Present-day ocean deoxygenation has major implications for marine ecosystems and biogeochemical cycling in the oceans. Chromium isotopes are used as a proxy to infer changes in past oceanic redox state. Chromium isotopes in carbonates, including the prime proxy carrier foraminifera, were initially thought to record the seawater composition during crystallisation. However, the uptake of Cr into foraminiferal tests and carbonates is still poorly understood and recent studies question this assumption. We assess whether Cr in foraminiferal calcite is taken up during biomineralisation, has a postdepositional origin or is a combination of the two. Laser Ablation-MC-ICP-MS analyses and NanoSIMS imaging of individual tests were used to characterise the distribution of Cr in both planktic and benthic foraminifera. Foraminifera in sediment core-top samples have up to two orders of magnitude more Cr than sediment trap, plankton net, and culture samples. In cultured specimens, Cr is incorporated in foraminiferal tests at low concentrations (0.04-0.13 ppm) with a distribution coefficient of ~250 ± 43 (2SE) which is an upper estimate due to substantial loss of dissolved Cr during the experiment. Part of the Cr signal in sedimentary foraminifera may be primary, but this primary signal is likely often overprinted by the uptake of Cr from bottom and pore waters. In sediment samples, there is no significant isotopic offset between individual species and bulk foraminiferal calcite from the same size fraction. The >500 µm fraction has a heavier isotopic composition than the smaller 250-500 µm fraction with an offset of-0.3 to-0.5‰ due to an increase in surface area to volume. We propose that Cr in foraminifera is predominantly post-depositional and records bottom/pore water signals. This is contrary to current interpretations of the foraminiferal Cr isotope proxy as a surface seawater redox proxy.

No abstract

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 was 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.

Bleaching, the loss of algal symbionts, occurs in marine photosymbiotic organisms at water temperatures minimally exceeding average summer SST (sea surface temperatures). Pre-adaptation allows organisms to persist under warmer conditions, providing the tolerance can be carried to new habitats. Here we provide evidence for the existence of such adaptation in the benthic foraminifera Pararotalia calcariformata. This species occurs at a thermally polluted site in the Mediterranean, where water temperatures reach a maxima daily average of 36 °C during the summer. To test whether this occurrence represents a widespread adaptation, we conducted manipulative experiments exposing this species from an unpolluted site to elevated temperatures (20–42 °C). It was kept in co-culture with the more thermally sensitive foraminifera Amphistegina lobifera in two experiments (20–36 °C). Reduced photosynthetic activity in A. lobifera occurred at 32 °C whereas photochemical stress in P. calcariformata was first observed during exposure to 36 °C. Pararotalia calcariformata survived all treatment conditions and grew under 36 °C. The photosymbiosis in P. calcariformata is unusually thermally tolerant. These observations imply that marine eukaryote-eukaryote photosymbiosis can respond to elevated temperatures by drawing on a pool of naturally occurring pre-adaptations. It also provides a perspective on the massive occurrence of symbiont-bearing foraminifera in the early Cenozoic hothouse climate.

The Eastern Mediterranean is experiencing a large-scale invasion of alien tropical species from the Red Sea. This “Lessepsian invasion” began with the opening of the Suez Canal and is promoted by the ongoing oceanic warming. The environmental differences between the Red Sea and the Mediterranean act as a buffer allowing the invasion of certain species. This provides an opportunity to study the differences in temperature sensitivity between two sibling species of the cosmopolitian foraminifera Amphistegina. Both species are very common in the Red Sea. Whilest, only one is a successful invader and the other is absent in the Eastern Mediterranean. Here we show that the two species are different in their temperature sensitivity, which explains their selective invasion into the Mediterranean. These differences demonstrate that in respect to climate change resilient marine species can be distinguished by their ability to compensate for temperature changes by adjusting their physiological performance and by having tolerance to a wider temperature range. Moreover, we demonstrate that selective filtering mechanisms during invasion can prefer species that are more resilient to colder rather than expected warmer temperatures.