Matthias López Correa

Earth's climate history serves as a natural laboratory for testing the effect of warm climates on the biosphere. The Cretaceous period featured a prolonged greenhouse climate characterized by higher-than-modern atmospheric CO2 concentrations and mostly ice-free poles. In such a climate, shallow seas in low latitudes probably became very hot, especially during the summers. At the same time, life seems to have thrived there in reef-like ecosystems built by rudists, an extinct group of bivalve molluscs. To test the seasonal temperature variability in this greenhouse period, and whether temperature extremes exceed the maximum tolerable temperatures of modern marine molluscs, we discuss a detailed sclerochronological (incrementally sampled) dataset of seasonal scale variability in shell chemistry from fossil rudist (Torreites sanchezi and Vaccinites vesiculosus) and oyster (Oscillopha figari) shells from the late Campanian (75-million-year-old) low latitude (3° S paleolatitude) Saiwan site in present-day Oman. We combine trace element data and microscopy to screen fossil shells for diagenesis, before sampling well-preserved sections of a Torreites sanchezi rudist specimen for clumped isotope analysis. Based on this specimen alone, we identify a strong seasonal variability in temperature of 19.2 ± 3.8 to 44.2 ± 4.0 °C in the seawater at the Saiwan site. The oxygen isotopic composition of the seawater (δ18Osw) varied from −4.62 ± 0.86 ‰ VSMOW in winter to +0.86 ± 1.6 ‰ VSMOW in summer. We use this information in combination with age modelling to infer temperature seasonality from incrementally sampled oxygen isotope profiles sourced from the literature, sampling multiple shells and species in the assemblage. We find that, on average, the Saiwan seawater experienced strong seasonal fluctuations in monthly temperature (18.7 ± 3.8 to 42.6 ± 4.0 °C seasonal range) and water isotopic composition (−4.33 ± 0.86 to 0.59 ± 1.03 ‰ VSMOW). The latter would strongly bias the interpretation of stable oxygen isotopes in shell carbonate without independent control on either temperature or seawater composition. Combining our seasonal temperature estimates with shell chronologies based on seasonal cyclicity in stable isotope records and daily variability in trace element data, we show that T. sanchezi rudists record temperatures during the hottest periods of the year as well as during the winters, which were characterized by cooler temperatures and enhanced influx of freshwater. Both O. figari and V. vesiculosus plausibly stopped growing during these seasonal extremes. This study aims to demonstrate how high-resolution geochemical records through fossil mollusc shells can shed light on the variability in past warm ecosystems and open the discussion about the limits of life in the shallow marine realm during greenhouse climates. Future work should apply the clumped isotope paleothermometer on incrementally sampled bio-archives to explore the upper-temperature limits experienced by calcifiers in different environments throughout geological history.

Earth’s climate history serves as a natural laboratory for testing the effect of warm climates on the biosphere. The Cretaceous period featured a prolonged greenhouse climate characterized by higher-than-modern atmospheric CO₂ concentrations and mostly ice-free poles. In such a climate, shallow seas in low latitudes probably became very hot, especially during the summers. At the same time, life seems to have thrived there in reef-like ecosystems built by rudists, an extinct group of bivalve molluscs. To test the seasonal temperature variability in this greenhouse period, and whether temperature extremes exceed the maximum tolerable temperatures of modern marine molluscs, we discuss a detailed sclerochronological (incrementally sampled) dataset of seasonal scale variability in shell chemistry from fossil rudist (Torreites sanchezi and Vaccinites vesiculosus) and oyster (Oscillopha figari) shells from the late Campanian (75-million-year-old) low latitude (3° S paleolatitude) Saiwan site in present-day Oman. We combine trace element data and microscopy to screen fossil shells for diagenesis, before sampling well-preserved sections of a Torreites sanchezi rudist specimen for clumped isotope analysis. Based on this specimen alone, we identify a strong seasonal variability in temperature of 19.2 ± 3.8 °C to 44.2 ± 4.0 °C in the seawater at the Saiwan site. The oxygen isotopic composition of the seawater (δ¹⁸O_sw) varied from -4.62 ± 0.86 ‰ VSMOW in winter to +0.86 ± 1.6 ‰ VSMOW in summer. We use this information in combination with age modelling to infer temperature seasonality from incrementally sampled oxygen isotope profiles sourced from the literature, sampling multiple shells and species in the assemblage. We find that, on average, the Saiwan seawater experienced strong seasonal fluctuations in monthly temperature (18.7 ± 3.8 to 42.6 ± 4.0 °C seasonal range) and water isotopic composition (-4.33 ± 0.86 to 0.59 ± 1.03 ‰ VSMOW). The latter would strongly bias the interpretation of stable oxygen isotopes in shell carbonate without independent control on either temperature or seawater composition. Combining our seasonal temperature estimates with shell chronologies based on seasonal cyclicity in stable isotope records and daily variability in trace element data, we show that T. sanchezi rudists record temperatures during the hottest periods of the year as well as during the winters, which were characterized by cooler temperatures and enhanced influx of freshwater. Both O. figari and V. vesiculosus plausibly stopped growing during these seasonal extremes. This study aims to demonstrate how high-resolution geochemical records through fossil mollusc shells can shed light on the variability in past warm ecosystems and open the discussion about the limits of life in the shallow marine realm during greenhouse climates. Future work should apply the clumped isotope paleothermometer on incrementally sampled bio-archives to explore the upper-temperature limits experienced by calcifiers in different environments throughout geological history.

Fibrous dolomite widely formed in Neoproterozoic marine sedimentary environments, but apparently disappeared in the Phanerozoic. Here, fibrous dolomite is recognised in a Miocene methane seep limestone (Marmorito, Italy) by synchrotron X-ray diffraction, length slow crystal optics and primary zonation under cathodoluminescence, which is unexpected. Low δ¹³C values and their negative correlation with MgCO₃ contents indicate a formation driven by highly alkaline pore waters and catalysis of dissolved sulphide generated by sulphate-driven anaerobic oxidation of methane. Cementing cavities of reefal carbonate, Neoproterozoic fibrous dolomite might have formed under sulphate-reducing conditions like Quaternary reef microbialites. Since the cavities of Neoproterozoic reefs were restricted microenvironments, the formation of fibrous dolomite was possibly favoured by catalysis similar to its Miocene seep counterpart. Our findings reinforce the concept of penecontemporaneous dolomite formation by sulphide catalysis and contribute to our understanding of the environmental conditions of the Neoproterozoic.

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.

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.

Barnacle borings were found in six heterocoral skeletons. They are present as small (up to 2.9 mm long), slender, pouch-shaped borings with tapering, slit-like openings. The investigated borings were made by acrothoracican cirripedes, which mostly do not have a shell and bore into hard substrates to protect their “naked” bodies. The first occurrence of borings in skeletons of the heterocoral Oligophylloides Różkowska, 1969 from the Upper Devonian Tafilalt Platform was reported recently by Weyer in 2016. Here, the authors present the results of a detailed study of heterocoral remains with numerous acrothoracican borings from Jebel Bou Ifarherioun (Famennian, Anti-Atlas, Morocco). The borings were found on the basal part but also on broken branches and stems of the heterocoral corallum and occurred post mortem. There is no indication of a syn vivo coral-barnacle interaction with borings in tissue-covered areas. The authors used micro-CT scans to visualize the 3D morphology of the pits, their orientation, and distribution. Additionally, the 3D morphology of an assemblage of 75 pits was used to carry out ordination and cluster analyses, which showed that previously proposed ichnospecies may be a continuum of morphological variability. In the basis of measurements by the present authors, the studied borings do not fit any known ichnotaxa. The absence of bourrelets excludes the possibility that the borings studied belong to the ichnogenus Rogerella Saint-Seine, 1951. Hence, the results seem to contradict a synonymization that was proposed by Bromley and D’Alessandro (1987) and subsequent authors and leave room for further research and discussion on this topic. Although the inferred boring organism is a filter feeder and, thus, depends on currents, the authors did not find a preferential orientation of the borings. The samples considered here are the best-preserved Devonian barnacle borings to date.

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.