ForCenS-LGM: a dataset of planktonic foraminifera species assemblage composition for the Last Glacial Maximum.

Species assemblage composition of marine microfossils offers the possibility to investigate ecological and climatological change on time scales inaccessible using conventional observations. Planktonic foraminifera - calcareous zooplankton - have an excellent fossil record and are used extensively in palaeoecology and palaeoceanography. During the Last Glacial Maximum (LGM; 19,000 - 23,000 years ago), the climate was in a radically different state. This period is therefore a key target to investigate climate and biodiversity under different conditions than today. Studying LGM climate and ecosystems indeed has a long history, yet the most recent global synthesis of planktonic foraminifera assemblage composition is now nearly two decades old. Here we present the ForCenS-LGM dataset with 2,365 species assemblage samples collected using standardised methods and with harmonised taxonomy. The data originate from marine sediments from 664 sites and present a more than 50% increase in coverage compared to previous work. The taxonomy is compatible with the most recent global core top dataset, enabling direct investigation of temporal changes in foraminifera biogeography and facilitating seawater temperature reconstructions.

The global genetic diversity of planktonic foraminifera reveals the structure of cryptic speciation in plankton.

The nature and extent of diversity in the plankton has fascinated scientists for over a century. Initially, the discovery of many new species in the remarkably uniform and unstructured pelagic environment appeared to challenge the concept of ecological niches. Later, it became obvious that only a fraction of plankton diversity had been formally described, because plankton assemblages are dominated by understudied eukaryotic lineages with small size that lack clearly distinguishable morphological features. The high diversity of the plankton has been confirmed by comprehensive metabarcoding surveys, but interpretation of the underlying molecular taxonomies is hindered by insufficient integration of genetic diversity with morphological taxonomy and ecological observations. Here we use planktonic foraminifera as a study model and reveal the full extent of their genetic diversity and investigate geographical and ecological patterns in their distribution. To this end, we assembled a global data set of ~7600 ribosomal DNA sequences obtained from morphologically characterised individual foraminifera, established a robust molecular taxonomic framework for the observed diversity, and used it to query a global metabarcoding data set covering ~1700 samples with ~2.48 billion reads. This allowed us to extract and assign 1 million reads, enabling characterisation of the structure of the genetic diversity of the group across ~1100 oceanic stations worldwide. Our sampling revealed the existence of, at most, 94 distinct molecular operational taxonomic units (MOTUs) at a level of divergence indicative of biological species. The genetic diversity only doubles the number of formally described species identified by morphological features. Furthermore, we observed that the allocation of genetic diversity to morphospecies is uneven. Only 16 morphospecies disguise evolutionarily significant genetic diversity, and the proportion of morphospecies that show genetic diversity increases poleward. Finally, we observe that MOTUs have a narrower geographic distribution than morphospecies and that in some cases the MOTUs belonging to the same morphospecies (cryptic species) have different environmental preferences. Overall, our analysis reveals that even in the light of global genetic sampling, planktonic foraminifera diversity is modest and finite. However, the extent and structure of the cryptic diversity reveals that genetic diversification is decoupled from morphological diversification, hinting at different mechanisms acting at different levels of divergence.

A multicentennial mode of North Atlantic climate variability throughout the Last Glacial Maximum.

Paleoclimate proxy records from the North Atlantic region reveal substantially greater multicentennial temperature variability during the Last Glacial Maximum (LGM) compared to the current interglacial. As there was no obvious change in external forcing, causes for the increased variability remain unknown. Exploiting LGM simulations with a comprehensive coupled climate model along with high-resolution proxy records, we introduce an oscillatory mode of multicentennial variability, which is associated with moderate variations in the Atlantic meridional overturning circulation and depends on the large-scale salinity distribution. This self-sustained mode is amplified by sea-ice feedbacks and induces maximum surface temperature variability in the subpolar North Atlantic region. Characterized by a distinct climatic imprint and different dynamics, the multicentennial oscillation has to be distinguished from Dansgaard-Oeschger variability and emerges only under full LGM climate forcing. The potential of multicentennial modes of variability to emerge or disappear in response to changing climate forcing may have implications for future climate change.

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.

Plankton response to global warming is characterized by non-uniform shifts in assemblage composition since the last ice age.

Biodiversity is expected to change in response to future global warming. However, it is difficult to predict how species will track the ongoing climate change. Here we use the fossil record of planktonic foraminifera to assess how biodiversity responded to climate change with a magnitude comparable to future anthropogenic warming. We compiled time series of planktonic foraminifera assemblages, covering the time from the last ice age across the deglaciation to the current warm period. Planktonic foraminifera assemblages shifted immediately when temperature began to rise at the end of the last ice age and continued to change until approximately 5,000 years ago, even though global temperature remained relatively stable during the last 11,000 years. The biotic response was largest in the mid latitudes and dominated by range expansion, which resulted in the emergence of new assemblages without analogues in the glacial ocean. Our results indicate that the plankton response to global warming was spatially heterogeneous and did not track temperature change uniformly over the past 24,000 years. Climate change led to the establishment of new assemblages and possibly new ecological interactions, which suggests that current anthropogenic warming may lead to new, different plankton community composition.

Complex spatio-temporal structure of the Holocene Thermal Maximum.

Inconsistencies between Holocene climate reconstructions and numerical model simulations question the robustness of climate models and proxy temperature records. Climate reconstructions suggest an early-middle Holocene Thermal Maximum (HTM) followed by gradual cooling, whereas climate models indicate continuous warming. This discrepancy either implies seasonal biases in proxy-based climate reconstructions, or that the climate model sensitivity to forcings and feedbacks needs to be reevaluated. Here, we analyze a global database of Holocene paleotemperature records to investigate the spatiotemporal structure of the HTM. Continental proxy records at mid and high latitudes of the Northern Hemisphere portray a "classic" HTM (8-4 ka). In contrast, marine proxy records from the same latitudes reveal an earlier HTM (11-7ka), while a clear temperature anomaly is missing in the tropics. The results indicate a heterogeneous response to climate forcing and highlight the lack of globally synchronous HTM.

Publisher Correction: A global database of Holocene paleotemperature records.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Author Correction: A global database of Holocene paleotemperature records.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Publisher Correction: A global database of Holocene paleotemperature records.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

A global database of Holocene paleotemperature records.

A comprehensive database of paleoclimate records is needed to place recent warming into the longer-term context of natural climate variability. We present a global compilation of quality-controlled, published, temperature-sensitive proxy records extending back 12,000 years through the Holocene. Data were compiled from 679 sites where time series cover at least 4000 years, are resolved at sub-millennial scale (median spacing of 400 years or finer) and have at least one age control point every 3000 years, with cut-off values slackened in data-sparse regions. The data derive from lake sediment (51%), marine sediment (31%), peat (11%), glacier ice (3%), and other natural archives. The database contains 1319 records, including 157 from the Southern Hemisphere. The multi-proxy database comprises paleotemperature time series based on ecological assemblages, as well as biophysical and geochemical indicators that reflect mean annual or seasonal temperatures, as encoded in the database. This database can be used to reconstruct the spatiotemporal evolution of Holocene temperature at global to regional scales, and is publicly available in Linked Paleo Data (LiPD) format.

Genetic and morphological divergence in the warm-water planktonic foraminifera genus Globigerinoides.

The planktonic foraminifera genus Globigerinoides provides a prime example of a species-rich genus in which genetic and morphological divergence are uncorrelated. To shed light on the evolutionary processes that lead to the present-day diversity of Globigerinoides, we investigated the genetic, ecological and morphological divergence of its constituent species. We assembled a global collection of single-cell barcode sequences and show that the genus consists of eight distinct genetic types organized in five extant morphospecies. Based on morphological evidence, we reassign the species Globoturborotalita tenella to Globigerinoides and amend Globigerinoides ruber by formally proposing two new subspecies, G. ruber albus n.subsp. and G. ruber ruber in order to express their subspecies level distinction and to replace the informal G. ruber "white" and G. ruber "pink", respectively. The genetic types within G. ruber and Globigerinoides elongatus show a combination of endemism and coexistence, with little evidence for ecological differentiation. CT-scanning and ontogeny analysis reveal that the diagnostic differences in adult morphologies could be explained by alterations of the ontogenetic trajectories towards final (reproductive) size. This indicates that heterochrony may have caused the observed decoupling between genetic and morphological diversification within the genus. We find little evidence for environmental forcing of either the genetic or the morphological diversification, which allude to biotic interactions such as symbiosis, as the driver of speciation in Globigerinoides.

Global change drives modern plankton communities away from the pre-industrial state.

The ocean-the Earth's largest ecosystem-is increasingly affected by anthropogenic climate change1,2. Large and globally consistent shifts have been detected in species phenology, range extension and community composition in marine ecosystems3-5. However, despite evidence for ongoing change, it remains unknown whether marine ecosystems have entered an Anthropocene6 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 coverage7,8. 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 age9 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.

Icebergs not the trigger for North Atlantic cold events.

Abrupt climate change is a ubiquitous feature of the Late Pleistocene epoch. In particular, the sequence of Dansgaard-Oeschger events (repeated transitions between warm interstadial and cold stadial conditions), as recorded by ice cores in Greenland, are thought to be linked to changes in the mode of overturning circulation in the Atlantic Ocean. Moreover, the observed correspondence between North Atlantic cold events and increased iceberg calving and dispersal from ice sheets surrounding the North Atlantic has inspired many ocean and climate modelling studies that make use of freshwater forcing scenarios to simulate abrupt change across the North Atlantic region and beyond. On the other hand, previous studies identified an apparent lag between North Atlantic cooling events and the appearance of ice-rafted debris over the last glacial cycle, leading to the hypothesis that iceberg discharge may be a consequence of stadial conditions rather than the cause. Here we further establish this relationship and demonstrate a systematic delay between pronounced surface cooling and the arrival of ice-rafted debris at a site southwest of Iceland over the past four glacial cycles, implying that in general icebergs arrived too late to have triggered cooling. Instead we suggest that--on the basis of our comparisons of ice-rafted debris and polar planktonic foraminifera--abrupt transitions to stadial conditions should be considered as a nonlinear response to more gradual cooling across the North Atlantic. Although the freshwater derived from melting icebergs may provide a positive feedback for enhancing and or prolonging stadial conditions, it does not trigger northern stadial events.