Plant functional trait change across a warming tundra biomeThe tundra is warming more rapidly than any other biome on Earth, and the potential ramifications are far-reaching because of global feedback effects between vegetation and climate. A better understanding of how environmental factors shape plant structure and function is crucial for predicting the consequences of environmental change for ecosystem functioning. Here we explore the biome-wide relationships between temperature, moisture and seven key plant functional traits both across space and over three decades of warming at 117 tundra locations. Spatial temperature-trait relationships were generally strong but soil moisture had a marked influence on the strength and direction of these relationships, highlighting the potentially important influence of changes in water availability on future trait shifts in tundra plant communities. Community height increased with warming across all sites over the past three decades, but other traits lagged far behind predicted rates of change. Our findings highlight the challenge of using space-for-time substitution to predict the functional consequences of future warming and suggest that functions that are tied closely to plant height will experience the most rapid change. They also reveal the strength with which environmental factors shape biotic communities at the coldest extremes of the planet and will help to improve projections of functional changes in tundra ecosystems with climate warming. Environment-trait relationships across the tundra biomeWe found strong spatial associations between temperature and community height, SLA and LDMC (Fig. 2a, Extended Data Fig. 2 and Supplementary Table 3) across the 117 survey sites. Both height and SLA increased with summer temperature, but the temperaturetrait relationship for SLA was much stronger at wetter than at drier sites. LDMC was negatively related to temperature, and

Tundra ecosystems are vulnerable to hydrocarbon development, in part because small-scale, low-intensity disturbances can affect vegetation, permafrost soils, and wildlife out of proportion to their spatial extent. Scaling up to include human residents, tightly integrated arctic social-ecological systems (SESs) are believed similarly susceptible to industrial impacts and climate change. In contrast to northern Alaska and Canada, most terrestrial and aquatic components of West Siberian oil and gas fields are seasonally exploited by migratory herders, hunters, fishers, and domesticated reindeer (Rangifer tarandus L.). Despite anthropogenic fragmentation and transformation of a large proportion of the environment, recent socioeconomic upheaval, and pronounced climate warming, we find the Yamal-Nenets SES highly resilient according to a few key measures. We detail the remarkable extent to which the system has successfully reorganized in response to recent shocks and evaluate the limits of the system's capacity to respond. Our analytical approach combines quantitative methods with participant observation to understand the overall effects of rapid land use and climate change at the level of the entire Yamal system, detect thresholds crossed using surrogates, and identify potential traps. Institutional constraints and drivers were as important as the documented ecological changes. Particularly crucial to success is the unfettered movement of people and animals in space and time, which allows them to alternately avoid or exploit a wide range of natural and anthropogenic habitats. However, expansion of infrastructure, concomitant terrestrial and freshwater ecosystem degradation, climate change, and a massive influx of workers underway present a looming threat to future resilience. tundra disturbance ͉ Rangifer tarandus ͉ reindeer nomadism ͉ oil and gas activities ͉ remote sensing

Climate warming is altering the diversity of plant communities but it remains unknown which species will be lost or gained under warming, especially considering interactions with other factors such as herbivory and nutrient availability. Here, we experimentally test effects of warming, mammalian herbivory and fertilization on tundra species richness and investigate how plant functional traits affect losses and gains. We show that herbivory reverses the impact of warming on diversity: in the presence of herbivores warming increases species richness through higher species gains and lower losses, while in the absence of herbivores warming causes higher species losses and thus decreases species richness. Herbivores promote gains of short-statured species under warming, while herbivore removal and fertilization increase losses of short-statured and resource-conservative species through light limitation. Our results demonstrate that both rarity and traits forecast species losses and gains, and mammalian herbivores are essential for preventing trait-dependent extinctions and mitigate diversity loss under warming and eutrophication.

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Summary1. Warm-adapted low elevation plants are expected to exhibit considerable range shifts to higher altitudes and latitudes as a result of climate warming and increased nutrient loads. However, empirical studies show that the magnitude and direction of plant responses are highly species-and site-specific, suggesting that several additional drivers interact with warmer climate. 2. We experimentally tested the interactive effects of climate warming, mammalian herbivory and soil fertility on low elevation plants. Seedlings of three warm-adapted lowland forbs (Epilobium angustifolium, Silene dioica and Solidago virgaurea) were transplanted to an open tundra site with native mountain tundra vegetation, and the effects of full factorial combinations of herbivore exclosures, warming and fertilization on transplant survival, growth and flowering were studied for two growing seasons. We also investigated the response of native vegetation biomass to the same treatments and compared it with the responses of transplanted lowland forbs. 3. Effects of both warming and fertilization on the transplanted lowland forbs strongly hinged on herbivore exclusion, resulting in 2-13-fold increase in biomass in warmed and fertilized plots without herbivores compared with warmed and fertilized plots with herbivores present, the magnitude depending on the species. While warm-adapted transplants benefited from warming, the native tundra plant community biomass did not respond to warming treatment. 4. Our results show that grazing limits the growth of transplants under warmer and more productive conditions, indicating that the expansion of lowland plant species to higher altitudes with warming may be hampered by mammalian herbivory. Furthermore, our results also suggest that migration of warm-adapted species into lightly grazed high-altitude tundra ecosystems might transform these communities to be more responsive to warmer climate and nutrient loads. Studies that do not consider species' upward shifts from lower altitudes might thus have underestimated vegetation responses to global warming, as well as the potential of herbivory to influence these responses.

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Motivation The Tundra Trait Team (TTT) database includes field‐based measurements of key traits related to plant form and function at multiple sites across the tundra biome. This dataset can be used to address theoretical questions about plant strategy and trade‐offs, trait–environment relationships and environmental filtering, and trait variation across spatial scales, to validate satellite data, and to inform Earth system model parameters. Main types of variable contained The database contains 91,970 measurements of 18 plant traits. The most frequently measured traits (> 1,000 observations each) include plant height, leaf area, specific leaf area, leaf fresh and dry mass, leaf dry matter content, leaf nitrogen, carbon and phosphorus content, leaf C:N and N:P, seed mass, and stem specific density. Spatial location and grain Measurements were collected in tundra habitats in both the Northern and Southern Hemispheres, including Arctic sites in Alaska, Canada, Greenland, Fennoscandia and Siberia, alpine sites in the European Alps, Colorado Rockies, Caucasus, Ural Mountains, Pyrenees, Australian Alps, and Central Otago Mountains (New Zealand), and sub‐Antarctic Marion Island. More than 99% of observations are georeferenced. Time period and grain All data were collected between 1964 and 2018. A small number of sites have repeated trait measurements at two or more time periods. Major taxa and level of measurement Trait measurements were made on 978 terrestrial vascular plant species growing in tundra habitats. Most observations are on individuals (86%), while the remainder represent plot or site means or maximums per species. Software format csv file and GitHub repository with data cleaning scripts in R; contribution to TRY plant trait database (www.try-db.org) to be included in the next version release.

The causes of a greening trend detected in the Arctic using the normalized difference vegetation index (NDVI) are still poorly understood. Changes in NDVI are a result of multiple ecological and social factors that affect tundra net primary productivity. Here we use a 25 year time series of AVHRR-derived NDVI data (AVHRR: advanced very high resolution radiometer), climate analysis, a global geographic information database and ground-based studies to examine the spatial and temporal patterns of vegetation greenness on the Yamal Peninsula, Russia. We assess the effects of climate change, gas-field development, reindeer grazing and permafrost degradation. In contrast to the case for Arctic North America, there has not been a significant trend in summer temperature or NDVI, and much of the pattern of NDVI in this region is due to disturbances. There has been a 37% change in early-summer coastal sea-ice concentration, a 4% increase in summer land temperatures and a 7% change in the average time-integrated NDVI over the length of the satellite observations. Gas-field infrastructure is not currently extensive enough to affect regional NDVI patterns. The effect of reindeer is difficult to quantitatively assess because of the lack of control areas where reindeer are excluded. Many of the greenest landscapes on the Yamal are associated with landslides and drainage networks that have resulted from ongoing rapid permafrost degradation. A warming climate and enhanced winter snow are likely to exacerbate positive feedbacks between climate and permafrost thawing. We present a diagram that summarizes the social and ecological factors that influence Arctic NDVI. The NDVI should be viewed as a powerful monitoring tool that integrates the cumulative effect of a multitude of factors affecting Arctic land-cover change. ReferencesBelopukova E B, Leibman M O and Tukacheva L A 1989 The effect of climatic parameters on ground temperature regime and active layer depth on the Yamal Peninsula Engineering and Engineering Construction Studies in the Yakutia Republic (Yakutsk: NTO) Bhatt U S, Walker D A, Raynolds M and Comiso J 2007 The relationship between sea ice variability and Arctic tundra on the pan-Arctic, regional, and site scales Eos Trans. AGU 88 (52) Fall Meet. Suppl. Abstract U41C-0612 Bhatt U S, Walker D A, Raynolds M K and Comiso J 2008 Circumpolar and regional analysis of the relationship between sea-ice variability, summer land-surface temperatures, Arctic tundra greenness and large-scale climate drivers