A 2-million-year-old ecosystem in Greenland uncovered by environmental DNA.

Late Pliocene and Early Pleistocene epochs 3.6 to 0.8 million years ago1 had climates resembling those forecasted under future warming2. Palaeoclimatic records show strong polar amplification with mean annual temperatures of 11-19 °C above contemporary values3,4. The biological communities inhabiting the Arctic during this time remain poorly known because fossils are rare5. Here we report an ancient environmental DNA6 (eDNA) record describing the rich plant and animal assemblages of the Kap København Formation in North Greenland, dated to around two million years ago. The record shows an open boreal forest ecosystem with mixed vegetation of poplar, birch and thuja trees, as well as a variety of Arctic and boreal shrubs and herbs, many of which had not previously been detected at the site from macrofossil and pollen records. The DNA record confirms the presence of hare and mitochondrial DNA from animals including mastodons, reindeer, rodents and geese, all ancestral to their present-day and late Pleistocene relatives. The presence of marine species including horseshoe crab and green algae support a warmer climate than today. The reconstructed ecosystem has no modern analogue. The survival of such ancient eDNA probably relates to its binding to mineral surfaces. Our findings open new areas of genetic research, demonstrating that it is possible to track the ecology and evolution of biological communities from two million years ago using ancient eDNA.

Divergence of Arctic shrub growth associated with sea ice decline.

Arctic sea ice extent (SIE) is declining at an accelerating rate with a wide range of ecological consequences. However, determining sea ice effects on tundra vegetation remains a challenge. In this study, we examined the universality or lack thereof in tundra shrub growth responses to changes in SIE and summer climate across the Pan-Arctic, taking advantage of 23 tundra shrub-ring chronologies from 19 widely distributed sites (56°N to 83°N). We show a clear divergence in shrub growth responses to SIE that began in the mid-1990s, with 39% of the chronologies showing declines and 57% showing increases in radial growth (decreasers and increasers, respectively). Structural equation models revealed that declining SIE was associated with rising air temperature and precipitation for increasers and with increasingly dry conditions for decreasers. Decreasers tended to be from areas of the Arctic with lower summer precipitation and their growth decline was related to decreases in the standardized precipitation evapotranspiration index. Our findings suggest that moisture limitation, associated with declining SIE, might inhibit the positive effects of warming on shrub growth over a considerable part of the terrestrial Arctic, thereby complicating predictions of vegetation change and future tundra productivity.

Common mechanisms explain nitrogen-dependent growth of Arctic shrubs over three decades despite heterogeneous trends and declines in soil nitrogen availability.

Heterogeneity has been observed in the responses of Arctic shrubs to climate variability over recent decades, which may reflect landscape-scale variability in belowground resources. At a northern fringe of tall shrub expansion (Yuribei, Yamal Peninsula, Russia), we sought to determine the mechanisms relating nitrogen (N) limitation to shrub growth over decadal time. We analysed the ratio of 15 N to 14 N isotopes in wood rings of 10 Salix lanata individuals (399 measurements) to reconstruct annual point-based bioavailable N between 1980 and 2013. We applied a model-fitting/model-selection approach with a suite of competing ecological models to assess the most-likely mechanisms that explain each shrub's individual time-series. Shrub δ15 N time-series indicated declining (seven shrubs), increasing (two shrubs) and no trend (one shrub) in N availability. The most appropriate model for all shrubs included N-dependent growth of linear rather than saturating form. Inclusion of plant-soil feedbacks better explained ring width and δ15 N for eight of 10 individuals. Although N trajectories were individualistic, common mechanisms of varying strength confirmed the N-dependency of shrub growth. The linear mechanism may reflect intense scavenging of scarce N; the importance of plant-soil feedbacks suggests that shrubs subvert the microbial bottleneck by actively controlling their environment.

Sensitivity of global terrestrial ecosystems to climate variability.

The identification of properties that contribute to the persistence and resilience of ecosystems despite climate change constitutes a research priority of global relevance. Here we present a novel, empirical approach to assess the relative sensitivity of ecosystems to climate variability, one property of resilience that builds on theoretical modelling work recognizing that systems closer to critical thresholds respond more sensitively to external perturbations. We develop a new metric, the vegetation sensitivity index, that identifies areas sensitive to climate variability over the past 14 years. The metric uses time series data derived from the moderate-resolution imaging spectroradiometer (MODIS) enhanced vegetation index, and three climatic variables that drive vegetation productivity (air temperature, water availability and cloud cover). Underlying the analysis is an autoregressive modelling approach used to identify climate drivers of vegetation productivity on monthly timescales, in addition to regions with memory effects and reduced response rates to external forcing. We find ecologically sensitive regions with amplified responses to climate variability in the Arctic tundra, parts of the boreal forest belt, the tropical rainforest, alpine regions worldwide, steppe and prairie regions of central Asia and North and South America, the Caatinga deciduous forest in eastern South America, and eastern areas of Australia. Our study provides a quantitative methodology for assessing the relative response rate of ecosystems--be they natural or with a strong anthropogenic signature--to environmental variability, which is the first step towards addressing why some regions appear to be more sensitive than others, and what impact this has on the resilience of ecosystem service provision and human well-being.

Community Assembly and Climate Mismatch in Late Quaternary Eastern North American Pollen Assemblages.

Plant community response to climate change ranges from synchronous tracking to strong mismatch. Explaining this variation in climate change response is critical for accurate global change modeling. Here we quantify how closely assemblages track changes in climate (match/mismatch) and how broadly climate niches are spread within assemblages (narrow/broad ecological tolerance, or "filtering") using data for the past 21,000 years for 531 eastern North American fossil pollen assemblages. Although climate matching has been strong over the last 21 millennia, mismatch increased in 30% of assemblages during the rapid climate shifts between 14.5 and 10 ka. Assemblage matching rebounded toward the present day in 10%-20% of assemblages. Climate-assemblage mismatch was greater in tree-dominated and high-latitude assemblages, consistent with persisting populations, slower dispersal rates, and glacial retreat. In contrast, climate matching was greater for assemblages comprising taxa with higher median seed mass. More than half of the assemblages were climatically filtered at any given time, with peak filtering occurring at 8.5 ka for nearly 80% of assemblages. Thus, vegetation assemblages have highly variable rates of climate mismatch and filtering over millennial scales. These climate responses can be partially predicted by species' traits and life histories. These findings help constrain predictions for plant community response to contemporary climate change.

Changes in timing of seasonal peak photosynthetic activity in northern ecosystems.

Seasonality in photosynthetic activity is a critical component of seasonal carbon, water, and energy cycles in the Earth system. This characteristic is a consequence of plant's adaptive evolutionary processes to a given set of environmental conditions. Changing climate in northern lands (>30°N) alters the state of climatic constraints on plant growth, and therefore, changes in the seasonality and carbon accumulation are anticipated. However, how photosynthetic seasonality evolved to its current state, and what role climatic constraints and their variability played in this process and ultimately in carbon cycle is still poorly understood due to its complexity. Here, we take the "laws of minimum" as a basis and introduce a new framework where the timing (day of year) of peak photosynthetic activity (DOYPmax ) acts as a proxy for plant's adaptive state to climatic constraints on its growth. Our analyses confirm that spatial variations in DOYPmax reflect spatial gradients in climatic constraints as well as seasonal maximum and total productivity. We find a widespread warming-induced advance in DOYPmax (-1.66 ± 0.30 days/decade, p < 0.001) across northern lands, indicating a spatiotemporal dynamism of climatic constraints to plant growth. We show that the observed changes in DOYPmax are associated with an increase in total gross primary productivity through enhanced carbon assimilation early in the growing season, which leads to an earlier phase shift in land-atmosphere carbon fluxes and an increase in their amplitude. Such changes are expected to continue in the future based on our analysis of earth system model projections. Our study provides a simplified, yet realistic framework based on first principles for the complex mechanisms by which various climatic factors constrain plant growth in northern ecosystems.

Effects of sea ice on Arctic biota: an emerging crisis discipline.

The rapid decline in Arctic sea ice (ASI) extent, area and volume during recent decades is occurring before we can understand many of the mechanisms through which ASI interacts with biological processes both at sea and on land. As a consequence, our ability to predict and manage the effects of this enormous environmental change is limited, making this a crisis discipline Here, we propose a framework to study these effects, defining direct effects as those acting on life-history events of Arctic biota, and indirect effects, where ASI acts upon biological systems through chains of events, normally involving other components of the physical system and/or biotic interactions. Given the breadth and complexity of ASI's effects on Arctic biota, Arctic research requires a truly multidisciplinary approach to address this issue. In the absence of effective global efforts to tackle anthropogenic global warming, ASI will likely continue to decrease, compromising the conservation of many ASI-related taxonomic groups and ecosystems. Mitigation actions will rely heavily on the knowledge acquired on the mechanisms and components involved with the biological effects of ASI.

Predictability in community dynamics.

The coupling between community composition and climate change spans a gradient from no lags to strong lags. The no-lag hypothesis is the foundation of many ecophysiological models, correlative species distribution modelling and climate reconstruction approaches. Simple lag hypotheses have become prominent in disequilibrium ecology, proposing that communities track climate change following a fixed function or with a time delay. However, more complex dynamics are possible and may lead to memory effects and alternate unstable states. We develop graphical and analytic methods for assessing these scenarios and show that these dynamics can appear in even simple models. The overall implications are that (1) complex community dynamics may be common and (2) detailed knowledge of past climate change and community states will often be necessary yet sometimes insufficient to make predictions of a community's future state.

Sea ice, rain-on-snow and tundra reindeer nomadism in Arctic Russia.

Sea ice loss is accelerating in the Barents and Kara Seas (BKS). Assessing potential linkages between sea ice retreat/thinning and the region's ancient and unique social-ecological systems is a pressing task. Tundra nomadism remains a vitally important livelihood for indigenous Nenets and their large reindeer herds. Warming summer air temperatures have been linked to more frequent and sustained summer high-pressure systems over West Siberia, Russia, but not to sea ice retreat. At the same time, autumn/winter rain-on-snow (ROS) events have become more frequent and intense. Here, we review evidence for autumn atmospheric warming and precipitation increases over Arctic coastal lands in proximity to BKS ice loss. Two major ROS events during November 2006 and 2013 led to massive winter reindeer mortality episodes on the Yamal Peninsula. Fieldwork with migratory herders has revealed that the ecological and socio-economic impacts from the catastrophic 2013 event will unfold for years to come. The suggested link between sea ice loss, more frequent and intense ROS events and high reindeer mortality has serious implications for the future of tundra Nenets nomadism.

Warming-induced upslope advance of subalpine forest is severely limited by geomorphic processes.

Forests are expected to expand into alpine areas because of climate warming, causing land-cover change and fragmentation of alpine habitats. However, this expansion will only occur if the present upper treeline is limited by low-growing season temperatures that reduce plant growth. This temperature limitation has not been quantified at a landscape scale. Here, we show that temperature alone cannot realistically explain high-elevation tree cover over a >100-km(2) area in the Canadian Rockies and that geologic/geomorphic processes are fundamental to understanding the heterogeneous landscape distribution of trees. Furthermore, upslope tree advance in a warmer scenario will be severely limited by availability of sites with adequate geomorphic/topographic characteristics. Our results imply that landscape-to-regional scale projections of warming-induced, high-elevation forest advance into alpine areas should not be based solely on temperature-sensitive, site-specific upper-treeline studies but also on geomorphic processes that control tree occurrence at long (centuries/millennia) timescales.

Remote assessment of locally important ecological features across landscapes: how representative of reality?

The local ecological footprinting tool (LEFT) uses globally available databases, modeling, and algorithms to, remotely assess locally important ecological features across landscapes based on five criteria: biodiversity (beta-diversity), vulnerability (threatened species), fragmentation, connectivity, and resilience. This approach can be applied to terrestrial landscapes at a 300-m resolution within a given target area. Input is minimal (latitude and longitude) and output is a computer-generated report and series of maps that both individually and synthetically depict the relative value of each ecological criteria. A key question for any such tool, however, is how representative is the remotely obtained output compared to what is on the ground. Here, we present the results from comparing remotely- vs. field-generated outputs from the LEFT tool on two distinct study areas for beta-diversity and distribution of threatened species (vulnerability), the two fields computed by LEFT for which such an approach is feasible. The comparison method consists of a multivariate measure of similarity between two fields based on discrete wavelet transforms, and reveals consistent agreement across a wide range of spatial scales. These results suggest that remote assessment tools such as LEFT hold great potential for determining key ecological features across landscapes and for being utilized in preplanning biodiversity assessment tools.

Greening of Svalbard.

Svalbard, located between 76°30'N and 80°50'N, is among the regions in the world with the most rapid temperature increase. We processed a cloud-free time-series of MODIS-NDVI for Svalbard. The dataset is interpolated to daily data during the 2000-2022 period with 232 m pixel resolution. The onset of growth, with a clear phenological definition, has been mapped each year. Then the integrated NDVI from the onset (O) of growth each year to the time of average (2000-2022) peak (P) of growth (OP NDVI) have been calculated. OP NDVI has previously shown high correlation with field-based tundra productivity. Daily mean temperature data from 11 meteorological stations are compared with the NDVI data. The OP NDVI values show very high and significant correlation with growing degree days computed from onset to time of peak of growth for all the meteorological stations used. On average for the entire Svalbard, the year 2016 first had the highest greening (OP NDVI values) recorded since the year 2000, then the greening in 2018 surpassed 2016, then 2020 surpassed 2018, and finally 2022 was the year with the overall highest greening by far for the whole 2000-2022 period. This shows a rapid recent greening of Svalbard very strongly linked to temperature increase, although there are regional differences: the eastern parts of Svalbard show the largest variability between years, most likely due to variability in the timing of sea-ice break-up in adjacent areas. Finally, we find that areas dominated by manured moss-tundra in the polar desert zone require new methodologies, as moss does not share the seasonal NDVI dynamics of tundra communities.

Large herbivore diversity slows sea ice-associated decline in arctic tundra diversity.

Biodiversity is declining globally in response to multiple human stressors, including climate forcing. Nonetheless, local diversity trends are inconsistent in some taxa, obscuring contributions of local processes to global patterns. Arctic tundra diversity, including plants, fungi, and lichens, declined during a 15-year experiment that combined warming with exclusion of large herbivores known to influence tundra vegetation composition. Tundra diversity declined regardless of experimental treatment, as background growing season temperatures rose with sea ice loss. However, diversity declined slower with large herbivores than without them. This difference was associated with an increase in effective diversity of large herbivores as formerly abundant caribou declined and muskoxen increased. Efforts that promote herbivore diversity, such as rewilding, may help mitigate impacts of warming on tundra diversity.

Plant traits and associated data from a warming experiment, a seabird colony, and along elevation in Svalbard.

The Arctic is warming at a rate four times the global average, while also being exposed to other global environmental changes, resulting in widespread vegetation and ecosystem change. Integrating functional trait-based approaches with multi-level vegetation, ecosystem, and landscape data enables a holistic understanding of the drivers and consequences of these changes. In two High Arctic study systems near Longyearbyen, Svalbard, a 20-year ITEX warming experiment and elevational gradients with and without nutrient input from nesting seabirds, we collected data on vegetation composition and structure, plant functional traits, ecosystem fluxes, multispectral remote sensing, and microclimate. The dataset contains 1,962 plant records and 16,160 trait measurements from 34 vascular plant taxa, for 9 of which these are the first published trait data. By integrating these comprehensive data, we bridge knowledge gaps and expand trait data coverage, including on intraspecific trait variation. These data can offer insights into ecosystem functioning and provide baselines to assess climate and environmental change impacts. Such knowledge is crucial for effective conservation and management in these vulnerable regions.

The role of large wild animals in climate change mitigation and adaptation.

Two major environmental challenges of our time are responding to climate change and reversing biodiversity decline. Interventions that simultaneously tackle both challenges are highly desirable. To date, most studies aiming to find synergistic interventions for these two challenges have focused on protecting or restoring vegetation and soils but overlooked how conservation or restoration of large wild animals might influence the climate mitigation and adaptation potential of ecosystems. However, interactions between large animal conservation and climate change goals may not always be positive. Here, we review wildlife conservation and climate change mitigation in terrestrial and marine ecosystems. We elucidate general principles about the biome types where, and mechanisms by which, positive synergies and negative trade-offs between wildlife conservation and climate change mitigation are likely. We find that large animals have the greatest potential to facilitate climate change mitigation at a global scale via three mechanisms: changes in fire regime, especially in previously low-flammability biomes with a new or intensifying fire regime, such as mesic grasslands or warm temperate woodlands; changes in terrestrial albedo, particularly where there is potential to shift from closed canopy to open canopy systems at higher latitudes; and increases in vegetation and soil carbon stocks, especially through a shift towards below-ground carbon pools in temperate, tropical and sub-tropical grassland ecosystems. Large animals also contribute to ecosystem adaptation to climate change by promoting complexity of trophic webs, increasing habitat heterogeneity, enhancing plant dispersal, increasing resistance to abrupt ecosystem change and through microclimate modification.

Pleistocene Arctic megafaunal ecological engineering as a natural climate solution?

Natural climate solutions (NCS) in the Arctic hold the potential to be implemented at a scale able to substantially affect the global climate. The strong feedbacks between carbon-rich permafrost, climate and herbivory suggest an NCS consisting of reverting the current wet/moist moss and shrub-dominated tundra and the sparse forest-tundra ecotone to grassland through a guild of large herbivores. Grassland-dominated systems might delay permafrost thaw and reduce carbon emissions-especially in Yedoma regions, while increasing carbon capture through increased productivity and grass and forb deep root systems. Here we review the environmental context of megafaunal ecological engineering in the Arctic; explore the mechanisms through which it can help mitigate climate change; and estimate its potential-based on bison and horse, with the aim of evaluating the feasibility of generating an ecosystem shift that is economically viable in terms of carbon benefits and of sufficient scale to play a significant role in global climate change mitigation. Assuming a megafaunal-driven ecosystem shift we find support for a megafauna-based arctic NCS yielding substantial income in carbon markets. However, scaling up such projects to have a significant effect on the global climate is challenging given the large number of animals required over a short period of time. A first-cut business plan is presented based on practical information-costs and infrastructure-from Pleistocene Park (northeastern Yakutia, Russia). A 10 yr experimental phase incorporating three separate introductions of herds of approximately 1000 individuals each is costed at US$114 million, with potential returns of approximately 0.3-0.4% yr-1 towards the end of the period, and greater than 1% yr-1 after it. Institutional friction and the potential role of new technologies in the reintroductions are discussed. This article is part of the theme issue 'Climate change and ecosystems: threats, opportunities and solutions'.

Summer warming explains widespread but not uniform greening in the Arctic tundra biome.

Arctic warming can influence tundra ecosystem function with consequences for climate feedbacks, wildlife and human communities. Yet ecological change across the Arctic tundra biome remains poorly quantified due to field measurement limitations and reliance on coarse-resolution satellite data. Here, we assess decadal changes in Arctic tundra greenness using time series from the 30 m resolution Landsat satellites. From 1985 to 2016 tundra greenness increased (greening) at ~37.3% of sampling sites and decreased (browning) at ~4.7% of sampling sites. Greening occurred most often at warm sampling sites with increased summer air temperature, soil temperature, and soil moisture, while browning occurred most often at cold sampling sites that cooled and dried. Tundra greenness was positively correlated with graminoid, shrub, and ecosystem productivity measured at field sites. Our results support the hypothesis that summer warming stimulated plant productivity across much, but not all, of the Arctic tundra biome during recent decades.