment, but were reduced, after summer, in the lowered pH treatment. This was consistent with measured shell net dissolution and observed loss of periostracum, as well as uncompensated extracellular acidosis in the lowered pH treatment indicating a progressive insufficiency in acid-base regulation capacity. However, based on the present dataset, we cannot elucidate if these decreases in growth and regulation capacities after summer are a consequence of lower pH levels during that period or a consequence of a combined effect of acidification and warming. To summarize, while ocean acidification will potentially contribute to lower growth rates, especially in summer when mussels are exposed to sub-optimal conditions, ocean warming will likely pose more serious threats to Mediterranean mussels in this region in the coming decades, This work was funded by the EC FP7 project “Mediterranean Sea Acidification in a changing climate” (MedSeA; grant agreement 265103), the French program PNEC (Programme national environnement côtier; Institut national des sciences de l'univers) and the EC FP7 project “European Project on Ocean Acidification” (EPOCA; grant agreement 211384)

We show that the Li/Mg systematics of a large suite of aragonitic coral skeletons, representing a wide range of species inhabiting disparate environments, provides a robust proxy for ambient seawater temperature. The corals encompass both zooxanthellate and azooxanthellate species (Acropora sp., Porites sp., Cladocora caespitosa, Lophelia pertusa, Madrepora oculata and Flabellum impensum) collected from shallow, intermediate, and deep-water habitats, as well as specimens cultured in tanks under temperature-controlled conditions. The Li/Mg ratios observed in corals from these diverse tropical, temperate, and deep-water environments are shown to be highly correlated with temperature, giving an exponential temperature relationship of: Li/Mg (mmol/mol) = 5.41 exp (−0.049 * T) (r² = 0.975, n = 49). Based on the standard error of the Li/Mg versus temperature correlation, we obtain a typical precision of ±0.9 °C for the wide range of species analysed, similar or better than that of other less robust coral temperature proxies such as Sr/Ca ratios.The robustness and species independent character of the Li/Mg temperature proxy is shown to be the result of the normalization of Li to Mg, effectively eliminating the precipitation efficiency component such that temperature remains as the main controller of coral Li/Mg compositions. This is inferred from analysis of corresponding Li/Ca and Mg/Ca ratios with both ratios showing strong microstructure-related co-variations between the fibrous aragonite and centres of calcification, a characteristic that we attribute to varying physiological controls on growth rate. Furthermore, Li/Ca ratios show an offset between more rapidly growing zooxanthellate and azooxanthellate corals, and hence only an approximately inverse relationship to seawater temperature. Mg/Ca ratios show very strong physiological controls on growth rate but no significant dependence with temperature, except possibly for Acropora sp. and Porites sp. A strong positive correlation is nevertheless found between Li/Ca and Mg/Ca ratios at similar temperatures, indicating that both Li and Mg are subject to control by similar growth mechanisms, specifically the mass fraction of aragonite precipitated during calcification, which is shown to be consistent with a Rayleigh-based elemental fractionation model.The highly coherent array defined by Li/Mg versus temperature is thus largely independent of coral calcification mechanisms, with the strong temperature dependence reflecting the greater sensitivity of the KdLi/Ca partition coefficient relative to KdMg/Ca. Accordingly, Li/Mg ratios exhibit a highly coherent exponential correlation with temperature, thereby providing a more robust tool for reconstructing paleo-seawater temperatures.

The boron isotope systematics has been determined for azooxanthellate scleractinian corals from a wide range of both deep-sea and shallow-water environments. The aragonitic coral species, Caryophyllia smithii, Desmophyllum dianthus, Enallopsammia rostrata, Lophelia pertusa, and Madrepora oculata, are all found to have relatively high d11B compositions ranging from 23.2‰ to 28.7‰. These values lie substantially above the pH-dependent inorganic seawater borate equilibrium curve, indicative of strong up-regulation of pH of the internal calcifying fluid (pHcf), being elevated by ~0.6–0.8 units (?pH) relative to ambient seawater. In contrast, the deep-sea calcitic coral Corallium sp. has a significantly lower d11B composition of 15.5‰, with a corresponding lower ?pH value of ~0.3 units, reflecting the importance of mineralogical control on biological pH up-regulation. The solitary coral D. dianthus was sampled over a wide range of seawater pHT and shows an approximate linear correlation with ?pHDesmo = 6.43 - 0.71pHT (r2 = 0.79). An improved correlation is however found with the closely related parameter of seawater aragonite saturation state, where ?pHDesmo = 1.09 - 0.14Oarag (r2 = 0.95), indicating the important control that carbonate saturation state has on calcification. The ability to up-regulate internal pHcf, and consequently Ocf, of the calcifying fluid is therefore a process present in both azooxanthellate and zooxanthellate aragonitic corals, and is attributed to the action of Ca2+-ATPase in modulating the proton gradient between seawater and the site of calcification. These findings also show that the boron isotopic compositions (d11Bcarb) of aragonitic corals are highly systematic and consistent with direct uptake of the borate species within the biologically controlled extracellular calcifying medium. We also show that the relatively strong up-regulation of pH and consequent elevation of the internal carbonate saturation state (Ocf ~8.5 to ~13) at the site of calcification by cold-water corals, facilitates calcification at or in some cases below the aragonite saturation horizon, providing a greater ability to adapt to the already low and now decreasing carbonate ion concentrations. Although providing greater resilience to the effects of ocean acidification and enhancing rates of calcification with increasing temperature, the process of internal pHcf up-regulation has an associated energetic cost, and therefore growth-rate cost, of ~10% per 0.1 pH unit decrease in seawater pHT. Furthermore, as the aragonite saturation horizon shoals with rapidly increasing pCO2 and Oarag < 1, increased dissolution of the exposed skeleton will ultimately limit their survival in the deep oceans.

Special issue The APLABES Programme: Physical, Chemical and Biological Characterization of Deep-Water Coral Ecosystems from the Ionian Sea (Mediterranean).-- 16 pages, 13 figures, 3 tables, 2 appendix, The aragonitic skeletons of bathyal cold-water corals have a high potential as geochemical in situ archives for paleoceanography. Oxygen isotopes and stable carbon isotopes (δ18O and δ13C) have been analyzed, as well as trace and minor element compositions (e.g. Mg/Ca, Sr/Ca, U/Ca, B/Ca and P/Ca) in Lophelia pertusa, one of the most important frame-builders at the Santa Maria di Leuca (SML) deep-water coral hotspot in the Central Mediterranean. The Apulian Bank is swept by strong currents of the Adriatic Deep Water Outflow. The temperature of 13.9 °C is the highest temperature recorded for L. pertusa and provides an important end-member of environmental conditions for geochemical analyses on living Atlantic and Mediterranean cold-water corals. Temperature and salinity (38.77 PSU) are stable throughout the year, and thus virtually no changes should be observed in the stable oxygen isotope signal-if the coral precipitates its skeleton in equilibrium with seawater. We measured various marine properties, such as the seawater oxygen isotope composition (δ18Osw), stable carbon isotope composition (δ13CDIC) of dissolved inorganic carbon (DIC), and dissolved inorganic nutrient concentrations (PO4, NO3, NO2, NH3 and SiO2). Bottom water at the coral sites shows a mean oxygen isotope composition of 1.47‰ δ18Osw-VSMOW, and δ13CDIC showed a mean of 1.1‰ VPDB. A section of a living L. pertusa with a thick theca calcification was probed with a Merchantek MicroMill at a high spatial sampling resolution with 10 samples per 1 mm. This reduced the signal-smoothing inherent to conventional sampling. The δ18Oag of coral aragonite ranges between -2.0‰ and +2.8‰ VPDB and the δ13Cag ranges between -7.77‰ and +1.47‰ VPDB. The Gaussian data distribution for both parameters, including heavy equilibrium values, suggests the completeness of the captured isotopic variability. The strict linear correlation of δ13C and δ18O displays a strong 'kinetic' vital effect. The intercept of the δ13C/δ18O correlation line with the δ13CDIC-composition permits recognition of δ18O equilibrium values of aragonite and thus reconstruction of water temperatures despite strong disequilibrium precipitation. Since the environmental parameters (T, S and δ18Osw) are stable, the entire isotopic signal of the coral must be driven by biological fractionation and might reflect growth speed variations, potentially related to pH variations and changes in the saturation state of the calcifying fluid or seasonally varying nutrient availability. Laser ablation tracks show a trace element composition dependent to microstructural zones (fibrous aragonite vs. centres of calcification). The parabolic relation of the classical temperature proxies Mg/Ca and U/Ca point to trace element vital effects, rendering them unreliable in L. pertusa. The P/Ca ratio shows similar values as Desmophyllum dianthus, for which a linear dependence with seawater phosphate (DIP) has been previously demonstrated. Consequently L. pertusa might be an additional nutrient recorder at bathyal depths. From the same site we also analysed the stable isotopic composition of the Last Glacial pectinid bivalve Pseudamussium peslutrae, which has been radiocarbon-dated (AMS-14C) at 26.3 ka 14C yr BP. The isotope values of the shell calcite document a strongly differing glacial temperature-salinity regime preceding the Holocene coral growth above a prominent hiatus, This work has been carried out within the FIRB APLABES-project, the ESF Euromargins MOUNDFORCE-project, the EU HERMES-and Hermione-projects (EC contract no. GOCE-CT-2005-511234) and the FISR MICENA-project. We would like to thank Prof. Dr. A. Freiwald (GZN, University of Erlangen-Nuremberg) for further financial and logistical support under the 6th Framework Program of the European Commission

Terra Nova, 22, 494–500, 2010 Abstract New detailed swath bathymetry and bottom samples from Coral Patch and Ormonde seamounts provide constraints on the emplacement of the Madeira hotspot in the Eastern Atlantic Ocean. Swath bathymetric data document that Coral Patch is a composite structure, made up of at least nine distinct volcanic centres. Lithified pelagic carbonates infilling fissures in lava blocks constrain a minimal age for the volcanism in the Early Miocene and represent the first documentation of Coral Patch acting as an offshore terrigenous-starved seamount. At Coral Patch, as already observed at the Ormonde seamount, volcanism was emplaced on top of a pre-existing relief resulting from the regional tectonic compressive regime.

This paper presents the first compilation of information on the spatial distribution of scleractinian cold-water corals in the Gulf of Cádiz based on literature research and own observations (video footage, sediment samples). Scleractinian cold-water corals are widely distributed along the Spanish and Moroccan margins in the Gulf of Cádiz, where they are mainly associated with mud volcanoes, diapiric ridges, steep fault escarpments, and coral mounds. Dendrophyllia cornigera, Dendrophyllia alternata, Eguchipsammia cornucopia, Madrepora oculata and Lophelia pertusa are the most abundant reef-forming species. Today, they are almost solely present as isolated patches of fossil coral and coral rubble. The absence of living scleractinian corals is likely related to a reduced food supply caused by low productivity and diminished tidal effects. In contrast, during the past 48 kyr scleractinian corals were abundant in the Gulf of Cádiz, although their occurrence demonstrates no relationship with main climatic or oceanographic changes. Nevertheless, there exists a conspicuous relationship when the main species are considered separately. Dendrophylliids are associated with periods of relatively stable and warm conditions. The occurrence of L. pertusa mainly clusters within the last glacial when bottom current strength in the Gulf of Cádiz was enhanced and long-term stable conditions existed in terms of temperature. Madrepora oculata shows a higher tolerance to abrupt environmental changes. © 2009 Elsevier Ltd. All rights reserved.

The parasitic foraminifer Hyrrokkin sarcophagapredominantly infests the cold-water coral Lophelia pertusaand the co-occurring bivalve Acesta excavata, showing a commensal or parasitic behaviour. It occurs also on some other corals (e.g., Caryophyllia sarsiae), bivalves (e.g., Delectopecten vitreus) and sponges (Geodiasp.), typically in aphotic environments. The aim of the study is to describe its traces from various host substrates, to characterise its parasitic behaviour and to map the geographical distribution of the genus Hyrrokkin. Epoxy-resin casts of H. sarcophagatraces in A. excavata C. sarsiae, D. vitreusand L. pertusa,and of H. carnivoratraces in A. excavata, were SEM analysed. The boring pattern is in all cases characterised by a shallow groove of up to 7 mm in diameter (max. 2 mm deep), from which ‘whip’-shaped extensions protrude vertically into the substrate. In A. excavatathe foraminifer can penetrate the entire valve to the mantle cavity, producing a thick shaft of fused ‘whips’. This parasitic attack is answered by a strong callus formation of the mollusc. One individual foraminifer can repeatedly bypass the organic-rich callus, resulting in a thick aragonite pinnacle. The trace surface texture is xenoglyph and changes with the penetrated host-microstructures. This is especially obvious on deeply penetrating trace portions (e.g., ‘whip’-shaped filaments) and is a strong indication for a chemical penetration mode (etching). The trace of Hyrrokkin is described as Kardopomorphos polydioryxigen. n., isp. n. On the substrates without the shaft, related to parasitic behaviour, Hyrrokkinmight feed directly on adjacent external host tissue. H. sarcophagais known along the North Atlantic continental margin from polar to subtropical latitudes and H. carnivoraoccurs on the continental margin of Mauritania, Congo and Guinea. In the Mediterranean we could document the parasitism of H. sarcophagafrom Last Glacial A. excavata.Traces or detached foraminifer tests occur in Early Pleistocene cold-water coral deposits on Sicily and Rhodes. Recent H. sarcophagahas not been observed above 11°C and is scarce near 5°C water temperature. Hyrrokkinsp. was reported from the Canadian Pacific on fossil sponges and was observed on Acesta patagonicain the Beagle Channel (Chile).

Acesta excavata is the largest known bivalve associated with the Lophelia pertusa communities along the northeast Atlantic continental margin, but has also been found along steep cliffs, devoid of bathyal corals, and on more subdued topographies. During the Pleistocene, including the last glacial, A. excavata was widespread in the Mediterranean, but is in the Recent fauna known only from four Mediterranean sites. A new possible live-record from the Canyon du Var off Nice is presented. Radiocarbon dating (AMS 14C-method) on an A. excavata assemblage from the Strait of Sicily yielded a Late Pleistocene age of 39.9 ka. Biometric parameters indicate that Pleistocene Mediterranean shells achieved about the same maximum sizes as their Recent Atlantic counterparts, but individuals have been slightly smaller on average. The subspecies Acesta excavata sublaevis Nordsieck, based on a subfossil juvenile Mediterranean shell, is not considered a valid taxon. Mineralogical composition and ultrastructure of A. excavata have been analyzed using thin-sections, Feigel staining, x-ray diffractometry (XRD) and SEM-imaging. Its wide distribution, large size (15 cm) and simple shell architecture makes it a prime candidate for palaeoenvironmental studies in cold-water coral settings.

The carnivorous sponge Asbestopluma hypogea, was known only from shallow submarine caves (-15 to -26 m) in the Western Mediterranean Sea and the Adriatic. Herein A. hypogea is reported from outside of caves, on seamounts in the Alboran Sea (-167 m), off the Balearics (-100 m) and north off Sicily (-660 m), and along steep bathyal escarpments in the Strait of Sicily (~700 m). These deeper ROV-based findings of A. hypogea are conform to the typical deep-sea occurrence of the Cladorhizidae.