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

Arctic shrub expansion is occurring across large parts of the tundra biome and its potential ecological repercussions have been widely discussed. But while the term “shrub expansion” often implicitly refers to an increase in tall, deciduous species such as birch and willow, several studies have also found a strong increase in evergreen dwarf shrubs in response to warming, a fact which has received far less attention. The effects of an evergreen dwarf shrub expansion are markedly different from the effects of an increase in taller, deciduous species. While deciduous shrubs may increase carbon (C) cycling through changes in albedo, litter input, and snow depth, the low stature of evergreen dwarf shrubs means that they are unlikely to influence snow cover. They also produce more recalcitrant litter, which reduces microbial activity. Furthermore, recent research suggests that ericoid mycorrhiza associated with evergreen shrubs may help to decelerate litter and soil organic matter turnover rates through the production of melanized hyphae that resist decomposition. Through selective browsing, herbivores may promote evergreen shrubs and facilitate C storage. Synthesis. In this mini review, we argue that basing predictions of how shrub expansion will affect tundra ecosystems on characteristics only applicable to tall deciduous shrubs hampers our understanding of the complex feedbacks related to Arctic vegetation shifts.

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.

Summary One of the most palpable effects of warming in Arctic ecosystems is shrub expansion above the tree line. However, previous studies have found that reindeer can influence plant community responses to warming and inhibit shrubification of the tundra.We revisited grazed (ambient) and ungrazed study plots (exclosures), at the southern as well as the northern limits of the Swedish alpine region, to study long‐term grazing effects and vegetation changes in response to increasing temperatures between 1995 and 2011, in two vegetation types (shrub heath and mountain birch forest).In the field layer at the shrub heath sites, evergreen dwarf shrubs had increased in cover from 26% to 49% but were unaffected by grazing. Deciduous dwarf and tall shrubs also showed significant, though smaller, increases over time. At the birch forest sites, the increase was similar for evergreen dwarf shrubs (20–48%) but deciduous tall shrubs did not show the same consistent increase over time as in the shrub heath.The cover and height of the shrub layer were significantly greater in exclosures at the shrub heath sites, but no significant treatment effects were found on species richness or diversity.July soil temperatures and growing season thawing degree days (TDD) were higher in exclosures at all but one site, and there was a significant negative correlation between mean shrub layer height and soil TDD at the shrub heath sites. Synthesis. This study shows that shrub expansion is occurring rapidly in the Scandes mountain range, both above and below the tree line. Tall, deciduous shrubs had benefitted significantly from grazing exclosure, both in terms of cover and height, which in turn lowered summer soil temperatures. However, the overriding vegetation shift across our sites was the striking increase in evergreen dwarf shrubs, which were not influenced by grazing. As the effects of an increase in evergreen dwarf shrubs and more recalcitrant plant litter may to some degree counteract some of the effects of an increase in deciduous tall shrubs, herbivore influence on shrub interactions is potentially of great importance for shaping arctic shrub expansion and its associated ecosystem effects.

The majority of variation in six traits critical to the growth, survival and reproduction of plant species is thought to be organised along just two dimensions, corresponding to strategies of plant size and resource acquisition. However, it is unknown whether global plant trait relationships extend to climatic extremes, and if these interspecific relationships are confounded by trait variation within species. We test whether trait relationships extend to the cold extremes of life on Earth using the largest database of tundra plant traits yet compiled. We show that tundra plants demonstrate remarkably similar resource economic traits, but not size traits, compared to global distributions, and exhibit the same two dimensions of trait variation. Three quarters of trait variation occurs among species, mirroring global estimates of interspecific trait variation. Plant trait relationships are thus generalizable to the edge of global trait-space, informing prediction of plant community change in a warming world.

1. Large herbivores influence plant community structure and ecosystem processes in many ecosystems. In large parts of the Arctic, reindeer (or caribou) are the only large herbivores present. Recent studies show that reindeer have the potential to mitigate recent warming-induced shrub encroachment in the Arctic and the associated greening of high-latitude ecosystems. This will potentially have large scale consequences for ecosystem productivity and carbon cycling.2. To date, information on variation in the interactions between reindeer and plants across Arctic landscapes has been scarce. We utilized a network of experimental sites across a latitudinal gradient in the Scandinavian mountains where reindeer have been excluded from 59 study plots for at least 15 years. We used this study system to test the effect of long-term exclusion of reindeer on the abundance of major plant functional groups, the greenness indexes Leaf Area Index (LAI) andNormalized Difference Vegetation Index (NDVI), soil mineral nitrogen (N) and phosphorous (P), and species richness, and to determine whether the effect of reindeer exclusion is dependent on reindeer density, productivity, soil fertility or climate.3. We found that NDVI and LAI, lichen and deciduous shrub abundances were largely reduced while soil mineral N was enhanced by reindeer. The direction and amplitude of other plant functional group responses to reindeer exclusion differed between forest and tundra as well as shrub-and herbaceous-dominated vegetation. Higher reindeer densities were related to decreased plant species richness in low-productive sites and to increased species richness in productive sites. The relative reduction in LAI and associated absolute reductions of deciduousshrubs in response to reindeer were positively related to reindeer density, while the relative reduction in NDVI was not. Further, relative reductions in LAI and NDVI in response to reindeer were unrelated to climate and soil fertility. Synthesis.Our results provide long-term experimental evidence highlighting the role of reindeer density in regulating plant species richness, global climate change induced greenness patterns and shrub encroachment at regional scales in the Arctic. These findings emphasize the need to consider reindeer in models predicting vegetation patterns and changes in high-latitude ecosystems. | 2725Journal of Ecology SUNDQVIST eT al.

Aim Plant functional groups are widely used in community ecology and earth system modelling to describe trait variation within and across plant communities. However, this approach rests on the assumption that functional groups explain a large proportion of trait variation among species. We test whether four commonly used plant functional groups represent variation in six ecologically important plant traits. Location Tundra biome. Time period Data collected between 1964 and 2016. Major taxa studied 295 tundra vascular plant species. Methods We compiled a database of six plant traits (plant height, leaf area, specific leaf area, leaf dry matter content, leaf nitrogen, seed mass) for tundra species. We examined the variation in species‐level trait expression explained by four traditional functional groups (evergreen shrubs, deciduous shrubs, graminoids, forbs), and whether variation explained was dependent upon the traits included in analysis. We further compared the explanatory power and species composition of functional groups to alternative classifications generated using post hoc clustering of species‐level traits. Results Traditional functional groups explained significant differences in trait expression, particularly amongst traits associated with resource economics, which were consistent across sites and at the biome scale. However, functional groups explained 19% of overall trait variation and poorly represented differences in traits associated with plant size. Post hoc classification of species did not correspond well with traditional functional groups, and explained twice as much variation in species‐level trait expression. Main conclusions Traditional functional groups only coarsely represent variation in well‐measured traits within tundra plant communities, and better explain resource economic traits than size‐related traits. We recommend caution when using functional group approaches to predict tundra vegetation change, or ecosystem functions relating to plant size, such as albedo or carbon storage. We argue that alternative classifications or direct use of specific plant traits could provide new insights for ecological prediction and modelling.

Plant communities in Arctic and alpine areas are changing due to higher temperatures and longer vegetation periods and it is uncertain how this will affect plant-herbivore dynamics. For instance, relatively fast-growing, deciduous shrub species that are the most responsive to warming may also be the most targeted by herbivores such as reindeer, giving less palatable evergreen shrubs the chance to expand. Using herbivore exclosures, we have studied how two grasslands with contrasting nutrient and moisture regimes, a dry, nutrient-poor alpine grass heath and a wet, productive low herb meadow, changed between 1995 and 2012, in grazed and ungrazed conditions. At the grass heath, evergreen low shrub abundance had more than doubled, regardless of grazer treatment, whereas at the low herb meadow, evergreen shrubs had increased only outside exclosures while deciduous tall shrubs and forbs were significantly more abundant inside exclosures. Deciduous tall shrubs were also significantly taller in exclosures. These contrasting findings suggest that the impact of herbivores is to a great deal determined by their influence on competitive interactions between plant species, and therefore depends on the underlying composition of the plant community. Consequently, as the balance in these competitive interactions is shifting due to climate warming, we conclude that the potential of herbivory to influence this balance is considerable yet highly site dependent.