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 plant functional groups mediating climate impacts on carbon and biodiversity of alpine grasslands.

Plant removal experiments allow assessment of the role of biotic interactions among species or functional groups in community assembly and ecosystem functioning. When replicated along climate gradients, they can assess changes in interactions among species or functional groups with climate. Across twelve sites in the Vestland Climate Grid (VCG) spanning 4 °C in growing season temperature and 2000 mm in mean annual precipitation across boreal and alpine regions of Western Norway, we conducted a fully factorial plant functional group removal experiment (graminoids, forbs, bryophytes). Over six years, we recorded biomass removed, soil microclimate, plant community composition and structure, seedling recruitment, ecosystem carbon fluxes, and reflectance in 384 experimental and control plots. The dataset consists of 5,412 biomass records, 360 species-level biomass records, 1,084,970 soil temperature records, 4,771 soil moisture records, 17,181 plant records covering 206 taxa, 16,656 seedling records, 3,696 ecosystem carbon flux measurements, and 1,244 reflectance measurements. The data can be combined with longer-term climate data and plant population, community, ecosystem, and functional trait data collected within the VCG.

Patterns of free amino acids in tundra soils reflect mycorrhizal type, shrubification, and warming.

The soil nitrogen (N) cycle in cold terrestrial ecosystems is slow and organically bound N is an important source of N for plants in these ecosystems. Many plant species can take up free amino acids from these infertile soils, either directly or indirectly via their mycorrhizal fungi. We hypothesized that plant community changes and local plant community differences will alter the soil free amino acid pool and composition; and that long-term warming could enhance this effect. To test this, we studied the composition of extractable free amino acids at five separate heath, meadow, and bog locations in subarctic and alpine Scandinavia, with long-term (13 to 24 years) warming manipulations. The plant communities all included a mixture of ecto-, ericoid-, and arbuscular mycorrhizal plant species. Vegetation dominated by grasses and forbs with arbuscular and non-mycorrhizal associations showed highest soil free amino acid content, distinguishing them from the sites dominated by shrubs with ecto- and ericoid-mycorrhizal associations. Warming increased shrub and decreased moss cover at two sites, and by using redundancy analysis, we found that altered soil free amino acid composition was related to this plant cover change. From this, we conclude that the mycorrhizal type is important in controlling soil N cycling and that expansion of shrubs with ectomycorrhiza (and to some extent ericoid mycorrhiza) can help retain N within the ecosystems by tightening the N cycle.

Legacy effects of herbivory on treeline dynamics along an elevational gradient.

Treelines are expected to expand into alpine ecosystems with global warming, but herbivory may delay this expansion. This study quantifies long-term effects of temporally varying sheep densities on birch recruitment and growth in the treeline ecotone. We examined treeline ecotone successional trajectories and legacy effects in a replicated experimental setup, where enclosures were present for 14 years with three different sheep densities (0, 25, 80 sheep km-2). Before and after the enclosures were present, the site had an ambient sheep density of 20-25 km-2. We sampled field data 4 years after enclosure removal and compared these to data sampled 8 and 9 years after enclosure erection. We sampled data on birch browsing pressure, birch distribution across life-stages (recruits, saplings, and mature trees), and birch annual radial growth. Fourteen years of increased or decreased sheep density had observable legacy effects depending on birch life-stage. Birch recruit prevalence decreased in areas, where sheep were reintroduced after being absent for 14 years. For the same areas, sapling and mature tree prevalence increased, indicating that these areas have entered alternative successional trajectories compared to areas, where sheep were present the whole time. Birch annual radial growth showed a lag effect of 2 years after enclosure removal, with growth decreasing in areas where sheep had been absent for 14 years and increasing where sheep densities were high. Thus, decadal-scale absences of herbivores can leave legacy effects due to increased numbers of trees that have high resistance to later-introduced herbivore browsing.

Do trade-offs govern plant species' responses to different global change treatments?

Plants are subject to trade-offs among growth strategies such that adaptations for optimal growth in one condition can preclude optimal growth in another. Thus, we predicted that a plant species that responds positively to one global change treatment would be less likely than average to respond positively to another treatment, particularly for pairs of treatments that favor distinct traits. We examined plant species' abundances in 39 global change experiments manipulating two or more of the following: CO2 , nitrogen, phosphorus, water, temperature, or disturbance. Overall, the directional response of a species to one treatment was 13% more likely than expected to oppose its response to a another single-factor treatment. This tendency was detectable across the global data set, but held little predictive power for individual treatment combinations or within individual experiments. Although trade-offs in the ability to respond to different global change treatments exert discernible global effects, other forces obscure their influence in local communities.

Ontogenetic niche shifts in a locally endangered tree species (Olea europaea subsp. cuspidata) in a disturbed forest in Northern Ethiopia: Implications for conservation.

Understanding the responses of different ontogenetic stages to environmental and human disturbance factors is essential for developing efficient conservation strategies for endangered plant species. We examined how three ontogenetic stages of a locally endangered tree species, Olea europaea subsp. cuspidata, responded to environmental factors and human disturbance in Hugumburda dry Afromontane forest in Ethiopia. We counted individual seedlings, saplings and adults of O. europaea in 70 20 × 20 m quadrats over ca. 2.8 ha, and measured biotic (woody species richness, canopy cover, aboveground tree biomass, herbaceous cover), abiotic (soil and topographic variables), and human disturbance factors (logging and tracks). To detect ontogenetic niche shifts, we compared observed vs. simulated locations of trees in the three life stages and how they related to the environmental and human disturbance factors. We found that the population structure of O. europaea showed generally low recruitment, with few seedlings per hectare compared with the abundance of saplings and adults. The probability of finding O. europaea individuals was influenced by biotic (woody species richness) and abiotic (soil depth, slope) environmental conditions and human disturbance (logging intensity), but the direction, strength and shape of the relationships differed between seedling, sapling and adult life stages, indicating ontogenetic niche shifts. All life stages showed a positive relationship with elevation. The observed environmental niches of the different lifestages of O. europaea, and their association with human disturbance levels, should be considered when conservation strategies are developed for this species. Human disturbance in terms of logging decreases the abundance of saplings, but may facilitate emerging seedlings through creation of gaps with improved light conditions. Recruitment is, however, very low in the study area, and seedlings should be protected from browsing to enhance survival. Woody species richness in general should be conserved to optimize conditions also for O. europaea saplings.

Experimental warming differentially affects vegetative and reproductive phenology of tundra plants.

Rapid climate warming is altering Arctic and alpine tundra ecosystem structure and function, including shifts in plant phenology. While the advancement of green up and flowering are well-documented, it remains unclear whether all phenophases, particularly those later in the season, will shift in unison or respond divergently to warming. Here, we present the largest synthesis to our knowledge of experimental warming effects on tundra plant phenology from the International Tundra Experiment. We examine the effect of warming on a suite of season-wide plant phenophases. Results challenge the expectation that all phenophases will advance in unison to warming. Instead, we find that experimental warming caused: (1) larger phenological shifts in reproductive versus vegetative phenophases and (2) advanced reproductive phenophases and green up but delayed leaf senescence which translated to a lengthening of the growing season by approximately 3%. Patterns were consistent across sites, plant species and over time. The advancement of reproductive seasons and lengthening of growing seasons may have significant consequences for trophic interactions and ecosystem function across the tundra.

Macroecological context predicts species' responses to climate warming.

Context-dependencies in species' responses to the same climate change frustrate attempts to generalize and make predictions based on experimental and observational approaches in biodiversity science. Here, we propose predictability may be enhanced by explicitly incorporating macroecological context into analyses of species' responses to climate manipulations. We combined vascular plant species' responses to an 8-year, 12-site turf transplant climate change experiment set in southwestern Norway with climate niche data from the observed 151 species. We used the difference between a species' mean climate across their range and climate conditions at the transplant site ("climate differences") to predict colonization probability, extinction probability, and change in abundance of a species at a site. In analyses across species that ignore species-specific patterns, colonization success increased as species' distribution optima were increasingly warmer than the experimental target site. Extinction probability increased as species' distribution optima were increasingly colder than the target site. These patterns were reflected in change in abundance analyses. We found weak responses to increased precipitation in these oceanic climates. Climate differences were better predictors of species' responses to climate manipulations than range size. Interestingly, similar patterns were found when analyses focused on variation in species-specific responses across sites. These results provide an experimental underpinning to observational studies that report thermophilization of communities and suggest that space-for-time substitutions may be valid for predicting species' responses to climate warming, given other conditions are accounted for (e.g., soil nutrients). Finally, we suggest that this method of putting climate change experiments into macroecological context has the potential to generalize and predict species' responses to climate manipulations globally.

Biotic rescaling reveals importance of species interactions for variation in biodiversity responses to climate change.

Generality in understanding biodiversity responses to climate change has been hampered by substantial variation in the rates and even directions of response to a given change in climate. We propose that such context dependencies can be clarified by rescaling climate gradients in terms of the underlying biological processes, with biotic interactions as a particularly important process. We tested this rescaling approach in a replicated field experiment where entire montane grassland communities were transplanted in the direction of expected temperature and/or precipitation change. In line with earlier work, we found considerable variation across sites in community dynamics in response to climate change. However, these complex context dependencies could be substantially reduced or eliminated by rescaling climate drivers in terms of proxies of plant-plant interactions. Specifically, bryophytes limited colonization by new species into local communities, whereas the cover of those colonists, along with bryophytes, were the primary drivers of local extinctions. These specific interactions are relatively understudied, suggesting important directions for future work in similar systems. More generally, the success of our approach in explaining and simplifying landscape-level variation in climate change responses suggests that developing and testing proxies for relevant underlying processes could be a fruitful direction for building more general models of biodiversity response to climate change.

Plant traits and vegetation data from climate warming experiments along an 1100 m elevation gradient in Gongga Mountains, China.

Functional trait data enhance climate change research by linking climate change, biodiversity response, and ecosystem functioning, and by enabling comparison between systems sharing few taxa. Across four sites along a 3000-4130 m a.s.l. gradient spanning 5.3 °C in growing season temperature in Mt. Gongga, Sichuan, China, we collected plant functional trait and vegetation data from control plots, open top chambers (OTCs), and reciprocally transplanted vegetation turfs. Over five years, we recorded vascular plant composition in 140 experimental treatment and control plots. We collected trait data associated with plant resource use, growth, and life history strategies (leaf area, leaf thickness, specific leaf area, leaf dry matter content, leaf C, N and P content and C and N isotopes) from local populations and from experimental treatments. The database consists of 6,671 plant records and 36,743 trait measurements (increasing the trait data coverage of the regional flora by 500%) covering 11 traits and 193 plant taxa (ca. 50% of which have no previous published trait data) across 37 families.

Quantifying the roles of seed dispersal, filtering, and climate on regional patterns of grassland biodiversity.

Seed dispersal and local filtering interactively govern community membership and scale up to shape regional vegetation patterns, but data revealing how and why particular species are excluded from specific communities in nature are scarce. This lack of data is a missing link between our theoretical understanding of how diversity patterns can form and how they actually form in nature, and it hampers our ability to predict community responses to climate change. Here, we compare seed, seedling, and adult plant communities at 12 grassland sites with different climates in southern Norway to examine how community membership is interactively shaped by seed dispersal and local filtering, and how this process varies with climate across sites. To do this, we divide species at each site into two groups: locally transient species, which occur as seeds but are rare or absent as adults (i.e., they arrive but are filtered out), and locally persistent species, which occur consistently as adults in annual vegetation surveys. We then ask how and why locally transient species are disfavored during community assembly. Our results led to four main conclusions: (1) the total numbers of seeds and species that arrived, but failed to establish locally persistent populations, rose with temperature, indicating an increase in the realized effects of local filtering on community assembly, as well as an increase in the number of species poised to rapidly colonize those warmer sites if local conditions change in their favor, (2) locally transient species were selectively filtered out during seedling emergence, but not during seedling establishment, (3) selective filtering was partly driven by species climate preferences, exemplified by the poor performance of seeds dispersing outside of their realized climate niches into colder and drier foreign climates, and (4) locally transient species had traits that likely made them better dispersers (i.e., smaller seeds) but poorer competitors for light (i.e., shorter statures and less persistent clonal connections) than locally persistent species, potentially explaining why these species arrived to new sites but did not establish locally persistent adult populations. Our study is the first to combine seed, seedling, and adult survey data across sites to rigorously characterize how seed dispersal and local filtering govern community membership and shape climate-associated vegetation patterns.

Global change effects on plant communities are magnified by time and the number of global change factors imposed.

Global change drivers (GCDs) are expected to alter community structure and consequently, the services that ecosystems provide. Yet, few experimental investigations have examined effects of GCDs on plant community structure across multiple ecosystem types, and those that do exist present conflicting patterns. In an unprecedented global synthesis of over 100 experiments that manipulated factors linked to GCDs, we show that herbaceous plant community responses depend on experimental manipulation length and number of factors manipulated. We found that plant communities are fairly resistant to experimentally manipulated GCDs in the short term (<10 y). In contrast, long-term (≥10 y) experiments show increasing community divergence of treatments from control conditions. Surprisingly, these community responses occurred with similar frequency across the GCD types manipulated in our database. However, community responses were more common when 3 or more GCDs were simultaneously manipulated, suggesting the emergence of additive or synergistic effects of multiple drivers, particularly over long time periods. In half of the cases, GCD manipulations caused a difference in community composition without a corresponding species richness difference, indicating that species reordering or replacement is an important mechanism of community responses to GCDs and should be given greater consideration when examining consequences of GCDs for the biodiversity-ecosystem function relationship. Human activities are currently driving unparalleled global changes worldwide. Our analyses provide the most comprehensive evidence to date that these human activities may have widespread impacts on plant community composition globally, which will increase in frequency over time and be greater in areas where communities face multiple GCDs simultaneously.

Author Correction: Warming shortens flowering seasons of tundra plant communities.

In the version of this Article originally published, the following sentence was missing from the Acknowledgements: "This work was supported by the Norwegian Research Council SnoEco project, grant number 230970". This text has now been added.

Disturbance and the elevation ranges of woody plant species in the mountains of Costa Rica.

AIM: To understand how disturbance-here defined as a transient reduction in competition-can shape plant distributions along elevation gradients. Theory suggests that disturbance may increase elevation ranges, especially at the lower range limits, through reduced competitive exclusion. Nevertheless, to date this relationship remains unclear. LOCATION: Mountains of Costa Rica. METHODS: We compared the elevation range of woody stems over 10 cm dbh ("trees") observed in plots along two transects spanning a range of elevations in secondary (regrowth) and old-growth forest (409 and 249 species, respectively). We also estimated these elevation ranges using nationwide data. In addition, we examined the influence of stem size and plot scale basal area (as a measure of competition) on species elevation range limits in the two gradients. RESULTS: In general, tree species ranges increased with elevation. Species in the secondary forest had broader elevation ranges (100-318 m broader than species in the old-growth forest; Wilcoxon: p-value <.001). Also, in the secondary transect, individuals with greater diameters had broader elevation ranges than those observed as smaller trees (137 m broader; Kruskal-Wallis: p-value = .03). The lower range limit of species occurred more frequently in plots with lower (vs. higher) basal area than expected by chance in both forest types. We also observed higher elevation upper limits in old growth, but not in secondary forests, with lower (vs. higher) basal area. MAIN CONCLUSION: Disturbance relaxes the constraints imposed by competition and extends effective elevation ranges of species, particularly those in secondary forest, to warmer and cooler climates (minimum increase equivalent to about 0.6-1.4°C). Thus, suitable disturbance may assist species persistence under climate change. We believe this is the first study indicating a consistent relation between disturbance and woody plant species distributions along elevation gradients.

Warming shortens flowering seasons of tundra plant communities.

Advancing phenology is one of the most visible effects of climate change on plant communities, and has been especially pronounced in temperature-limited tundra ecosystems. However, phenological responses have been shown to differ greatly between species, with some species shifting phenology more than others. We analysed a database of 42,689 tundra plant phenological observations to show that warmer temperatures are leading to a contraction of community-level flowering seasons in tundra ecosystems due to a greater advancement in the flowering times of late-flowering species than early-flowering species. Shorter flowering seasons with a changing climate have the potential to alter trophic interactions in tundra ecosystems. Interestingly, these findings differ from those of warmer ecosystems, where early-flowering species have been found to be more sensitive to temperature change, suggesting that community-level phenological responses to warming can vary greatly between biomes.

Transplants, Open Top Chambers (OTCs) and Gradient Studies Ask Different Questions in Climate Change Effects Studies.

Long-term monitoring, space-for-time substitutions along gradients, and in situ temperature manipulations are common approaches to understand effects of climate change on alpine and arctic plant communities. Although general patterns emerge from studies using different approaches, there are also some inconsistencies. To provide better estimates of plant community responses to future warming across a range of environments, there have been repeated calls for integrating different approaches within single studies. Thus, to examine how different methods in climate change effect studies may ask different questions, we combined three climate warming approaches in a single study in the Hengduan Mountains of southwestern China. We monitored plant communities along an elevation gradient using the space-for-time approach, and conducted warming experiments using open top chambers (OTCs) and plant community transplantation toward warmer climates along the same gradient. Plant species richness and abundances were monitored over 5 years addressing two questions: (1) how do plant communities respond to the different climate warming approaches? (2) how can the combined approaches improve predictions of plant community responses to climate change? The general trend across all three approaches was decreased species richness with climate warming at low elevations. This suggests increased competition from immigrating lowland species, and/or from the species already growing inside the plots, as indicated by increased biomass, vegetation height or proportion of graminoids. At the coldest sites, species richness decreased in OTCs and along the gradient, but increased in the transplants, suggesting that plant communities in colder climates are more open to invasion from lowland species, with slow species loss. This was only detected in the transplants, showing that different approaches, may yield different results. Whereas OTCs may constrain immigration of new species, transplanted communities are rapidly exposed to new neighbors that can easily colonize the small plots. Thus, different approaches ask slightly different questions, in particular regarding indirect climate change effects, such as biotic interactions. To better understand both direct and indirect effects of climate change on plant communities, we need to combine approaches in future studies, and if novel interactions are of particular interest, transplants may be a better approach than OTCs.

Intraspecific Trait Variation and Phenotypic Plasticity Mediate Alpine Plant Species Response to Climate Change.

In a rapidly changing climate, alpine plants may persist by adapting to new conditions. However, the rate at which the climate is changing might exceed the rate of adaptation through evolutionary processes in long-lived plants. Persistence may depend on phenotypic plasticity in morphology and physiology. Here we investigated patterns of leaf trait variation including leaf area, leaf thickness, specific leaf area, leaf dry matter content, leaf nutrients (C, N, P) and isotopes (δ13C and δ15N) across an elevation gradient on Gongga Mountain, Sichuan Province, China. We quantified inter- and intra-specific trait variation and the plasticity in leaf traits of selected species to experimental warming and cooling by using a reciprocal transplantation approach. We found substantial phenotypic plasticity in most functional traits where δ15N, leaf area, and leaf P showed greatest plasticity. These traits did not correspond with traits with the largest amount of intraspecific variation. Plasticity in leaf functional traits tended to enable plant populations to shift their trait values toward the mean values of a transplanted plants' destination community, but only if that population started with very different trait values. These results suggest that leaf trait plasticity is an important mechanism for enabling plants to persist within communities and to better tolerate changing environmental conditions under climate change.

Accelerated increase in plant species richness on mountain summits is linked to warming.

Globally accelerating trends in societal development and human environmental impacts since the mid-twentieth century 1-7 are known as the Great Acceleration and have been discussed as a key indicator of the onset of the Anthropocene epoch 6 . While reports on ecological responses (for example, changes in species range or local extinctions) to the Great Acceleration are multiplying 8, 9 , it is unknown whether such biotic responses are undergoing a similar acceleration over time. This knowledge gap stems from the limited availability of time series data on biodiversity changes across large temporal and geographical extents. Here we use a dataset of repeated plant surveys from 302 mountain summits across Europe, spanning 145 years of observation, to assess the temporal trajectory of mountain biodiversity changes as a globally coherent imprint of the Anthropocene. We find a continent-wide acceleration in the rate of increase in plant species richness, with five times as much species enrichment between 2007 and 2016 as fifty years ago, between 1957 and 1966. This acceleration is strikingly synchronized with accelerated global warming and is not linked to alternative global change drivers. The accelerating increases in species richness on mountain summits across this broad spatial extent demonstrate that acceleration in climate-induced biotic change is occurring even in remote places on Earth, with potentially far-ranging consequences not only for biodiversity, but also for ecosystem functioning and services.

Asynchrony among local communities stabilises ecosystem function of metacommunities.

Temporal stability of ecosystem functioning increases the predictability and reliability of ecosystem services, and understanding the drivers of stability across spatial scales is important for land management and policy decisions. We used species-level abundance data from 62 plant communities across five continents to assess mechanisms of temporal stability across spatial scales. We assessed how asynchrony (i.e. different units responding dissimilarly through time) of species and local communities stabilised metacommunity ecosystem function. Asynchrony of species increased stability of local communities, and asynchrony among local communities enhanced metacommunity stability by a wide range of magnitudes (1-315%); this range was positively correlated with the size of the metacommunity. Additionally, asynchronous responses among local communities were linked with species' populations fluctuating asynchronously across space, perhaps stemming from physical and/or competitive differences among local communities. Accordingly, we suggest spatial heterogeneity should be a major focus for maintaining the stability of ecosystem services at larger spatial scales.

Greater temperature sensitivity of plant phenology at colder sites: implications for convergence across northern latitudes.

Warmer temperatures are accelerating the phenology of organisms around the world. Temperature sensitivity of phenology might be greater in colder, higher latitude sites than in warmer regions, in part because small changes in temperature constitute greater relative changes in thermal balance at colder sites. To test this hypothesis, we examined up to 20 years of phenology data for 47 tundra plant species at 18 high-latitude sites along a climatic gradient. Across all species, the timing of leaf emergence and flowering was more sensitive to a given increase in summer temperature at colder than warmer high-latitude locations. A similar pattern was seen over time for the flowering phenology of a widespread species, Cassiope tetragona. These are among the first results highlighting differential phenological responses of plants across a climatic gradient and suggest the possibility of convergence in flowering times and therefore an increase in gene flow across latitudes as the climate warms.