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Foraminiferal wall microstructures, consistent with the molecular-based high-rank classification, are critical to understanding foraminiferal evolution and advanced taxonomic relationships. Although test structures are well documented for recent, Cenozoic, and some Mesozoic foraminifera, the diagnostic characteristics of Paleozoic taxa are largely unexplored. The majority of calcareous Paleozoic foraminifera have been assigned to the Fusulinata based on questionable homogeneously “microgranular” test wall microstructures, which have never been sufficiently documented for most taxa. We investigated the test structures of exceptionally well-preserved Devonian (Eifelian) Semitextularia thomasi, representing the first calcareous true multichambered (serial) foraminifera, and compared this species with a large fusiform Permian representative of “true” fusulinids (Neoschwagerinidae). The tests of Semitextularia thomasi display lamellar structures that are not observed in any other fossil or recent foraminiferal group. The Paleozoic foraminifera, traditionally referred to one taxon (the class Fusulinata), possess at least three contrasting test wall microstructures, representing separate high-rank taxonomic groups. Fusulinata is most likely a highly polyphyletic group that is in need of taxonomic revision. The term Fusulinata, defined as including all Paleozoic calcareous forms except Miliolida and Lagenata, is not phylogenetically meaningful and should no longer be used or should be restricted to true complex fusulinids with microgranular test structures, which appeared in the Carboniferous.

The Cretaceous/Palaeogene mass extinction eradicated 76% of species on Earth 1 , 2 . It was caused by the impact of an asteroid 3 , 4 on the Yucatán carbonate platform in the southern Gulf of Mexico 66 million years ago 5 , forming the Chicxulub impact crater 6 , 7 . After the mass extinction, the recovery of the global marine ecosystem—measured as primary productivity—was geographically heterogeneous 8 ; export production in the Gulf of Mexico and North Atlantic–western Tethys was slower than in most other regions 8 – 11 , taking 300 thousand years (kyr) to return to levels similar to those of the Late Cretaceous period. Delayed recovery of marine productivity closer to the crater implies an impact-related environmental control, such as toxic metal poisoning 12 , on recovery times. If no such geographic pattern exists, the best explanation for the observed heterogeneity is a combination of ecological factors—trophic interactions 13 , species incumbency and competitive exclusion by opportunists 14 —and ‘chance’ 8 , 15 , 16 . The question of whether the post-impact recovery of marine productivity was delayed closer to the crater has a bearing on the predictability of future patterns of recovery in anthropogenically perturbed ecosystems. If there is a relationship between the distance from the impact and the recovery of marine productivity, we would expect recovery rates to be slowest in the crater itself. Here we present a record of foraminifera, calcareous nannoplankton, trace fossils and elemental abundance data from within the Chicxulub crater, dated to approximately the first 200 kyr of the Palaeocene. We show that life reappeared in the basin just years after the impact and a high-productivity ecosystem was established within 30 kyr, which indicates that proximity to the impact did not delay recovery and that there was therefore no impact-related environmental control on recovery. Ecological processes probably controlled the recovery of productivity after the Cretaceous/Palaeogene mass extinction and are therefore likely to be important for the response of the ocean ecosystem to other rapid extinction events. Micro- and nannofossil, trace fossil and geochemical evidence from the Chicxulub impact crater demonstrates that proximity to the asteroid impact site did not determine rates of recovery of marine ecosystems after the end-Cretaceous mass extinction.

Planktonic foraminifera are one of the most abundant and diverse protists in the oceans. Their utility as paleo proxies requires rigorous taxonomy and comparison with living and genetically related counterparts. We merge genetic and fossil evidence of \"Globigerinoides\", characterized by supplementary apertures on spiral side, in a new approach to trace their \"total evidence phylogeny\" since their first appearance in the latest Paleogene. Combined fossil and molecular genetic data indicate that this genus, as traditionally understood, is polyphyletic. Both datasets indicate the existence of two distinct lineages that evolved independently. One group includes \"Globigerinoides\" trilobus and its descendants, the extant \"Globigerinoides\" sacculifer, Orbulina universa and Sphaeroidinella dehiscens. The second group includes the Globigerinoides ruber clade with the extant G. conglobatus and G. elongatus and ancestors. In molecular phylogenies, the trilobus group is not the sister taxon of the ruber group. The ruber group clusters consistently together with the modern Globoturborotalita rubescens as a sister taxon. The re-analysis of the fossil record indicates that the first \"Globigerinoides\" in the late Oligocene are ancestral to the trilobus group, whereas the ruber group first appeared at the base of the Miocene with representatives distinct from the trilobus group. Therefore, polyphyly of the genus \"Globigerinoides\" as currently defined can only be avoided either by broadening the genus concept to include G. rubescens and a large number of fossil species without supplementary apertures, or if the trilobus group is assigned to a separate genus. Since the former is not feasible due to the lack of a clear diagnosis for such a broad genus, we erect a new genus Trilobatus for the trilobus group (type species Globigerina triloba Reuss) and amend Globoturborotalita and Globigerinoides to clarify morphology and wall textures of these genera. In the new concept, Trilobatus n. gen. is paraphyletic and gave rise to the Praeorbulina/Orbulina and Sphaeroidinellopsis/Sphaeroidinella lineages.

Foraminifera are a group of mostly marine protists with high taxonomic diversity. Species identification is often complex, as both morphological and molecular approaches can be challenging due to a lack of unique characters and reference sequences. An integrative approach combining state of the art morphological and molecular tools is therefore promising. In this study, we analysed large benthic Foraminifera of the genus Amphisorus from Western Australia and Indonesia. Based on previous findings on high morphological variability observed in the Soritidae and the discontinuous distribution of Amphisorus along the coast of western Australia, we expected to find multiple morphologically and genetically unique Amphisorus types. In order to gain detailed insights into the diversity of Amphisorus , we applied micro CT scanning and shotgun metagenomic sequencing. We identified four distinct morphotypes of Amphisorus , two each in Australia and Indonesia, and showed that each morphotype is a distinct genotype. Furthermore, metagenomics revealed the presence of three dinoflagellate symbiont clades. The most common symbiont was Fugacium Fr5, and we could show that its genotypes were mostly specific to Amphisorus morphotypes. Finally, we assembled the microbial taxa associated with the two Western Australian morphotypes, and analysed their microbial community composition. Even though each Amphisorus morphotype harboured distinct bacterial communities, sampling location had a stronger influence on bacterial community composition, and we infer that the prokaryotic community is primarily shaped by the microhabitat rather than host identity. The integrated approach combining analyses of host morphology and genetics, dinoflagellate symbionts, and associated microbes leads to the conclusion that we identified distinct, yet undescribed taxa of Amphisorus . We argue that the combination of morphological and molecular methods provides unprecedented insights into the diversity of foraminifera, which paves the way for a deeper understanding of their biodiversity, and facilitates future taxonomic and ecological work.

Representatives of the foraminifer Nummulites are important in Earth history for timing Cenozoic shallow-water carbonates. Taphonomic complexity explains the construction of carbonate buildups, but reproduction and life span of the constructing individuals are unknown. During the 15-month investigation period, asexually reproduced schizonts and gamonts showed equal proportions in the first half of this period, whereas gamonts predominated in the second half. Oscillations in cell growth are mainly caused by light intensities during chamber construction when minor differences in water depth increase the photosynthetic rate of endosymbiotic diatoms during neap tides. The continuous reproduction rate of N . venosus throughout the year is increased in subtropical calms by higher summer temperatures and the marginal input of inorganic nutrients during rainy seasons. The expected life span of both gamonts and schizonts are 18 months.

Oxygen isotope compositions of fossil foraminifera tests are commonly used proxies for ocean paleotemperatures, with reconstructions spanning the last 112 million years. However, the isotopic composition of these calcitic tests can be substantially altered during diagenesis without discernible textural changes. Here, we investigate fluid-mediated isotopic exchange in pristine tests of three modern benthic foraminifera species ( Ammonia sp ., Haynesina germanica , and Amphistegina lessonii ) following immersion into an 18 O-enriched artificial seawater at 90 °C for hours to days. Reacted tests remain texturally pristine but their bulk oxygen isotope compositions reveal rapid and species-dependent isotopic exchange with the water. NanoSIMS imaging reveals the 3-dimensional intra-test distributions of 18 O-enrichment that correlates with test ultra-structure and associated organic matter. Image analysis is used to quantify species level differences in test ultrastructure, which explains the observed species-dependent rates of isotopic exchange. Consequently, even tests considered texturally pristine for paleo-climatic reconstruction purposes may have experienced substantial isotopic exchange; critical paleo-temperature record re-examination is warranted. Paleoclimate reconstructions commonly use oxygen isotope compositions from fossil foraminifera tests as proxies. Here, the authors show that these tests exchange O-isotopes with surrounding fluids, with implications for paleotemperature records.

The ocean—the Earth’s largest ecosystem—is increasingly affected by anthropogenic climate change 1 , 2 . Large and globally consistent shifts have been detected in species phenology, range extension and community composition in marine ecosystems 3 – 5 . However, despite evidence for ongoing change, it remains unknown whether marine ecosystems have entered an Anthropocene 6 state beyond the natural decadal to centennial variability. This is because most observational time series lack a long-term baseline, and the few time series that extend back into the pre-industrial era have limited spatial coverage 7 , 8 . Here we use the unique potential of the sedimentary record of planktonic foraminifera—ubiquitous marine zooplankton—to provide a global pre-industrial baseline for the composition of modern species communities. We use a global compilation of 3,774 seafloor-derived planktonic foraminifera communities of pre-industrial age 9 and compare these with communities from sediment-trap time series that have sampled plankton flux since ad 1978 (33 sites, 87 observation years). We find that the Anthropocene assemblages differ from their pre-industrial counterparts in proportion to the historical change in temperature. We observe community changes towards warmer or cooler compositions that are consistent with historical changes in temperature in 85% of the cases. These observations not only confirm the existing evidence for changes in marine zooplankton communities in historical times, but also demonstrate that Anthropocene communities of a globally distributed zooplankton group systematically differ from their unperturbed pre-industrial state. Seafloor-derived planktonic foraminifera communities of pre-industrial age are compared with communities from sediment-trap time series and show that Anthropocene communities of a globally distributed zooplankton group differ from their unperturbed pre-industrial state.

A total of 25 surface sediment samples collected along a North-South transect (from 9.69N to 80E and 55.01S to 40E) in south western Indian Ocean have been used to study recent planktic foraminifera revealing the ecological preferences of the planktic foraminifer species in the area, particularly the coiling direction patterns in the planktic species Neoglobo-quadrina pachyderma (Ehrenberg). The coiling direction and their absolute abundance can be shown to be related to ambient temperature and salinity values. Dextrally coiled forms decrease towards higher latitude, while sinistrally coiled forms increase. Similar studies should be undertaken along various transects to further establish the extent of environmental control on coiling direction in planktic foraminifera.

Photosymbiosis, a mode of mixotrophy by algal endosymbiosis, provides key advantages to pelagic life in oligotrophic oceans. Despite its ecological importance, mechanisms underlying its emergence and association with the evolutionary success of photosymbiotic lineages remain unclear. We used planktonic foraminifera, a group of pelagic test-forming protists with an excellent fossil record, to reveal the history of symbiont acquisition among their three main extant clades. We used single-cell 18S rRNA gene amplicon sequencing to reveal symbiont identity and mapped the symbiosis on a phylogeny time-calibrated by fossil data. We show that the highly specific symbiotic interaction with dinoflagellates emerged in the wake of a major extinction of symbiont-bearing taxa at the end of the Eocene. In contrast, less specific and low-light-adapted symbioses with pelagophytes emerged 20 million years later, in multiple independent lineages in the Late Neogene, at a time when the vertical structure of pelagic ecosystems was transformed by global cooling. We infer that in foraminifera, photosymbiosis can evolve easily and that its establishment leads to diversification and ecological dominance to such an extent, that the proliferation of new symbioses is prevented by the incumbent lineages.