A dynamic subtropical coastal hotspot of benthic foraminifera in the Southeastern Mediterranean indicates early-stage tropicalization.

Due to ongoing ocean warming, subtropical environments are becoming accessible to tropical species. Among these environments are the vermetid reefs of the Southeastern Mediterranean (SEM). In the last decades, these valuable coastal habitats witnessed the proliferation of numerous alien species of tropical origin. Among the meiofauna thriving on these reefs are benthic foraminifera, single cell marine organisms that make a significant contribution to global carbonate production. It has been widely recognized that benthic foraminifera, among other invasive species, thrive in the macroalgal cover, and it has been suggested that their populations are becoming a significant new source of sediment substrate. Here, we report on the first systematic assessment of the population size of the benthic foraminifera, allowing a comparison with data from the native tropical habitat of these species. Our study is based on a seasonal sampling of benthic foraminifera from confined sampling areas at four sites along the vermetid reef platforms of the Israeli SEM coast. Our survey reveals a patchy distribution of each species with peak population densities exceeding 100,000 specimens per m2, making the SEM a hotspot of benthic foraminifera, with population densities comparable to tropical coral reef environments. The assemblages of the SEM hotspot are dominated by cosmopolitan foraminiferal taxa and tropical invaders from the Indo-Pacific (e.g., Amphistegina lobifera, Pararotalia calcariformata, soritids, and Hauerina diversa). In contrast to foraminiferal hotspots in the tropics, which are completely dominated by larger symbiont-bearing taxa, the SEM hotspot stands out due to high abundances of non-symbiont-bearing species Textularia agglutinans and small miliolids. An intriguing observation is the significant heterogeneity in composition and density of foraminiferal assemblages between the vermetid reefs' southern and northern areas (Israel), indicating that the productivity of the dominant species are also modulated by local yet unknown environmental factors.

Active microbial communities facilitate carbon turnover in brine pools found in the deep Southeastern Mediterranean Sea.

Discharge of gas-rich brines fuels productive chemosynthetic ecosystems in the deep sea. In these salty, methanic and sulfidic brines, microbial communities adapt to specific niches along the physicochemical gradients. However, the molecular mechanisms that underpin these adaptations are not fully known. Using metagenomics, we investigated the dense (∼106 cell ml-1) microbial communities that occupy small deep-sea brine pools found in the Southeastern Mediterranean Sea (1150 m water depth, ∼22 °C, ∼60 PSU salinity, sulfide, methane, ammonia reaching millimolar levels, and oxygen usually depleted), reaching high productivity rates of 685 µg C L-1 d-1 ex-situ. We curated 266 metagenome-assembled genomes of bacteria and archaea from the several pools and adjacent sediment-water interface, highlighting the dominance of a single Sulfurimonas, which likely fuels its autotrophy using sulfide oxidation or inorganic sulfur disproportionation. This lineage may be dominant in its niche due to genome streamlining, limiting its metabolic repertoire, particularly by using a single variant of sulfide: quinone oxidoreductase. These primary producers co-exist with ANME-2c archaea that catalyze the anaerobic oxidation of methane. Other lineages can degrade the necromass aerobically (Halomonas and Alcanivorax), or anaerobically through fermentation of macromolecules (e.g., Caldatribacteriota, Bipolaricaulia, Chloroflexota, etc). These low-abundance organisms likely support the autotrophs, providing energy-rich H2, and vital organics such as vitamin B12.

Exceptionally high levels of total mercury in deep-sea sharks of the Southeastern Mediterranean sea over the last ∼ 40 years.

Deep-sea habitats are currently recognized as a hot spot for mercury (Hg) accumulation from anthropogenic sources, resulting in elevated concentrations of total mercury (THg) in deep-sea megafauna. Among them, deep-sea sharks (Class Chondrichthyes) are characterized by high trophic position and extended longevity and are, therefore, at high risk for mercury contamination. Despite this, sharks are overexploited by fishing activity in increasingly deeper water, worldwide, imposing health risks to human consumption. While it is imperative to better understand long-term mercury contamination in deep-sea megafauna, few historical data sets exist to capture this process. Here we explore four decades (1985-2022) of THg accumulation in five species of deep-sea sharks (G. melastomus, E. spinax, S. rostratus, C. granulosus, and D. licha) of the ultra-oligotrophic Southeastern Mediterranean Sea (SEMS) sampled during 19 research cruises. We exhibited exceptionally high THg levels (per length/weight), the highest as 16.6 µg g-1 (wet wt.), almost entirely (98.9 %; n = 298 specimens) exceeding the limit for safe consumption (0.3-0.5 µg THg g-1 wet wt.). The maximal THg levels of the long-lived species D. licha and C. granulosus in the SEMS were enriched by a factor of âˆ¼ 7 and >10 compared to counterpart species from other oceanic areas, respectively. We attribute this to the ultra-oligotrophic conditions of the SEMS, which cause slower growth rates and dwarfism in deep-sea sharks, resulting in an extended exposure time to mercury contamination. In the long-lived species, C. granulosus and D. licha, a temporal increase of average THg levels of âˆ¼ 80 % was recorded between 1987-1999 and 2021-2022. This likely reflects the long-term accumulation of historical anthropogenic Hg in deep-sea environments, which is further amplified in marginal seas such as the Mediterranean, impacted by global air pollution crossroads and surrounded by land-based pollution sources. Future consumption of products from deep-sea sharks is potentially high risk to human health.

Far-field effects of the Nile damming on the silica cycle in the Southeastern Mediterranean Sea.

Silica plays a key role in the growth of silicifying primary producers (e.g., diatoms) and hence the ocean carbon pump. The Mediterranean Sea's eastern Levantine Basin (ELB) is a low silica (and low N and P) ultra-oligotrophic basin. Before 1965, Nile autumn floods were a major source of dissolved silica (DSi) and other nutrients to primary producers of the ELB continental shelf, also known as the Nilotic cell. The construction of the Aswan High Dam (AHD) in the mid-1960s, blocked these floods, drastically diminishing the autumn-diatom blooms offshore the Nile delta. However, the far-reaching and long-lasting effects of the Nile damming on the Si cycle in the ELB remain unclear. Here, we studied the changes in DSi in the surface water offshore Israel and the distribution of biogenic silica in deep-sea short sediment cores, collected hundreds of kilometers from the Nile outlet, at depths range of 1100-1900 m, offshore the ELB Israeli coast. We show post dam reduction and termination in flood related seasonality of DSi and a concurrent decrease (of up to 79 %) in biogenic silica (BSi) accumulation rates in surficial sediments relative to underlying sediments. These changes reflect the effects of Si (dissolved and particulate) retention by the AHD on diatoms production, export and burial in the ELB. This far-field effect was demonstrated in deep-sea areas subjected to intense lateral transport of resuspended sediments from the shelf via intermediate nepheloid layers and to coastal water intrusions, along the path of the pre-dam, flood plumes. Our core records show that the AHD worsened nutrient-diminished, exceptionally unfavorable conditions for diatoms that persisted in the deep ELB at least during the last four millennia.

Accumulation of total mercury in deep-sea sediments and biota across a bathymetric gradient in the Southeastern Mediterranean Sea.

This study explores the accumulation of total mercury (THg) in deep-sea sediments and demersal megafauna of the ultra-oligotrophic Southeastern Mediterranean Sea (SEMS) across bathymetric gradients in the range 35-1900 m, sampled in seven cruises during 2013, 2017-2021, and 2023. Measurements of THg were conducted in surficial (0.0-0.5 cm) and subsurface (9.0-10 cm) sediments, demersal sharks, demersal teleost fish, and benthic crustaceans. Sedimentary organic carbon and biota δ13C and δ15N values were determined to explore possible foraging habitats and dietary sources of THg. The results exhibit an increasing trend of THg in surficial sediments with increasing bottom depth, while in the subsurface, pre-industrial sediments, THg remains lower, slightly increasing with depth. Having no major terrestrial point sources in this area, this increasing trend of THg in surficial sediments across bathymetric gradients is controlled by atmospheric mercury deposition, scavenged by the biological pump, and by lateral transport of particulate Hg in winnowed fine particles from the shelf. Similarly, the THg in benthic crustaceans and demersal fish ranged between 0.02 and 2.71 µg g-1 wet weight (0.06 and 10.8 µg g-1 dry weight) and increased with muscle δ13C as a function of distance offshore, while presenting a low THg-δ15N bio-magnification power. Our results suggest that foraging habitats, longevity, and species-specific depth distribution control their muscle THg bioaccumulation. Despite this complexity, the pooling of THg in megafauna into specific deep zones reflected the trend of increasing anthropogenic THg across bathymetric gradients. Furthermore, many of the biota measurements exceeded safe consumption thresholds for Hg and therefore, should be considered carefully in the development and regulation of deep-sea trawling in this region.

Impact of combined seawater warming and triazine-type herbicide pollution on the physiology and potential toxicity of the dinoflagellate Alexandrium minutum.

Coastal phytoplankton communities are often exposed to multiple anthropogenic stressors simultaneously. Here, we experimentally examined how temperature increase (20-26 °C) and triazine-type herbicides pollution (500 ng terbutryn L-1), both recognized as emerging stressors, affect the abundance, physiology and selected saxitoxin gene expression in the toxic dinoflagellate Alexandrium minutum. The results show that A. minutum is more susceptible to terbutryn pollution with increasing temperatures, resulting in a significant decline in its abundance (∼80 %) and photosynthetic activity (∼40 %), while saxitoxin gene expression increased (1.5-2.5-fold). This suggests that in warming polluted coastal areas where A. minutum is often found, saxitoxin poisoning may occur even in the absence of a massive bloom. Our results recommend the development of science-based monitoring practices for algal dissolved toxins in coastal waters and estuaries, supporting environmental policies under warming and contaminated coastal regions.

How can past sea level be evaluated from traces of anthropogenic layers in ancient saltpans?

Footprints of human activities identified in the sedimentary sequence of submerged historical saltpans can reveal the history of the site and can indicate the relative sea level during its operational period. Saltpans are man-made constructions used continuously for salt production in the Mediterranean at least for the last 2000 years. The east Adriatic coast contains many such submerged remains, preserved and well-dated by historical archives. Sedimentological, microfossil and geochemical analyses of the sediments from cores drilled in the saltwork area at Brbinj, Dugi Otok, Croatia, enable the reconstruction of various past environmental conditions. The current study aims to: a) identify the anthropogenic unit in the sedimentary sequence deposited over time, b) determine its age, and c) use it as past sea-level limiting points. Basal units made of terra rossa soil materials were identified in the sedimentary records. These layers are located -120 ±7 cm below mean sea level next to the separation wall and -125 ±7 cm and -135 ±7 cm, respectively, in the inner pools, most likely representing a man-made pavement. The terra rossa layer is overlaid by a unit rich in faunal remains dominated by euryhaline foraminifera and ostracod species such as Ammonia veneta and Cyprideis torosa, representing the saltworks unit. The flooding of the saltpans by the rising sea is manifested by the deposition of an upper sedimentary unit dominated by remains of marine species. The base and the top of the saltwork unit are dated by Optically Stimulated Luminescence to 1040±50 CE and to 1390±30 CE, respectively. The study presents a new approach for obtaining footprints of human activities in ancient, submerged saltpans, by identifying and dating the indicative anthropogenic layers and using these for the reconstruction of paleo sea-level. The described method can be applied all around the Mediterranean.

The FORCIS database: A global census of planktonic Foraminifera from ocean waters.

Planktonic Foraminifera are unique paleo-environmental indicators through their excellent fossil record in ocean sediments. Their distribution and diversity are affected by different environmental factors including anthropogenically forced ocean and climate change. Until now, historical changes in their distribution have not been fully assessed at the global scale. Here we present the FORCIS (Foraminifera Response to Climatic Stress) database on foraminiferal species diversity and distribution in the global ocean from 1910 until 2018 including published and unpublished data. The FORCIS database includes data collected using plankton tows, continuous plankton recorder, sediment traps and plankton pump, and contains ~22,000, ~157,000, ~9,000, ~400 subsamples, respectively (one single plankton aliquot collected within a depth range, time interval, size fraction range, at a single location) from each category. Our database provides a perspective of the distribution patterns of planktonic Foraminifera in the global ocean on large spatial (regional to basin scale, and at the vertical scale), and temporal (seasonal to interdecadal) scales over the past century.

Diversity, activity, and abundance of benthic microbes in the Southeastern Mediterranean Sea.

Benthic microbes are key organisms in the oligotrophic Southeastern Mediterranean Sea (SEMS), yet their abundance, activity, and diversity in this rapidly changing basin are not fully understood. We investigated the prokaryotic and microfungal communities throughout years 2018-2020 at 27 stations (6-1900 m water depths, down to 20 cm below the sediment surface), in two transects with distinct downslope transport regimes, and along the eutrophic coastline. We estimated microbial abundance with flow cytometry, secondary production as leucine assimilation, and sequenced marker genes (the 16S rRNA and internal transcribed spacer) to assess diversity indices. The highest abundance (0.21 × 108 cells gr-1 sediment) was estimated at slope stations where we assumed substantial transport rates and found an accumulation of organic carbon. Secondary production was the highest nearshore (12 ± 4 ng C gr-1 h-1), and markedly declined offshore (0.5 ± 0.9 ng C gr-1 h-1). Populations of archaea (dominant Nitrososphaeria and Nanoarchaeia) and diverse bacteria were stable over three years, and taxonomic composition was dictated mainly by depth gradients. Saprotrophic and pathotrophic microfungi Ascomycota (70% ± 23%) and Basidiomycota (16% ± 18%) were prevalent, whereas parasitic chytrids were abundant nearshore. Our results highlight the role of downslope transport, which enriched the typical deep-sea communities with anaerobic lineages, in shaping microbial populations near the continental slope.

The combined effects of rising temperature and salinity may halt the future proliferation of symbiont-bearing foraminifera as ecosystem engineers.

Rising sea surface temperatures and extreme heat waves are affecting symbiont-bearing tropical calcifiers such as corals and Large Benthic Foraminifera (LBF). In many ecosystems, parallel to warming, global change unleashes a host of additional changes to the marine environment, and the combined effect of such multiple stressors may be far greater than those of temperature alone. One such additional stressor, positively correlated to temperature in evaporation-dominated shallow-water settings is rising salinity. Here we used laboratory culture experiments to evaluate the combined thermohaline tolerance of one of the most common LBF species and carbonate producer, Amphistegina lobifera. The experiments were done under ambient (39 psu) and modified (30, 45, 50 psu) salinities and at optimum (25 °C) and warm temperatures (32 °C). Calcification of the A. lobifera holobiont was evaluated by measuring alkalinity loss in the culturing seawater, as an indication of carbonate ion uptake. The vitality of the symbionts was determined by monitoring pigment loss of the holobiont and their photosynthetic performances by measuring dissolved oxygen. We further evaluated the growth of Peneroplis (P. pertusus and P. planatus), a Rhodophyta bearing LBF, which is known to tolerate high temperatures, under elevated salinities. The results show that the A. lobifera holobiont exhibits optimal performance at 39 psu and 25 °C, and its growth is significantly reduced upon exposure to 30, 45, 50 psu and under all 32 °C treatments. Salinity and temperature exhibit a significant interaction, with synergic effects observed in most treatments. Our results confirm that Peneroplis has a higher tolerance to elevated temperature and salinity compared to A. lobifera, implying that a further increase of salinity and temperatures may result in a regime shift from Amphistegina- to Peneroplis-dominated assemblages.

Dust-borne microbes affect Ulva ohnoi's growth and physiological state.

The marine macroalgae Ulva sp. is considered an ecosystem engineer in rocky shores of temperate waters worldwide. Ulva sp. harbors a rich diversity of associated microbial epibionts, which are known to affect the algae's typical morphological development and 'health'. We examined the interaction between airborne microbes derived from atmospheric aerosols and Ulva ohnoi growth and physiological state. Specifically, we measured U. ohnoi growth rates and photosynthetic efficiency (Fv/Fm), alongside its microbial epibionts abundance, activity and diversity following dust (containing nutrients and airborne microorganisms) or UV-treated dust (only nutrients) amendments to filtered seawater. Parallel incubations with epibionts-free U. ohnoi (treated with antibiotics that removed the algae epibionts) were also tested to specifically examine if dust-borne microbes can replenish the epibiont community of U. ohnoi. We show that viable airborne microbes can restore U. ohnoi natural microbial epibionts communities, thereby keeping the seaweed alive and 'healthy'. These results suggest that microbes delivered through atmospheric aerosols can affect epiphyte biodiversity in marine flora, especially in areas subjected to high annual atmospheric dust deposition such as the Mediterranean Sea.

Epiphytic benthic foraminiferal preferences for macroalgal habitats: Implications for coastal warming.

Considering the thermal limits of coastal macroalgae habitats in the South-Eastern Mediterranean, it is important to study the response of the associated meiofauna to better understand the expected feedback of ecosystems to future warming. In this study, we compared benthic foraminiferal assemblages from two common macroalgal habitats, Turf and Coralline algae, based on ecological monitoring of a thermally polluted station representing near future warming, and an undisturbed environment. None of the common local species is confined to a specific algal habitat. This implies that their existence is not threatened by the disappearance of the Coralline algae. However, most likely their community structure will be impacted with coastal warming. Species that are more affiliated with Coralline algae are highly thermally tolerant, thus their proliferation might be reduced with warming. Specifically, the negative response of Coralline algae to warming may limit the contribution of invasive species such as Pararotalia calcariformata.

Shell Growth of Large Benthic Foraminifera under Heavy Metals Pollution: Implications for Geochemical Monitoring of Coastal Environments.

This study was promoted by the recent efforts using larger benthic foraminiferal (LBF) shells geochemistry for the monitoring of heavy metals (HMs) pollution in the marine environment. The shell itself acts as a recorder of the ambient water chemistry in low to extreme HMs-polluted environments, allowing the monitoring of recent-past pollution events. This concept, known as sclerochronology, requires the addition of new parts (i.e., new shell) even in extreme pollution events. We evaluated the physiological resilience of three LBF species with different shell types and symbionts to enriched concentrations of Cd, Cu, and Pb at levels several folds higher than the ecological criteria maximum concentration (CMC) (165-166, 33-43, 1001-1206 µg L-1, respectively), which is derived from aquatic organisms' toxicity tests. The physiological response of the holobiont was expressed by growth rates quantified by the addition of new chambers (new shell parts), and by the chlorophyll a of the algal symbionts. The growth rate decrease varied between 0% and 30% compared to the unamended control for all HMs tested, whereas the algal symbionts exhibited a general non-fatal but significant response to Pb and Cu. Our results highlight that shell growth inhibition of LBF is predicted in extreme concentrations of 57 × CMC of Cu and 523 × CMC of Cd, providing a proof of concept for shell geochemistry monitoring, which is currently not used in the regulatory sectors.

Monitoring long-term spatial and temporal trends of the infaunal community characteristics along the shallow waters of the Mediterranean coast of Israel.

Sandy sediment and its infauna were annually sampled along the shallow waters of the Israeli coast during the 2005-2016 period, as a part of the Israeli National Environmental Program framework, aiming to detect anthropogenic interference in that province by monitoring changes in the species composition, abundance, and diversity of the infaunal communities and in accompanied abiotic parameters: the levels of total organic carbon and a series of heavy metals and the site-specific grain size distribution. The > 250-µm fraction of the fauna was segregated from the sampled sediment and was identified to species or higher taxonomic level. Three spatial biotopes were determined based on their unique faunal composition, Haifa Bay, Haifa harbor, and the southern coast. Species homogeneity among samples of each biotope was evaluated. Temporal and spatial changes of the species composition, abundance, and diversity were calculated for each biotope, mostly revealing random annual fluctuations. Only two minor temporal trends were observed: two spatially identical and temporally different faunal communities in the southern coast biotope, distinguishing the 2005-2007 and 2008-2016 periods, and a slight increase in the number of species across time in the two Haifa Bay provinces. Total organic carbon was highly correlated to the faunal composition with the highest organic carbon levels in the Haifa harbor biotope. The biotopes' mutually occurring abundant species were sufficient to determine biotope borders and the contribution of intermittently sampled rare species, including the zoogeographically Indo-Pacific originated ones was feeble, important only to identify species migration and faunistics. Practically, three sampling sites along the Israeli shallow soft substrate, corresponding to the defined spatial biotopes, are sufficient to monitor the effect of environmental changes. Seasonal sampling twice a year is recommended as well as more accurate species identification using molecular taxonomy.

Environmental status of Israel's Mediterranean coastal waters: Setting reference conditions and thresholds for nutrients, chlorophyll-a and suspended particulate matter.

Criteria for eutrophication related parameters to achieve and preserve good environmental status (GES) of the oligotrophic Israeli Mediterranean coast were proposed for nutrients, chlorophyll-a (Chl-a) and suspended particulate matter (SPM) concentrations. The criteria were derived from current conditions, the best choice for the area that has undergone large and irreversible ecological changes compared to the pristine background. A five-year data set (2010-2014, ca. 800 data points) was analyzed using statistical methods and best professional judgement. The coastal waters were divided into four provinces, data gaps were identified, and seasonal reference and threshold values for each province determined as the median and 1.5 times the median, respectively. Application of the derived criteria to data up to 2016 showed the coastal waters to be mainly in GES, with a few exceptions. Simplification of the proposed criteria for environmental management was addressed as well.

Thermal tolerance and range expansion of invasive foraminifera under climate changes.

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.

Foraminiferal single chamber analyses of heavy metals as a tool for monitoring permanent and short term anthropogenic footprints.

In order to establish environmentally sustainable industries there is a need for high-resolution temporal and spatial monitoring of heavy metal pollutants even at low concentrations before they become hazardous for local ecosystems. Here we present single chamber records of Cu, Zn and Pb in shells of two benthic foraminifera species with different shell types from two shallow coastal stations in Israel: An area adjacent to an electrical power plant and desalination factory (Hadera) and an industrially free nature reserve (Nachsholim). Records of both foraminifera species show elevated metal concentrations in Hadera clearly identifying the footprint of the local industrial facilities. Moreover, short-term events of elevated Cu and Pb concentrations were detected by single chamber analyses. This study demonstrates the potential of using heavy metals anomalies in foraminiferal single chambers as a tool for detecting the industrial footprint of coastal facilities as well as short term events of elevated heavy metals.

Impact of nutrient enrichment on productivity of coastal water along the SE Mediterranean shore of Israel - A bioassay approach.

The coastal waters of the southeastern Mediterranean-Sea (SEMS) are routinely enriched with naturally-occurring and anthropogenic land-based nutrient loads. These external inputs may affect autotrophic and heterotrophic microbial biomass and activity. Here, we conducted 13 microcosm bioassays with different additions of inorganic NO3-(N), PO4-(P) and Si(OH)4-(Si) in different seasons along the Mediterranean coast of Israel. Our results indicate that cyanobacteria are mainly N-limited, whereas N or Si (or both) limit pico-eukaryotes. Furthermore, the degree to which N affects phytoplankton depends on the ambient seawater's inorganic N and N:P characteristics. Heterotrophic bacteria displayed no response in all treatments, except when all nutrients were added simultaneously, suggesting a possible co-limitation by nutrients. These results contrast the N+P co-limitation of phytoplankton and the P-limitation of bacteria in the open waters of the SEMS. These observations enable the application for a better science-based environmental monitoring and policy implementation along the SEMS coast of Israel.

Co-existence of Methanogenesis and Sulfate Reduction with Common Substrates in Sulfate-Rich Estuarine Sediments.

The competition between sulfate reducing bacteria and methanogens over common substrates has been proposed as a critical control for methane production. In this study, we examined the co-existence of methanogenesis and sulfate reduction with shared substrates over a large range of sulfate concentrations and rates of sulfate reduction in estuarine systems, where these processes are the key terminal sink for organic carbon. Incubation experiments were carried out with sediment samples from the sulfate-methane transition zone of the Yarqon (Israel) estuary with different substrates and inhibitors along a sulfate concentrations gradient from 1 to 10 mM. The results show that methanogenesis and sulfate reduction can co-exist while the microbes share substrates over the tested range of sulfate concentrations and at sulfate reduction rates up to 680 µmol L-1 day-1. Rates of methanogenesis were two orders of magnitude lower than rates of sulfate reduction in incubations with acetate and lactate, suggesting a higher affinity of sulfate reducing bacteria for the available substrates. The co-existence of both processes was also confirmed by the isotopic signatures of δ34S in the residual sulfate and that of δ13C of methane and dissolved inorganic carbon. Copy numbers of dsrA and mcrA genes supported the dominance of sulfate reduction over methanogenesis, while showing also the ability of methanogens to grow under high sulfate concentration and in the presence of active sulfate reduction.

Geochemical signatures of benthic foraminiferal shells from a heat-polluted shallow marine environment provide field evidence for growth and calcification under extreme warmth.

Shallow marine calcifiers play an important role as marine ecosystem engineers and in the global carbon cycle. Understanding their response to warming is essential to evaluate the fate of marine ecosystems under global change scenarios. A rare opportunity to test the effect of warming acting on natural ecosystems is by investigation of heat-polluted areas. Here, we study growth and calcification in benthic foraminifera that inhabit a thermally polluted coastal area in Israel, where they are exposed to elevated temperatures reaching up to ~42°C in summer. Live specimens of two known heat-tolerant species Lachlanella sp. 1 and Pararotalia calcariformata were collected over a period of 1 year from two stations, representing thermally polluted and undisturbed (control) shallow hard bottom habitats. Single-chamber element ratios of these specimens were obtained using laser ablation, and the Mg/Ca of the most recently grown final chambers were used to calculate their calcification temperatures. Our results provide the first direct field evidence that these foraminifera species not only persist at extreme warm temperatures but continue to calcify and grow. Species-specific Mg/Ca thermometry indicates that P. calcariformata precipitate their shells at temperatures as high as 40°C and Lachlanella sp. 1 at least up to 36°C, but both species show a threshold for calcification at cold temperatures: calcification in P. calcariformata only occurred above 22°C and in Lachlanella sp. 1 above 15°C. Our observations from the heat-polluted area indicate that under future warming scenarios, calcification in heat-tolerant foraminifera species will not be inhibited during summer, but instead the temperature window for their calcification will be expanded throughout much of the year. The observed inhibition of calcification at low temperatures indicates that the role of heat-tolerant foraminifera in carbonate production will most likely increase in future decades.