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Acknowledgements: Acknowledgements: We thank D. H. Mann for his detailed and constructive comments; and T. Ager, J. Austin, T. B. Brand, A. Cooper, S. Funder, M. T. P. Gilbert, T. Jørgensen, N. J. Korsgaard, S. Liu, M. Meldgaard, P. V. S. Olsen, M. L. Siggaard-Andersen, J. Stenderup, S. A. Woodroffe and staff at the GeoGenetics Sequencing Core and National Park Service-Western Arctic National Parklands for help and support. E.W. and D.J.M. thank the staff at St. John’s College, Cambridge, for providing a stimulating environment for scientific discussion of the project. E.W. thanks Illumina for collaboration. The Lundbeck Foundation GeoGenetics Centre is supported by the Carlsberg Foundation (CF18-0024), the Lundbeck Foundation (R302-2018-2155), the Novo Nordisk Foundation (NNF18SA0035006), the Wellcome Trust (UNS69906) and GRF EXC CRS Chair (44113220)—Cluster of Excellence. The PhyloNorway plant genome database is part of the Norwegian Barcode of Life Network (https://www.norbol.org) funded by the Research Council of Norway (226134/F50), the Norwegian Biodiversity Information Centre (14-14, 70184209) and The Arctic University Museum of Norway. Metabarcoding sequencing was funded by the Central Public-Interest Scientific Institution Basal Research Fund, CAFS (2017B001 and 2020A001). B.D.S. is supported by the Wellcome Trust programme in Mathematical Genomics and Medicine (WT220023); F.R. by a Villum Fonden Young Investigator award (no. 00025300); D.J.M. by the Quest Archaeological Research Fund; P.M. by the Swedish Research Council (VR); R.D. by the Wellcome Trust (WT207492); and A.R. by a Marie Skłodowska-Curie Actions Individual Fellowship (MSCA-IF, 703542) and the Research Council of Norway (KLIMAFORSK, 294929). L.O. has received funding from the European Research Council (ERC) under the European Union’s Horizon 2020 research and innovation programme (no. 681605); I.G.A. and Y.L. from the ERC under the European Union’s Horizon 2020 research and innovation programme (no. 819192). J.I.S. and J.M. are supported by the Research Council of Norway. P.B.H. and N.R.E. acknowledge NERC funding (grant NE/P015093/1). D.W.B. was supported by a Marie Skłodowska-Curie Actions Incoming International Fellowship (MCIIF-40974). T.S.K. is funded by a Carlsberg Foundation Young Researcher Fellowship (CF19-0712)., During the last glacial-interglacial cycle, Arctic biotas experienced substantial climatic changes, yet the nature, extent and rate of their responses are not fully understood1-8. Here we report a large-scale environmental DNA metagenomic study of ancient plant and mammal communities, analysing 535 permafrost and lake sediment samples from across the Arctic spanning the past 50,000 years. Furthermore, we present 1,541 contemporary plant genome assemblies that were generated as reference sequences. Our study provides several insights into the long-term dynamics of the Arctic biota at the circumpolar and regional scales. Our key findings include: (1) a relatively homogeneous steppe-tundra flora dominated the Arctic during the Last Glacial Maximum, followed by regional divergence of vegetation during the Holocene epoch; (2) certain grazing animals consistently co-occurred in space and time; (3) humans appear to have been a minor factor in driving animal distributions; (4) higher effective precipitation, as well as an increase in the proportion of wetland plants, show negative effects on animal diversity; (5) the persistence of the steppe-tundra vegetation in northern Siberia enabled the late survival of several now-extinct megafauna species, including the woolly mammoth until 3.9 ± 0.2 thousand years ago (ka) and the woolly rhinoceros until 9.8 ± 0.2 ka; and (6) phylogenetic analysis of mammoth environmental DNA reveals a previously unsampled mitochondrial lineage. Our findings highlight the power of ancient environmental metagenomics analyses to advance understanding of population histories and long-term ecological dynamics.

A detailed, well-dated record of pollen and sedimentary ancient DNA (sedaDNA) for the period 15 000–9500 cal a bp describes changes at Lake Bolshoye Shchuchye in the Polar Ural Mountains, located far east of the classical Lateglacial sites in western Europe. Arctic tundra rapidly changed to lusher vegetation, possibly including both dwarf (Betula nana) and tree birch (B. pubescens), dated in our record to take place 14 565 cal a bp, coincident with the onset of the Bølling in western Europe; this was paralleled by increased summer temperatures. A striking feature is an early decline in Betula pollen and sedaDNA reads 300 years before the onset of the Younger Dryas (YD) in western Europe. Given the solid site chronology, this could indicate that the YD cooling started in Siberia and propagated westwards, or that the vegetation reacted to the inter-Allerød cooling at 13 100 cal a bp and did not recover during the late Allerød. During the YD, increases in steppe taxa such as Artemisia and Chenopodiaceae suggest drier conditions. At the onset of the Holocene, the vegetation around the lake reacted fast to the warmer conditions, as seen in the increase of arboreal taxa, especially Betula, and a decrease in herbs such as Artemisia and Cyperaceae. publishedVersion

The evolution of past global ice sheets is highly uncertain. One example is the missing ice problem during the Last Glacial Maximum (LGM, 26 000-19 000 years before present) – an apparent 8-28 m discrepancy between far-field sea level indicators and modelled sea level from ice sheet reconstructions. In the absence of ice sheet reconstructions, researchers often use marine δ18O proxy records to infer ice volume prior to the LGM. We present a global ice sheet reconstruction for the past 80 000 years, called PaleoMIST 1.0, constructed independently of far-field sea level and δ18O proxy records. Our reconstruction is compatible with LGM far-field sea-level records without requiring extra ice volume, thus solving the missing ice problem. However, for Marine Isotope Stage 3 (57 000-29 000 years before present) - a pre-LGM period - our reconstruction does not match proxy-based sea level reconstructions, indicating the relationship between marine δ18O and sea level may be more complex than assumed., The configuration of past ice sheets, and therefore sea level, is highly uncertain. Here, the authors provide a global reconstruction of ice sheets for the past 80,000 years that allows to test proxy based sea level reconstructions and helps to reconcile disagreements with sea level changes inferred from models.

A 24,000-year record of plant community dynamics, based on pollen and ancient DNA from the sediments (sedaDNA) of Lake Bolshoye Shchuchye in the Polar Ural Mountains, provides detailed information on the flora of the Last Glacial Maximum (LGM) and also changes in plant community composition and dominance. It greatly improves on incomplete records from short and fragmented stratigraphic sequences found in exposed sedimentary sections in the western Russian Arctic. In total, 162 plant taxa were detected by sedaDNA and 115 by pollen analysis. Several shifts in dominance between and within plant functional groups occurred over the studied period, but most taxa appear to have survived in situ. A diverse arctic-alpine herb flora characterised the interval ca. 24,000–17,000 cal years BP and persisted into the Holocene. Around 17,000 cal years BP, sedges (e.g. Carex) and bryophytes (e.g. Bryum, Aulacomnium) increased. The establishment of shrub-tundra communities of Dryas and Vaccinium sp., with potentially some Betula pubescens trees (influx ∼290 grains cm2 year−1), followed at ca. 15,000 cal years BP. Forest taxa such as Picea and ferns (e.g. Dryopteris fragrans, Gymnocarpium dryopteris) established near the lake from ca. 10,000 cal years BP, followed by the establishment of Larix trees from ca. 9000 cal years BP. Picea began to decline from ca. 7000 cal years BP. A complete withdrawal of forest tree taxa occurred by ca. 4000 cal years BP, presumably due to decreasing growing-season temperatures, allowing the expansion of dwarf-shrub tundra and a diverse herb community similar to the present-day vegetation mosaic. Contrary to some earlier comparative studies, sedaDNA and pollen from Lake Bolshoye Shchuchye showed high similarity in the timing of compositional changes and the occurrence of key plant taxa. The sedaDNA record revealed several features that the pollen stratigraphy and earlier palaeorecords in the region failed to detect; a sustained, long-term increase in floristic richness since the LGM until the early Holocene, turnover in grass and forb genera over the Pleistocene-Holocene transition, persistence of a diverse arctic-alpine flora over the late Quaternary, and a variable bryophyte flora through time. As pollen records are often limited by taxonomic resolution, differential productivity and dispersal, sedaDNA can provide improved estimates of floristic richness and is better able to distinguish between different plant assemblages. However, pollen remains superior at providing quantitative estimates of plant abundance changes and detecting several diverse groups (e.g. Poaceae, Cyperaceae, Asteraceae) which may be underreported in the sedaDNA. Joint use of the two proxies provided unprecedented floristic detail of past plant communities and helped to distinguish between long-distance transport of pollen and local presence, particularly for woody plant taxa.

The last deglaciation in northern Europe provides an opportunity to study the hydrologic component of abrupt climate shifts in a region with complex interactions between ice sheets and oceanic and atmospheric circulation. We use leaf wax hydrogen isotopes (δ2H) to reconstruct summer precipitation δ2H and aridity in southwestern Norway from 15.8 to 11.5 ka. We identify transitions to a more proximal moisture source before the ends of Heinrich Stadial 1 and the Younger Dryas, prior to local warming and increased primary productivity in both instances. We infer these changes in moisture delivery to southwestern Norway to be a response to northward shifts in the polar front caused by warm water intrusion into the North Atlantic, which preceded abrupt warming in the circum-North Atlantic. These results suggest that moisture transport pathways shift northward as warm surface ocean water reaches higher latitudes in the North Atlantic. publishedVersion

We reconstruct patterns of ice flow and retreat of the southwestern Scandinavian Ice Sheet, from 2900 field observations of glacial striae and elevation measurements of 60 ice-marginal-deltas from a high-resolution LiDAR DEM. During the Last Glacial Maximum, ice flow was towards the west across the entire area, including across several-hundred meter deep north-south oriented fjords. During deglaciation, ice flow adjusted to topography and the dominant flow direction switched towards the south-west. We use a shoreline diagram constructed from relative sea-level curves to establish the age of each delta, which allows us to constrain the timing of retreat with almost decadal precision. Rapid ice sheet retreat commenced at the onset of the Holocene at 11,600 cal years BP. Retreat rates were 160 m a −1 in the deepest fjords, 60–80 m a −1 in shallower fjords, and even slower for land-terminating margins. The fastest retreat rates, 240 m a −1 and 340 m a −1 , were experienced in the largest fjords, Hardangerfjorden and Sognefjorden, which border the study area to the south and north. Crosscutting glacial striae indicate that calving bays developed during retreat along the widest fjords. The combination of complex fjord topography with fast ice-margin retreat by iceberg calving, led to isolation of ice remnants on islands and peninsulas, a process that accelerated the overall rate of deglaciation. Ice-margin retreat paused between 11,300–11,100 cal years BP, probably due to cooling during the Preboreal Oscillation.

Marine-terminating glaciers and ice streams are important controls of ice sheet mass balance. However, understanding of their long-term response to external forcing is limited by relatively short observational records of present-day glaciers and sparse geologic evidence for paleo-glaciers. Here we use a high-resolution ice sheet model with an accurate representation of grounding line dynamics to study the deglaciation of the marine-based south-western Norwegian sector of the Scandinavian Ice Sheet and its sensitivity to ocean and atmosphere forcing. We find that the regional response to a uniform climate change is highly dependent on the local bedrock topography, consistent with ice sheet reconstructions. Our simulations suggest that ocean warming is able to trigger initial retreat in several fjords, but is not sufficient to explain retreat everywhere. Widespread retreat requires additional ice thinning driven by surface melt. Once retreat is triggered, the underlying bedrock topography and fjord width control the rate and extent of retreat, while multi-millennial changes over the course of deglaciation are modulated by surface melt. We suggest that fjord geometry, ice-ocean interactions and grounding line dynamics are vital controls of decadal-to centennial scale ice sheet mass loss. However, we postulate that atmospheric changes are the most important drivers of widespread ice sheet demise, and will likely trump oceanic influence on future ice sheet mass loss and resulting sea level rise over centennial and longer time scales. publishedVersion

Contrary to what Zwyns et al . claim on a bibliographical basis, the lithic industry of Byzovaya cannot belong to the Streletskayan complex or be considered as Upper Palaeolithic (UP). Direct analysis of northern assemblages and of Streletskayan technologies reveals incompatible features between these industries. Byzovaya is structured on specific Mousterian technologies and does not show any unique features of the UP.

During the Quaternary period, ice sheets centred over the Barents and Kara seas expanded several times onto mainland Russia and blocked northflowing rivers, such as the Yenissei, Ob, Pechora and Mezen. Large ice-dammed lakes with reversed outlets, e.g. toward the Caspian Sea, formed south of these ice sheets. Some lakes are reconstructed from shorelines and lacustrine sediments, others mainly from ice-sheet configuration. Ice-dammed lakes, considerably larger than any lake on Earth today, are reconstructed for the periods 90-80 and 60-50 ka. The ages are based on numerous optically stimulated luminescence (OSL) dates. During the global Last Glacial Maximum (LGM, about 20 ka) the Barents-Kara Ice Sheet was too small to block these eastern rivers, although in contrast to the 90-80 and 60-50 ka maxima, the Scandinavian Ice Sheet grew large enough to divert rivers and meltwater across the drainage divide from the Baltic Basin to the River Volga, and that way to the Caspian Sea. Climate modelling shows that the lakes caused lower summer temperatures on the continent and on the lower parts of the ice sheet. The final drainage of the best mapped lake is modelled, and it is concluded that it probably emptied within few months. We predict that this catastrophic outburst had considerable impact on sea-ice formation in the Arctic Ocean and on the climate of a much larger area. (C) 2003 Elsevier Ltd. All rights reserved. (Less)

Lake and peat deposits from the Timan Ridge, Arctic Russia, were pollen analysed, reconstructing the vegetation history and paleoenvironment since the Last Glacial Maximum (LGM) 20–18,000 years ago. The sites studied are located inside the margins of a large paleolake of about 20 km 2 , by us named Lake Timan. This lake developed in the Late Weichselian, more than 30,000 years after the deglaciation of this region, and was formed due to increased precipitation and warmer summers that accelerated the melting of stagnant ice within its catchment. The lake was drained during the early Holocene when the outlet rivers eroded the spillways. A new generation of much smaller lakes formed during the Holocene when the last remnants of buried glacier ice melted away causing the exposed floor of Lake Timan to subside. Since deglaciation, the following regional vegetation development has been recorded: (1) During the initial stage of Lake Timan, the dominant vegetation was discontinuous steppe/tundra, with patches of snow bed vegetation. (2) A dwarf-shrub tundra established during the Late Weichselian interstadial (Allerod), probably reflecting warmer and moister conditions. (3) The Younger Dryas cooling is recognised by a reversal to steppe/tundra and snowbeds on unstable mineral-soils, and higher palynological richness. (4) Soon after the transition into the Holocene, a birch-forest established on the Timan Ridge. (5) A cooling starting around 8200 cal. years BP initiated the deforestation of the exposed hills. In the most protected sites, birch trees persisted until later than 4000 years ago, reflecting a gradual development into the present treeless dwarf-shrub tundra.