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Observations of changes in phenology have provided some of the strongest signals of the effects of climate change on terrestrial ecosystems. The International Tundra Experiment (ITEX), initiated in the early 1990s, established a common protocol to measure plant phenology in tundra study areas across the globe. Today, this valuable collection of phenology measurements depicts the responses of plants at the colder extremes of our planet to experimental and ambient changes in temperature over the past decades. The database contains 150,434 phenology observations of 278 plant species taken at 28 study areas for periods of 1 to 26 years. Here we describe the full dataset to increase the visibility and use of these data in global analyses, and to invite phenology data contributions from underrepresented tundra locations. Portions of this tundra phenology database have been used in three recent syntheses, some datasets are expanded, others are from entirely new study areas, and the entirety of these data are now available at the Polar Data Catalogue., Arctic Science, 8 (3), ISSN:2368-7460

Open top chambers (OTCs) were adopted as the recommended warming mechanism by the International Tundra Experiment network in the early 1990s. Since then, OTCs have been deployed across the globe. Hundreds of papers have reported the impacts of OTCs on the abiotic environment and the biota. Here, we review the impacts of the OTC on the physical environment, with comments on the appropriateness of using OTCs to characterize the response of biota to warming. The purpose of this review is to guide readers to previously published work and to provide recommendations for continued use of OTCs to understand the implications of warming on low stature ecosystems. In short, the OTC is a useful tool to experimentally manipulate temperature; however, the characteristics and magnitude of warming varies greatly in different environments; therefore, it is important to document chamber performance to maximize the interpretation of biotic response. When coupled with long-term monitoring, warming experiments are a valuable means to understand the impacts of climate change on natural ecosystems. Open top chambers (OTCs) were adopted as the recommended warming mechanism by the International Tundra Experiment network in the early 1990s. Since then, OTCs have been deployed across the globe. Hundreds of papers have reported the im-pacts of OTCs on the abiotic environment and the biota. Here, we review the impacts of the OTC on the physical environment, with comments on the appropriateness of using OTCs to characterize the response of biota to warming. The purpose of this review is to guide readers to previously published work and to provide recommendations for continued use of OTCs to under -stand the implications of warming on low stature ecosystems. In short, the OTC is a useful tool to experimentally manipulate temperature; however, the characteristics and magnitude of warming varies greatly in different environments; therefore, it is important to document chamber performance to maximize the interpretation of biotic response. When coupled with long-term monitoring, warming experiments are a valuable means to understand the impacts of climate change on natural ecosystems.

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.

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., Nature Communications, 12 (1), ISSN:2041-1723

The research data repository of the Environmental Data Initiative (EDI) is a signatory of the FAIR Data Principles. Building on over 30 years of data curation research and experience in the NSF-fun...

In their recent paper, Huang et al. (1) have amassed a fascinating time-series of xylem tissue formation across 826 individual trees, spanning 21 species and 79 Northern Hemisphere locations. Like many other aspects of plant growth, wood formation shows strong seasonality, but very little is known about the environmental triggers for seasonal growth resumption. With wood absorbing ∼15% of anthropogenic CO2 emissions annually, this extensive dataset provides an exciting opportunity to refine earth system models, but it does not provide compelling evidence that photoperiod is the dominant season cue. The crux of Huang et al.’s conclusions are derived from their observed positive relationship between the day of year of wood formation (DOYwf) and the photoperiod on the day of year of wood formation (Photoperiodwf), using an analysis that relies heavily on R 2 values (ref. 1, figures 3, S5, and S7). The authors used the response variable (DOYwf) to calculate several of their predictor variables: Photoperiodwf, cumulative forcing (FUwf), and chilling (CUwf) at DOYwf. There are two major problems with this approach. First, it can lead to regression models that appear to fit exceptionally well for the wrong reasons. Where photoperiod covaries positively with day of … [↵][1]1To whom correspondence may be addressed. Email: sarah.elmendorf{at}colorado.edu. [1]: #xref-corresp-1-1

The Niwot Ridge and Green Lakes Valley (NWT) long-term ecological research (LTER) site collects environmental observations spanning both alpine and subalpine regimes. The first observations began in 1952 and have since expanded to nearly 300 available datasets over an area of 99 km2 within the north-central Colorado Rocky Mountains that include hydrological (n = 101), biological (n = 79), biogeochemical (n = 62), and geographical (n = 56) observations. The NWT LTER database is well suited to support hydrologic investigations that require long-term and interdisciplinary data sets. Experimentation and data collection at the NWT LTER are designed to characterize ecological responses of high-mountain environments to changes in climate, nutrients, and water availability. In addition to the continuation of the many legacy NWT datasets, expansion of the breadth and utility of the NWT LTER database is driven by new initiatives including (a) a catchment-scale sensor network of soil moisture, temperature, humidity, and snow-depth observations to understand hydrologic connectivity and (b) snow-albedo alteration experiments using black carbon to evaluate the effects of snow-disappearance on ecosystems. Together, these observational and experimental datasets provide a substantial foundation for hydrologic studies seeking to understand and predict changes to catchment and local-scale process interactions.

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., It is unclear whether plant trait relationships found at the global scale extend to climatic extremes. Here the authors analyse six major aboveground traits to show that known plant trait relationships extend to the tundra biomes and exhibit the same two dimensions of variation detected at the global scale.

The research data repository of the Environmental Data Initiative (EDI) is building on over 30 years of data curation research and experience in the National Science Foundation-funded US Long-Term Ecological Research (LTER) Network. It provides mature functionalities, well established workflows, and now publishes all ‘long-tail’ environmental data. High quality scientific metadata are enforced through automatic checks against community developed rules and the Ecological Metadata Language (EML) standard. Although the EDI repository is far along in making its data findable, accessible, interoperable, and reusable (FAIR), representatives from EDI and the LTER are developing best practices for the edge cases in environmental data publishing. One of these is the vast amount of imagery taken in the context of ecological research, ranging from wildlife camera traps to plankton imaging systems to aerial photography. Many images are used in biodiversity research for community analyses (e.g., individual counts, species cover, biovolume, productivity), while others are taken to study animal behavior and landscape-level change. Some examples from the LTER Network include: using photos of a heron colony to measure provisioning rates for chicks (Clarkson and Erwin 2018) or identifying changes in plant cover and functional type through time (Peters et al. 2020). Multi-spectral images are employed to identify prairie species. Underwater photo quads are used to monitor changes in benthic biodiversity (Edmunds 2015). Sosik et al. (2020) used a continuous Imaging FlowCytobot to identify and measure phyto- and microzooplankton. Cameras at McMurdo Dry Valleys assess snow and ice cover on Antarctic lakes allowing estimation of primary production (Myers 2019). It has been standard practice to publish numerical data extracted from images in EDI; however, the supporting imagery generally has not been made publicly available. Our goal in developing best practices for documenting and archiving these images is for them to be discovered and re-used. Our examples demonstrate several issues. The research questions, and hence, the image subjects are variable. Images frequently come in logical sets of time series. The size of such sets can be large and only some images may be contributed to a dedicated specialized repository. Finally, these images are taken in a larger monitoring context where many other environmental data are collected at the same time and location. Currently, a typical approach to publishing image data in EDI are packages containing compressed (ZIP or tar) files with the images, a directory manifest with additional image-specific metadata, and a package-level EML metadata file. Images in the compressed archive may be organized within directories with filenames corresponding to treatments, locations, time periods, individuals, or other grouping attributes. Additionally, the directory manifest table has columns for each attribute. Package-level metadata include standard coverage elements (e.g., date, time, location) and sampling methods. This approach of archiving logical ‘sets’ of images reduces the effort of providing metadata for each image when most information would be repeated, but at the expense of not making every image individually searchable. The latter may be overcome if the provided manifest contains standard metadata that would allow searching and automatic integration with other images.

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.

As the Arctic warms, vegetation is responding, and satellite measures indicate widespread greening at high latitudes. This ‘greening of the Arctic’ is among the world’s most important large-scale ecological responses to global climate change. However, a consensus is emerging that the underlying causes and future dynamics of so-called Arctic greening and browning trends are more complex, variable and inherently scale-dependent than previously thought. Here we summarize the complexities of observing and interpreting high-latitude greening to identify priorities for future research. Incorporating satellite and proximal remote sensing with in-situ data, while accounting for uncertainties and scale issues, will advance the study of past, present and future Arctic vegetation change. acceptedVersion

Essential Biodiversity Variables (EBVs) allow observation and reporting of global biodiversity change, but a detailed framework for the empirical derivation of specific EBVs has yet to be developed. Here, we re-examine and refine the previous candidate set of species traits EBVs and show how traits related to phenology, morphology, reproduction, physiology and movement can contribute to EBV operationalization. The selected EBVs express intra-specific trait variation and allow monitoring of how organisms respond to global change. We evaluate the societal relevance of species traits EBVs for policy targets and demonstrate how open, interoperable and machine-readable trait data enable the building of EBV data products. We outline collection methods, meta(data) standardization, reproducible workflows, semantic tools and licence requirements for producing species traits EBVs. An operationalization is critical for assessing progress towards biodiversity conservation and sustainable development goals and has wide implications for data-intensive science in ecology, biogeography, conservation and Earth observation., Sí

The sensitivity of phenology to environmental drivers can vary across geography and species. As such, models developed to predict phenology are typically site- or taxon-specific. Generation of site- and taxon-specific models is limited by the intensive in-situ phenological monitoring effort required to generate sufficient data to parameterize each model. Where in-situ phenological observations exist, the data are often subject to analytical issues due to the limited duration of any individual monitoring program, spotty site- and species- level coverage, lack of standardized methodology, and infrequent or variable census intervals. Together, these characteristics constrain our ability to make phenological inferences outside of select sites and taxa where long-duration, intensive monitoring has occurred. In this study, we leveraged two national, standardized phenology datasets to develop a multi-species and multi-site state-space survival model of the onset of deciduous tree and shrub spring (leaf out) and fall (leaf-color) events across temperate ecoregions of the United States. We used data from two national-scale phenological databases, a 9-year, broadly distributed dataset from the USA National Phenology Network and a 4-year dataset from the National Ecological Observatory Network, to quantify regional and interspecific variation in sensitivity to environmental drivers for both spring and fall leaf phenophases. Spring leaf out was generally promoted by longer days, spring growing degree day accumulation, overwinter chilling, and was suppressed by frost events, whereas fall leaf color was promoted by shorter days and cold accumulation. The sensitivity to most environmental drivers tended to be more variable among species than among the regions as defined here (EPA ecoregions of North America, excluding desert and tropical areas). The results of this study lay the groundwork for incorporating the growing collection of phenological observations into a generalized framework for predicting the transition states for any species, in any location.

Motivation The BioTIME database contains raw data on species identities and abundances in ecological assemblages through time. These data enable users to calculate temporal trends in biodiversity within and amongst assemblages using a broad range of metrics. BioTIME is being developed as a community-led open-source database of biodiversity time series. Our goal is to accelerate and facilitate quantitative analysis of temporal patterns of biodiversity in the Anthropocene. Main types of variables included The database contains 8,777,413 species abundance records, from assemblages consistently sampled for a minimum of 2 years, which need not necessarily be consecutive. In addition, the database contains metadata relating to sampling methodology and contextual information about each record. Spatial location and grain BioTIME is a global database of 547,161 unique sampling locations spanning the marine, freshwater and terrestrial realms. Grain size varies across datasets from 0.0000000158 km2 (158 cm2) to 100 km2 (1,000,000,000,000 cm2). Time period and grain BioTIME records span from 1874 to 2016. The minimal temporal grain across all datasets in BioTIME is a year. Major taxa and level of measurement BioTIME includes data from 44,440 species across the plant and animal kingdoms, ranging from plants, plankton and terrestrial invertebrates to small and large vertebrates. Software format .csv and .SQL., Global Ecology and Biogeography, 27 (7), ISSN:1466-822X, ISSN:1466-8238

Altres ajuts europeus: P.A.W. was additionally supported by the European Union Fourth Environment and Climate Framework Programme (Project Number ENV4-CT970586)P.A.W. was additionally supported by the European Union Fourth Environment and Climate Framework Programme (Project Number ENV4-CT970586). The 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.

The Convention on Biological Diversity's strategic plan lays out five goals: “(A) address the underlying causes of biodiversity loss by mainstreaming biodiversity across government and society; (B) reduce the direct pressures on biodiversity and promote sustainable use; (C) improve the status of biodiversity by safeguarding ecosystems, species and genetic diversity; (D) enhance the benefits to all from biodiversity and ecosystem services; (E) enhance implementation through participatory planning, knowledge management and capacity building.” To meet and inform on the progress towards these goals, a globally coordinated approach is needed for biodiversity monitoring that is linked to environmental data and covers all biogeographic regions. During a series of workshops and expert discussions, we identified nine requirements that we believe are necessary for developing and implementing such a global terrestrial species monitoring program. The program needs to design and implement an integrated information chain from monitoring to policy reporting, to create and implement minimal data standards and common monitoring protocols to be able to inform Essential Biodiversity Variables (EBVs), and to develop and optimize semantics and ontologies for data interoperability and modelling. In order to achieve this, the program needs to coordinate diverse but complementary local nodes and partnerships. In addition, capacities need to be built for technical tasks, and new monitoring technologies need to be integrated. Finally, a global monitoring program needs to facilitate and secure funding for the collection of long-term data and to detect and fill gaps in under-observed regions and taxa. The accomplishment of these nine requirements is essential in order to ensure data is comprehensive, to develop robust models, and to monitor biodiversity trends over large scales. A global terrestrial species monitoring program will enable researchers and policymakers to better understand the status and trends of biodiversity.

Consistent with a warming climate, birds are shifting the timing of their migrations, but it remains unclear to what extent these shifts have kept pace with the changing environment. Because bird migration is primarily cued by annually consistent physiological responses to photoperiod, but conditions at their breeding grounds depend on annually variable climate, bird arrival and climate-driven spring events would diverge. We combined satellite and citizen science data to estimate rates of change in phenological interval between spring green-up and migratory arrival for 48 breeding passerine species across North America. Both arrival and green-up changed over time, usually in the same direction (earlier or later). Although birds adjusted their arrival dates, 9 of 48 species did not keep pace with rapidly changing green-up and across all species the interval between arrival and green-up increased by over half a day per year. As green-up became earlier in the east, arrival of eastern breeding species increasingly lagged behind green-up, whereas in the west—where green-up typically became later—birds arrived increasingly earlier relative to green-up. Our results highlight that phenologies of species and trophic levels can shift at different rates, potentially leading to phenological mismatches with negative fitness consequences.

Global change drivers influence ecological processes at multiple scales and manifest across most of Earth as changes in biodiversity, biogeochemical cycles, infectious disease incidence, and ecohydrology. Small-scale investigations provide compelling evidence of specific effects of global change on local systems, but are of limited use in modeling complex ecological processes at continental-to-global scales. Long-term observations distributed across a diversity of habitat types are needed to improve the ability to forecast ecological change at large spatial and temporal scales. This special issue introduces the Terrestrial Observation System (TOS) of the National Ecological Observatory Network (NEON), a long-term, continental-scale ecological research platform designed to deliver these large-scale datasets. The TOS measures biodiversity of key biota (soil microbes, insects, plants, small mammals), ecosystem productivity and biogeochemistry, infectious disease dynamics, phenology, and population dynamics. The articles in this special issue describe the scientific rationale for the sampling designs of the TOS, including an overview of protocols, locations, and frequencies of measurements. The science designs are a culmination of design requirements scoped by NEON and the National Science Foundation, best available practices put forth by the scientific community, input from technical working groups, and consideration of logistical and financial constraints by NEON staff. Within each site, measurements have been collocated to the extent possible to optimize linkages among different sampling elements. Integrated analyses of terrestrial observations with sensor-based, imagery, and remote-sensing data collected by other NEON subsystems can facilitate scaling of measured parameters to larger spatial and temporal scales. NEON is designed to collect data for 30 years, and make these data freely available on a public data portal (data.neonscience.org). Samples and specimens will be archived and available to the scientific community upon request. The open access approach to the Observatory will provide users with the resources necessary to map, understand, and predict the effects of global change drivers on ecological processes at a continental scale.

Two recent meta-analyses of local-scale biodiversity change over time, by the authors of the present paper, have been subject to a harsh critique. Here we use new data and analyses to respond to the main points of this critique. First, a central argument of the critique was that short-term time series lead to biased estimates of long-term biodiversity change. However, we show here that this conclusion was based entirely on two fundamental mistakes in the simulations used to support it. Second, we show that the critic's conclusion that there are negative relationships between temporal biodiversity change and study duration is entirely dependent on: (i) the unrealistic assumption that biodiversity change can be positive when study duration = 0; (ii) the use of only a subset of the available data; (iii) inclusion of a single outlier data point in a single study (out of 100 in this case); and/or (iv) a choice to use log ratios rather than slopes (when available) as the metric of temporal biodiversity change. In short, the evidence does not support the conclusion that studies of longer duration tend to find local diversity decline. Finally, the critique highlighted the obviously true fact that studies in the ecological literature represent a geographically biased sample of locations on Earth; this issue was noted in both original papers, and is relevant for all ecological data syntheses. This fact was used by the critics to cast doubt on our conclusion that, outside of areas converted to croplands or asphalt, the distribution of temporal biodiversity trends is centered on zero. As a scientific rule, future studies based on more or different data may cause us to modify our conclusion, but at present, alternative conclusions based on the geographic-bias argument rely entirely on speculation. In sum, the critique raises points of uncertainty typical of all ecological studies, but it falls far short of providing an evidence-based alternative interpretation for our results, which are now supported by syntheses of hundreds of individual data sets of temporal biodiversity change.

© 2016 Elmendorf et al. Phenology is an integrative science that comprises the study of recurring biological activities or events. In an era of rapidly changing climate, the relationship between the timing of those events and environmental cues such as temperature, snowmelt, water availability, or day length are of particular interest. This article provides an overview of the observer-based plant phenology sampling conducted by the U.S. National Ecological Observatory Network (NEON), the resulting data, and the rationale behind the design. Trained technicians will conduct regular in situ observations of plant phenology at all terrestrial NEON sites for the 30-yr life of the observatory. Standardized and coordinated data across the network of sites can be used to quantify the direction and magnitude of the relationships between phenology and environmental forcings, as well as the degree to which these relationships vary among sites, among species, among phenophases, and through time. Vegetation at NEON sites will also be monitored with tower-based cameras, satellite remote sensing, and annual high-resolution airborne remote sensing. Ground-based measurements can be used to calibrate and improve satellite-derived phenometrics. NEON's phenology monitoring design is complementary to existing phenology research efforts and citizen science initiatives throughout the world and will produce interoperable data. By collocating plant phenology observations with a suite of additional meteorological, biophysical, and ecological measurements (e.g., climate, carbon flux, plant productivity, population dynamics of consumers) at 47 terrestrial sites, the NEON design will enable continental-scale inference about the status, trends, causes, and ecological consequences of phenological change.

Temperature is increasing at unprecedented rates across most of the tundra biome. Remote-sensing data indicate that contemporary climate warming has already resulted in increased productivity over much of the Arctic, but plot-based evidence for vegetation transformation is not widespread. We analysed change in tundra vegetation surveyed between 1980 and 2010 in 158 plant communities spread across 46 locations. We found biome-wide trends of increased height of the plant canopy and maximum observed plant height for most vascular growth forms; increased abundance of litter; increased abundance of evergreen, low-growing and tall shrubs; and decreased abundance of bare ground. Intersite comparisons indicated an association between the degree of summer warming and change in vascular plant abundance, with shrubs, forbs and rushes increasing with warming. However, the association was dependent on the climate zone, the moisture regime and the presence of permafrost. Our data provide plot-scale evidence linking changes in vascular plant abundance to local summer warming in widely dispersed tundra locations across the globe. © 2012 Macmillan Publishers Limited. All rights reserved.

35 Abstract Understanding the sensitivity of tundra vegetation to climate warming is critical to forecasting future biodiversity and vegetation feedbacks to climate. In situ warming experiments accelerate climate change on a small scale to forecast responses of local plant communities. Limitations of this approach include the apparent site-specificity of results and uncertainty about the power of short-term studies to anticipate longer term change. We address these issues with a synthesis of 61 experimental warming studies, of up to 20 years duration, in tundra sites worldwide. The response of plant groups to warming often differed with ambient summer temperature, soil moisture and experimental duration. Shrubs increased with warming only where ambient temperature was high, whereas graminoids increased primarily in the coldest study sites. Linear increases in effect size over time were frequently observed. There was little indication of saturating or accelerating effects, as would be predicted if negative or positive vegetation feedbacks were common. These results indicate that tundra vegetation exhibits strong regional variation in response to warming, and that in vulnerable regions, cumulative effects of long-term warming on tundra vegetation - and associated ecosystem consequences - have the potential to be much greater than we have observed to date.

There is considerable debate among ecologists as to whether or not communities are saturated. In saturated communities, species richness should remain relatively constant over time, despite compositional turnover, because richness is negatively correlated with colonization and positively correlated with local extinction. Few studies have tested for saturation using temporal observational data as well as diversity-perturbation experiments. We analyzed 10 years of data for plant species richness at 71 sites on contrasting serpentine and non-serpentine soils within Californian (USA) grasslands. We also manipulated local richness and measured its effects on immigration and extinction. Consistent with saturation, we observed that richness was positively correlated with extinction rates and negatively correlated with colonization rates, and randomization tests confirmed that diversity fluctuated less than expected by chance. However, experimental species additions and removals did not affect extinction or colonization, suggesting that richness is not regulated by local species interactions. Instead, we propose three reasons why richness may fluctuate within narrow limits causing the appearance of saturation in temporal observational data sets: negatively autocorrelated patterns of biotic response to yearly conditions, differential affinities of particular species for local conditions, or stochastic abundance-dependent colonization and extinction rates. We illustrate the latter using a metacommunity model.

Nestedness occurs when species-poor assemblages contain a subset of the species that occur in more species-rich communities and is a commonly observed pattern in spatial data sets. Examination of nested distribution patterns across time rather than space are rarely conducted, even though they may have important implications for species coexistence. Nested temporal assemblages can occur when most species respond similarly to interannual variation in conditions. In contrast, assemblages might be non-nested when different sets of species occur in different years, either because of different resource requirements or as a result of competitive exclusion. Using eight years of plant occurrence data at 71 sites in California grasslands, we found strong signals of temporal nestedness with most species favored by similar conditions. High-quality years enabled the expansion of both grasses and forbs into locales where they were not found during poor-quality years. Native annual forb, exotic annual forb, and exotic annual grass species richness were all greatest in cool, wet years following hot, dry years. Together, these analyses support the hypothesis that, in the absence of community members that specialize on poor-quality years, interannual environmental variation can cause communities to form nested subsets across time much as they do across space.

Rapid climate warming has been linked to increasing shrub dominance in the Arctic tundra. Research now shows that climate–shrub growth relationships vary spatially and according to site characteristics such as soil moisture and shrub height. Rapid climate warming in the tundra biome has been linked to increasing shrub dominance1,2,3,4. Shrub expansion can modify climate by altering surface albedo, energy and water balance, and permafrost2,5,6,7,8, yet the drivers of shrub growth remain poorly understood. Dendroecological data consisting of multi-decadal time series of annual shrub growth provide an underused resource to explore climate–growth relationships. Here, we analyse circumpolar data from 37 Arctic and alpine sites in 9 countries, including 25 species, and ∼42,000 annual growth records from 1,821 individuals. Our analyses demonstrate that the sensitivity of shrub growth to climate was: (1) heterogeneous, with European sites showing greater summer temperature sensitivity than North American sites, and (2) higher at sites with greater soil moisture and for taller shrubs (for example, alders and willows) growing at their northern or upper elevational range edges. Across latitude, climate sensitivity of growth was greatest at the boundary between the Low and High Arctic, where permafrost is thawing4 and most of the global permafrost soil carbon pool is stored9. The observed variation in climate–shrub growth relationships should be incorporated into Earth system models to improve future projections of climate change impacts across the tundra biome.

Recent changes in climate have led to significant shifts in phenology, with many studies demonstrating advanced phenology in response to warming temperatures. The rate of temperature change is especially high in the Arctic, but this is also where we have relatively little data on phenological changes and the processes driving these changes. In order to understand how Arctic plant species are likely to respond to future changes in climate, we monitored flowering phenology in response to both experimental and ambient warming for four widespread species in two habitat types over 21 years. We additionally used long-term environmental records to disentangle the effects of temperature increase and changes in snowmelt date on phenological patterns. While flowering occurred earlier in response to experimental warming, plants in unmanipulated plots showed no change or a delay in flowering over the 21-year period, despite more than 1 °C of ambient warming during that time. This counterintuitive result was likely due to significantly delayed snowmelt over the study period (0.05-0.2 days/yr) due to increased winter snowfall. The timing of snowmelt was a strong driver of flowering phenology for all species - especially for early-flowering species - while spring temperature was significantly related to flowering time only for later-flowering species. Despite significantly delayed flowering phenology, the timing of seed maturation showed no significant change over time, suggesting that warmer temperatures may promote more rapid seed development. The results of this study highlight the importance of understanding the specific environmental cues that drive species' phenological responses as well as the complex interactions between temperature and precipitation when forecasting phenology over the coming decades. As demonstrated here, the effects of altered snowmelt patterns can counter the effects of warmer temperatures, even to the point of generating phenological responses opposite to those predicted by warming alone.

We review and synthesize recent developments in the study of the invasion of communities in heterogeneous environments, considering both the invasibility of the community and impacts to the community. We consider both empirical and theoretical studies. For each of three major kinds of environmental heterogeneity (temporal, spatial and invader-driven), we find evidence that heterogeneity is critical to the invasibility of the community, the rate of spread, and the impacts on the community following invasion. We propose an environmental heterogeneity hypothesis of invasions, whereby heterogeneity both increases invasion success and reduces the impact to native species in the community, because it promotes invasion and coexistence mechanisms that are not possible in homogeneous environments. This hypothesis could help to explain recent findings that diversity is often increased as a result of biological invasions. It could also explain the scale dependence of the diversity-invasibility relationship. Despite the undoubted importance of heterogeneity to the invasion of communities, it has been studied remarkably little and new research is needed that simultaneously considers invasion, environmental heterogeneity and community characteristics. As a young field, there is an unrivalled opportunity for theoreticians and experimenters to work together to build a tractable theory informed by data.

Inference about future climate change impacts typically relies on one of three approaches: manipulative experiments, historical comparisons (broadly defined to include monitoring the response to ambient climate fluctuations using repeat sampling of plots, dendroecology, and paleoecology techniques), and space-for-time substitutions derived from sampling along environmental gradients. Potential limitations of all three approaches are recognized. Here we address the congruence among these three main approaches by comparing the degree to which tundra plant community composition changes (i) in response to in situ experimental warming, (ii) with interannual variability in summer temperature within sites, and (iii) over spatial gradients in summer temperature. We analyzed changes in plant community composition from repeat sampling (85 plant communities in 28 regions) and experimental warming studies (28 experiments in 14 regions) throughout arctic and alpine North America and Europe. Increases in the relative abundance of species with a warmer thermal niche were observed in response to warmer summer temperatures using all three methods; however, effect sizes were greater over broad-scale spatial gradients relative to either temporal variability in summer temperature within a site or summer temperature increases induced by experimental warming. The effect sizes for change over time within a site and with experimental warming were nearly identical. These results support the view that inferences based on space-for-time substitution overestimate the magnitude of responses to contemporary climate warming, because spatial gradients reflect long-term processes. In contrast, in situ experimental warming and monitoring approaches yield consistent estimates of the magnitude of response of plant communities to climate warming.

We review and synthesize recent developments in the study of the spread of invasive species, emphasizing both empirical and theoretical approaches. Recent theoretical work has shown that invasive species spread is a much more complex process than the classical models suggested, as long range dispersal events can have a large influence on the rate of range expansion through time. Empirical work goes even further, emphasizing the role of spatial heterogeneity, temporal variability, other species, and evolution. As in some of the classic work on spread, the study of range expansion of invasive species provides unique opportunities to use differences between theory and data to determine the important underlying processes that control spread rates.

Global biodiversity is in decline. This is of concern for aesthetic and ethical reasons, but possibly also for practical reasons, as suggested by experimental studies, mostly with plants, showing that biodiversity reductions in small study plots can lead to compromised ecosystem function. However, inferring that ecosystem functions will decline due to biodiversity loss in the real world rests on the untested assumption that such loss is actually occurring at these small scales in nature. Using a global database of 168 published studies and >16,000 nonexperimental, local-scale vegetation plots, we show that mean temporal change in species diversity over periods of 5–261 y is not different from zero, with increases at least as likely as declines over time. Sites influenced primarily by plant species’ invasions showed a tendency for declines in species richness, whereas sites undergoing postdisturbance succession showed increases in richness over time. Other distinctions among studies had little influence on temporal richness trends. Although maximizing diversity is likely important for maintaining ecosystem function in intensely managed systems such as restored grasslands or tree plantations, the clear lack of any general tendency for plant biodiversity to decline at small scales in nature directly contradicts the key assumption linking experimental results to ecosystem function as a motivation for biodiversity conservation in nature. How often real world changes in the diversity and composition of plant communities at the local scale cause ecosystem function to deteriorate, or actually to improve, remains unknown and is in critical need of further study.

The rapidly warming temperatures in high-latitude and alpine regions have the potential to alter the phenology of Arctic and alpine plants, affecting processes ranging from food webs to ecosystem trace gas fluxes. The International Tundra Experiment (ITEX) was initiated in 1990 to evaluate the effects of expected rapid changes in temperature on tundra plant phenology, growth and community changes using experimental warming. Here, we used the ITEX control data to test the phenological responses to background temperature variation across sites spanning latitudinal and moisture gradients. The dataset overall did not show an advance in phenology; instead, temperature variability during the years sampled and an absence of warming at some sites resulted in mixed responses. Phenological transitions of high Arctic plants clearly occurred at lower heat sum thresholds than those of low Arctic and alpine plants. However, sensitivity to temperature change was similar among plants from the different climate zones. Plants of different communities and growth forms differed for some phenological responses. Heat sums associated with flowering and greening appear to have increased over time. These results point to a complex suite of changes in plant communities and ecosystem function in high latitudes and elevations as the climate warms.

When the distribution of species is limited by propagule supply, new populations may be initiated by seed addition, but identifying suitable sites for efficiently targeted seed addition remains a major challenge for restoration. In addition to the biotic or abiotic variables typically used in species distribution models, spatial isolation from conspecifics could help predict the suitability of unoccupied sites. Site suitability might be expected to increase with spatial isolation after other factors are accounted for, since isolation increases the chance that a site is unoccupied only because of propagule limitation. For two native annual forbs in Californian grasslands, we combined experimental seeding and niche modeling to ask whether suitability of unoccupied sites could be predicted by spatial variables (either distances from, or densities of, conspecific populations), either by themselves or in combination with niche models. We also asked whether experimental tests of these predictions held up not only in the short term (one year), but also in the longer term (three years). For Lasthenia californica, seed additions were only successful relatively near existing populations. For Lupinus nanus, seeding success was low and was positively related to the number of conspecifics within 1 km. For both species, a few previously unoccupied sites remained occupied three years after seeding, but this subset was not predictable based on either spatial or niche variables. Seed addition alone may be a limited means of native forb restoration if suitable unoccupied sites are either rare or unpredictable, or if they tend to be close to where the species already occurs.

Recent research using repeat photography, long-term ecological monitoring and dendrochronology has documented shrub expansion in arctic, high-latitude and alpine tundra ecosystems. Here, we (1) synthesize these findings, (2) present a conceptual framework that identifies mechanisms and constraints on shrub increase, (3) explore causes, feedbacks and implications of the increased shrub cover in tundra ecosystems, and (4) address potential lines of investigation for future research. Satellite observations from around the circumpolar Arctic, showing increased productivity, measured as changes in ‘greenness’, have coincided with a general rise in high-latitude air temperatures and have been partly attributed to increases in shrub cover. Studies indicate that warming temperatures, changes in snow cover, altered disturbance regimes as a result of permafrost thaw, tundra fires, and anthropogenic activities or changes in herbivory intensity are all contributing to observed changes in shrub abundance. A large-scale increase in shrub cover will change the structure of tundra ecosystems and alter energy fluxes, regional climate, soil–atmosphere exchange of water, carbon and nutrients, and ecological interactions between species. In order to project future rates of shrub expansion and understand the feedbacks to ecosystem and climate processes, future research should investigate the species or trait-specific responses of shrubs to climate change including: (1) the temperature sensitivity of shrub growth, (2) factors controlling the recruitment of new individuals, and (3) the relative influence of the positive and negative feedbacks involved in shrub expansion.

Species distribution models are critical tools for the prediction of invasive species spread and conservation of biodiversity. The majority of species distribution models have been built with environmental data. Community ecology theory suggests that species co-occurrence data could also be used to predict current and potential distributions of species. Species assemblages are the products of biotic and environmental constraints on the distribution of individual species and as a result may contain valuable information for niche modeling. We compared the predictive ability of distribution models of annual grassland plants derived from either environmental or community-composition data. Composition-based models were built with the presence or absence of species at a site as predictors of site quality, whereas environment-based models were built with soil chemistry, moisture content, above-ground biomass, and solar radiation as predictors. The reproductive output of experimentally seeded individuals of 4 species and the abundance of 100 species were used to evaluate the resulting models. Community-composition data were the best predictors of both the site-specific reproductive output of sown individuals and the site-specific abundance of existing populations. Successful community-based models were robust to omission of data on the occurrence of rare species, which suggests that even very basic survey data on the occurrence of common species may be adequate for generating such models. Our results highlight the need for increased public availability of ecological survey data to facilitate community-based modeling at scales relevant to conservation.

There is currently no consensus on how physical and biological factors affect competitive intensity. Tests of whether competitive intensity varies along axes of environmental change have commonly been conducted in systems with a single strong environmental gradient, such as productivity, a soil resource, or an environmental stress. Frequently, these same axes are associated with changes in species composition, yet few studies have asked whether shifts in the identity of competitors affect competitive intensity. We ask whether resources (nutrients, water), stressors (heavy metals, Ca:Mg ratio), productivity (aboveground biomass), or species identity (an ordination axis of plant community composition) were the best predictors of the intensity of competition in a heterogeneous grassland landscape that included multiple independent environmental gradients. The reproductive fitness of six annual plant species was measured in the presence and absence of competitors and used to calculate relative interaction intensity (RII). We found that RII was best predicted by community composition. Nutrient availability was also important, and a post hoc test showed that competitive intensity was best explained by the combined effects of community composition and nutrient availability. We argue that community composition may be the most effective metric for predicting competitive intensity in many ecosystems because it includes both the competitive effects of the local community and information about covarying environmental characteristics.

How introduced plants, which may be locally adapted to specific climatic conditions in their native range, cope with the new abiotic conditions that they encounter as exotics is not well understood. In particular, it is unclear what role plasticity versus adaptive evolution plays in enabling exotics to persist under new environmental circumstances in the introduced range. We determined the extent to which native and introduced populations of St. John's Wort (Hypericum perforatum) are genetically differentiated with respect to leaf-level morphological and physiological traits that allow plants to tolerate different climatic conditions. In common gardens in Washington and Spain, and in a greenhouse, we examined clinal variation in percent leaf nitrogen and carbon, leaf delta(13)C values (as an integrative measure of water use efficiency), specific leaf area (SLA), root and shoot biomass, root/shoot ratio, total leaf area, and leaf area ratio (LAR). As well, we determined whether native European H. perforatum experienced directional selection on leaf-level traits in the introduced range and we compared, across gardens, levels of plasticity in these traits. In field gardens in both Washington and Spain, native populations formed latitudinal clines in percent leaf N. In the greenhouse, native populations formed latitudinal clines in root and shoot biomass and total leaf area, and in the Washington garden only, native populations also exhibited latitudinal clines in percent leaf C and leaf delta(13)C. Traits that failed to show consistent latitudinal clines instead exhibited significant phenotypic plasticity. Introduced St. John's Wort populations also formed significant or marginally significant latitudinal clines in percent leaf N in Washington and Spain, percent leaf C in Washington, and in root biomass and total leaf area in the greenhouse. In the Washington common garden, there was strong directional selection among European populations for higher percent leaf N and leaf delta(13)C, but no selection on any other measured trait. The presence of convergent, genetically based latitudinal clines between native and introduced H. perforatum, together with previously published molecular data, suggest that native and exotic genotypes have independently adapted to a broad-scale variation in climate that varies with latitude.