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.

In sub/tropical waters, benthic foraminifera are among the most abundant epiphytic organisms inhabiting seagrass meadows. This study explored the nature of the association between foraminifera and the tropical seagrass species H. stipulacea, aiming to determine whether these interactions are facilitative or random. For this, we performed a "choice" experiment, where foraminifera could colonize H. stipulacea plants or plastic "seagrasses" plants. At the end of the experiment, a microbiome analysis was performed to identify possible variances in the microbial community and diversity of the substrates. Results show that foraminifera prefer to colonize H. stipulacea, which had a higher abundance and diversity of foraminifera than plastic seagrass plants, which increased over time and with shoot age. Moreover, H. stipulacea leaves have higher epiphytic microbial community abundance and diversity. These results demonstrate that seagrass meadows are important hosts of the foraminifera community and suggest the potential facilitative effect of H. stipulacea on epiphytic foraminifera, which might be attributed to a greater diversity of the microbial community inhabiting H. stipulacea.

In this study we revisited the Cretaceous of the Eratosthenes Seamount (ESM) from IODP LEG 160 Hole 967E, updating the chronology, depositional environment, and paleobathymetry of the ESM. Our goal was also to address the spatio-temporal distribution of organic matter and, by comparison with the eastern margins of the Levant Basin, discuss basin-wide controls on its deposition and preservation.

The investigated core has a relatively continuous Cretaceous succession from the Aptian to the Danian. By identifying the Pα Zone we conclude that the base of the Paleocene is included in the 967E section. A total of 17 ages were identified, with low sedimentation rates of 0.04–

2.37 cm/kyr. Petrographic analysis revealed pronounced differences, from micritic pellets, benthic foraminifera, algae, and mollusks in the bottom part of the core, to deep water facies in the uppermost Cretaceous. Paleobathymetry ranged between 0.5 and 5 m in the Aptian to 300–600 m in the Maastrichtian.

Both high productivity and OMZ conditions prevailed during the Cenomanian–Turonian, as well as elsewhere in the eastern Mediterranean. While the low core recovery might affect reliable interpretation, we present two alternative explanations for the high vs. low organic content of the Campanian–Maastrichtian in the eastern Levant Basin vs. the ESM: (1) Location relative to upwelling cells; (2) Relative bathymetry, with the ESM representing a paleohigh for much of the Cretaceous, similar to the organic-poor anticline deposits in the eastern Levant, where the massive intervals of organic-rich carbonates accumulated only in the synclines of the Syrian Arc deformational belt.

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.

Global warming permits range expansions of tropical marine species into mid-latitude habitats, where they are, however, faced with cold winter temperatures. Therefore, tolerance to cold temperatures may be the key adaptation controlling zonal range expansion in tropical marine species. Here we investigated the molecular and physiological response to cold and heat stress in a tropical symbiont-bearing foraminifera that has successfully invaded the Eastern Mediterranean. Our physiological measurements indicate thermal tolerance of the diatom symbionts but a decrease in growth for the foraminifera host under both cold and warm stress. The combined (‘holobiont’) transcriptome revealed an asymmetric response in short-term gene expression under cold versus warm stress. Cold stress induced major reorganization of metabolic processes, including regulation of genes involved in photosynthesis. Analyses limited to genes that are inferred to belong to the symbionts confirm that the observed pattern is due to changes in the regulation of photosynthesis-related genes and not due to changes in the abundance of the symbionts. In contrast to cold stress, far fewer genes change expression under heat stress and those that do are primarily related to movement and cytoskeleton. This implies that under cold stress, cellular resources are allocated to the maintenance of photosynthesis, and the key to zonal range shifts of tropical species could be the cold tolerance of the symbiosis.

The recent growing interest in Late Cretaceous biserial planktonic foraminifera (heterohelicids) has greatly enhanced their use as paleoceanographic and biostratigraphic markers. Pseudotextularia nuttalli, one of the most common cosmopolitan planktonic foraminifera, has an exceptionally long evolutionary range (Turonian-Maastrichtian). The image processing procedure we developed in this study enabled us to document changes in the growth pattern of Ps. nuttalli over time and between different oceanic provinces, and to identify possible speciation events within this lineage. The analysis was complemented by morphometric measurements of the penultimate chamber and test.

The morphometric analyses do not indicate an early speciation event within this lineage, and thus, do not support splitting to Planoheterohelix praenuttalli (Turonian-Coniacian) and Pseudotextularia nuttalli (Coniacian-Maastrichtian). Our results indicate a gradual phyletic increase in mean test size from the Santonian through the Maastrichtian, along with increasing morphological diversification. This trend seems to coincide with the global cooling that culminated in the Late Cretaceous.

Comparing between Ps. nuttalli from different oceanic provinces reveals that the Southern Tethys specimens are significantly less developed than their counterparts from the two tropical oceanic localities. This adaptive responsiveness suggests a negative correlation between test size and high productivity conditions in the upper water column.

The case of Ps. nuttalli lineage exemplifies how extraordinary long-range species (>30 My) can evolutionarily change over time from “primitive” to “developed” forms, without showing a morphological gap. Such species may also show adaptive morphological diversification in response to different environments and, thus, are valuable for palaeoceanographic and paleoecologic reconstructions.

The recovery time of marine productivity following the Cretaceous-Paleogene (KPg) mass extinction varies tremendously with location (hundreds to millions of years), with possible delays in the tropics as compared to higher latitudes. This heterogeneity is based on prevalent oligo- to mesotrophic marine environments. While highly productive eutrophic environments are less prevalent, they play a greater role in the carbon cycle. Here we present data from a eutrophic region within the tropical southern Tethys. Records of both organic matter and calcite production in this locality exhibit stability across the KPg boundary. In addition, our study points to a remarkably rapid recovery (<11 kyr), even possibly continuous high productivity across the KPg boundary, despite the tropical location. Moreover, the characteristic KPg δ¹³C negative excursion is observed in our locality, but is independent of high productivity, possibly indicating a reduction in the δ¹³C of the DIC of the ocean due to the input of light carbon from the atmosphere. Thus, this study provides new insight into the functioning of eutrophic ecosystems during environmental stress imposed by the ecological crisis.

Mercury (Hg) anomalies linked to Large Igneous Provinces (LIP) volcanism have been identified in sediments across all five major mass extinctions in Earth's history. This study tests whether Hg in marine sediments is a reliable proxy linking Deccan Traps volcanic eruptions to late Maastrichtian global climate warming and the mass extinction at the Cretaceous-Paleogene boundary (KPB). Our primary test site is the Elles section in Tunisia, the auxiliary Global Stratotype Section and Point (GSSP) to El Kef. Elles has the most complete marine sedimentary record and a high average sedimentation rate of ~4.7 cm/ky. We chose the Hor Hahar section in Israel to corroborate the geographic distribution of Hg fallout from Deccan volcanism. Reliability of the Hg proxy over the last 550 ky of the Maastrichtian to early Danian was evaluated based on high-resolution age control (orbital cyclostratigraphy), stable isotope climate record, Hg concentrations, biotic turnover and mass extinction. These results were correlated with the pulsed Deccan eruptive history constrained previously by U-Pb zircon geochronology.

Our results support Hg as robust proxy for Deccan volcanism with large Hg spikes marking “extreme event” (EE) pulsed eruptions correlative with climate warming peaks separated by steady, less intense eruptions. Long-term global climate warming began near ~350 ky pre-KPB, reached maximum warming (3–4 °C) between 285 and 200 ky pre-KPB, followed by gradual cooling and rapid temperature drop between 45 and 25 ky pre-KPB. During the last 25 ky before the KPB, multiple Hg EE eruptions correlate with hyperthermal warming that culminated in the rapid mass extinction at Elles during ≤1000 years of the Cretaceous. These latest Cretaceous Hg peaks may correlate with massive, distal, Deccan-sourced lava flows (>1000 km long) that traversed the Indian subcontinent and flowed into the Bay of Bengal, bracketing the mass extinction. These results support Deccan volcanism as a primary driver of the end-Cretaceous mass extinction.

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.

Following a multi-proxy analysis of the Upper Cretaceous high-productivity sequence from proximal and distal basins in Israel, Meilijson et al. (Paleobiology 42:77–97, 2016) provided evidence indicating that different benthic foraminifera species could survive and sustain large populations under long-term anoxic to dysoxic bottom water conditions. They proposed that massive blooms of triserial (buliminid) benthic foraminifera with distinct apertural and test morphologies during the Campanian managed to survive anoxic conditions by their capability to sequester diatom chloroplasts (kleptoplastidy) and associate with bacteria, in a similar manner as their modern analogues. This advantageous capability as well as other adaptations such as using nitrate instead of oxygen for their respiratory pathways, or changes in food type arriving to the seafloor, were all affected by the substantial shift in the depositional environment following the Campanian/Maastrichtian boundary. However, several of the hypothesis and assumptions presented in this chapter called for a continued study of the Upper Cretaceous deposits in the Levant, to better constrain the oceanographic and bottom water process in which these organisms lived.

Here we report on a high-resolution investigation focused on the inorganic geochemical properties of two sections within the high-productivity setting of the Late Cretaceous in the Levant. Benthic foraminiferal assemblages were compared with the trace metal enrichment, bottom water renewal and water column oxygen levels, on a high-productivity seafloor. Our work focused on the occurrence and distribution of redox-sensitive/sulphide-forming trace metals obtained by analytical approaches (bulk sediment composition, ED- and WD-XRF), in the organic-rich sediments. On basis of the bulk sediment geochemistry, a principal component analysis distinguished between two factors for both sections: Factor 1 mirrors the degree of bottom-water oxygenation (Cu, S, Ni, Zn, Cr and Corg) and includes elements representing enhanced phosphorite deposition (P₂O₅, U, As, Mo, Y). Factor 2 reflects the interplay between the input of biogenic carbonate (Ca) and terrigenous material (TiO₂, Rb, SiO₂, Fe₂O₃(t), Ce, Ga, V and Al₂O₃). An additional factor was used in the distal and deeper of the two sections to distinguish between times in which dominance of siliceous or calcareous biogenic sedimentation occurred. We observe that the lowest part of the Maastrichtian contained the strongest reducing conditions, whereas the upper part was affected by a lesser degree of oxygen deficiency. Geochemical results of the molybdenum-to-organic carbon ratio reveal a change in the water mass circulation to more restrictive condition within the lower Maastrichtian, which coincides with reported sea-level rise. Based on factor analysis of the elemental distribution we demonstrate a clear connection between diatom abundance and peaks in the abundance of foraminifera species thought to have used kleptoplastidy as a morphological adaptation to cope with environmental instability, advocating previous assumptions and hypothesis. Additionally, it is evident that along the section in which fluctuations in the relative abundance of primary producers occurred, substantial shifts also transpired in the relative contribution of terrigenic material to the accumulating sediments. The synchronous occurrence of abnormally high numbers of low-diversity benthic foraminifera demonstrates the existence and success of functional adaptations. Our identification of morphological adaptations in Praebulimina prolixa, which are identical to those recognized in modern diatom-based kleptoplastidy of benthic foraminifera, acts as the missing link for understanding this complex system. It does so by tying between productivity, oceanography, continental-marine interactions and remarkable biochemical reciprocity and adaptiveness of present and deep-time ecosystems.