The Arctic-alpine biome is warming rapidly, resulting in a gradual replacement of low statured species by taller woody species in many tundra ecosystems. In northwest North America, the remotely sensed normalized difference vegetation index (NDVI), suggests an increase in productivity of the Arctic and alpine tundra and a decrease in productivity of boreal forests. However, the responses of contrasting shrub species growing at the same sites to climate drivers remain largely unexplored.

Here, we test growth, climate, and NDVI relationships of two contrasting species: the expanding tall deciduous shrub Salix pulchra and the circumarctic evergreen dwarf shrub Cassiope tetragona from an alpine tundra site in the Pika valley in the Kluane Region, southwest Yukon Territories, Canada.

We found that annual growth variability of both species at this site is strongly driven by early summer temperatures, despite their contrasting traits and habitats. Shrub growth chronologies for both species were correlated with the regional climate signal and showed spatial correspondence with interannual variation in NDVI in surrounding alpine and Arctic regions. Our results suggest that early summer warming represents a common driver of vegetation change for contrasting shrub species growing in different habitats in the same alpine environments.

The Arctic and alpine biome is rapidly warming, which might be causing an encroachment of relatively tall woody shrub vegetation into tundra ecosystems, which will probably result in an overall positive feedback to climate warming. This encroachment is, however, believed to remain limited to the relatively warm parts of the biome, where taller shrubs may displace shorter species. Still, climate sensitivity of shrub growth strongly differs between species and sites and High Arctic dwarf shrub species may respond rapidly to increasing temperatures in absence of taller species. In addition, it remains largely unknown whether shrubs from different functional groups from the same sites respond similarly to climate drivers. In the present study we examined the climate-growth relationships of six different site-species combinations: one evergreen and one deciduous shrub species at two alpine sites, and one evergreen dwarf shrub species at two High Arctic sites. We compared linear mixed models for each combination, explaining existing shrub growth data with site-speci c interpolated monthly and seasonal climate data from the gridded meteorological dataset CRU TS4.00. Shrub growth rates were found to be sensitive to summer climate for all species at all sites. Continued and projected warming is thus likely to stimulate a further encroachment of shrubs in these systems, at least through a densi cation of existing stands. Dwarf shrub growth strongly responded to the recent warming at both High Arctic sites, contrasting with previous work suggesting that shrub expansion might remain limited to warmer tundra regions. At the alpine sites, growth of evergreen shrubs was found to be more dependent on summer climate than growth of deciduous shrubs, perhaps because these evergreen species are less prone to herbivory. However, biome-wide generalizations at the func- tional group level may be dif cult to interpolate to the species level. Micro-site conditions, such as the determination of growing season length and winter soil temperatures, and in uence on growing season soil moisture by snow depth, may determine the strength of the climate-growth relationships found.

Siepielski et al. (Reports, 3 March 2017, p. 959) claim that “precipitation drives global variation in natural selection.” This conclusion is based on a meta-analysis of the relationship between climate variables and natural selection measured in wild populations of invertebrates, plants, and vertebrates. Three aspects of this analysis cause concern: (i) lack of within-year climate variables, (ii) low and variable estimates of covariance relationships across taxa, and (iii) a lack of mechanistic explanations for the patterns observed; association is not causation.

1. Climate warming is predicted to alter ecological boundaries in high‐latitude ecosystems including the elevational or latitudinal extent of tall shrubs in Arctic and alpine tundra. Over 60 studies from 128 locations around the tundra biome have investigated shrub expansion in tundra ecosystems; however, only six studies test whether shrublines are actually advancing up hill‐slopes or northward into tundra where tall shrubs are currently absent.

2. We test the hypothesis that willow shrublines have expanded to higher elevations in relation to climate across a 50 × 50 km area in the Kluane Region of the southwest Yukon Territory, Canada by surveying of 379 shrubs at 14 sites and sampling of 297 of the surveyed shrubs at 10 sites. We compared growth and recruitment to climate variables to test the climate sensitivity of shrub increase using annual radial growth analysis, age distributions and repeat field surveys to estimate the current rate of shrubline advance.

3. We found consistent and increasing rates of recruitment of alpine willows, with estimates of faster advancing shrublines on shallower hill‐slopes. Mortality was extremely low across the elevation gradient. Aspect, elevation and species identity did not explain variation in recruitment patterns, suggesting a regional factor, such as climate, as the driver of the observed shrubline advance.

4. Annual radial growth of willows was best explained by variation in summer temperatures, and recruitment pulses by winter temperatures. Measured recruitment rates are ~20 ± 5 individuals per hectare per decade (M ± SE) and measured rates of increased shrub cover of ~5 ± 1% per decade (M ± SE) measured at the Pika Camp site between field surveys in 2009 and 2013. Our results suggest that shrubline will continue to advance over the next 50 years, if growing conditions remain suitable. However, if future conditions differ between summer and winter seasons, this could lead to contrasting trajectories for recruitment vs. growth, and influence the vegetation change observed on the landscape.

5. Synthesis. Our findings in the context of a review of the existing literature indicate that elevational and latitudinal shrublines, like treelines, are advancing in response to climate warming; however, the trajectories of change will depend on the climate drivers controlling recruitment vs. growth.

Lightweight drones have emerged recently as a remote sensing survey tool of choice for ecologists, conservation practitioners and environmental scientists. In published work, there are plentiful details on the parameters and settings used for successful data capture, but in contrast there is a dearth of information describing the operational complexity of drone deployment. Information about the practices of flying in the field, whilst currently lacking, would be useful for others embarking on new drone‐based investigations. As a group of drone‐piloting scientists, we have operated lightweight drones for research in over 25 projects, in over 10 countries, and in polar, desert, coastal and tropical ecosystems, with many hundreds of hours of flying experience between us. The purpose of this paper was to document the lesser‐reported methodological pitfalls of drone deployments so that other scientists can understand the spectrum of considerations that need to be accounted for prior to, and during drone survey flights. Herein, we describe the most common challenges encountered, alongside mitigation and remediation actions that increase the chances of safe and successful data capture. Challenges are grouped into the following categories: (i) pre‐flight planning, (ii) flight operations, (iii) weather, (iv) redundancy, (v) data quality, (vi) batteries. We also discuss the importance of scientists undertaking ethical assessment of their drone practices, to identify and mitigate potential conflicts associated with drone use in particular areas. By sharing our experience, our intention is that the paper will assist those embarking on new drone deployments, increasing the efficacy of acquiring high‐quality data from this new proximal aerial viewpoint.

Woody shrubs have increased in biomass and expanded into new areas throughout the Pan-Arctic tundra biome in recent decades, which has been linked to a biome-wide observed increase in productivity. Experimental, observational, and socio-ecological research suggests that air temperature-and to a lesser degree precipitation-trends have been the predominant drivers of this change. However, a progressive decoupling of these drivers from Arctic vegetation productivity has been reported, and since 2010, vegetation productivity has also been declining. We created a protocol to (a) identify the suite of controls that may be operating on shrub growth and expansion, and (b) characterise the evidence base for controls on Arctic shrub growth and expansion. We found evidence for a suite of 23 proximal controls that operate directly on shrub growth and expansion; the evidence base focused predominantly on just four controls (air temperature, soil moisture, herbivory, and snow dynamics). 65% of evidence was generated in the warmest tundra climes, while 24% was from only one of 28 floristic sectors. Temporal limitations beyond 10 years existed for most controls, while the use of space-for-time approaches was high, with 14% of the evidence derived via experimental approaches. The findings suggest the current evidence base is not sufficiently robust or comprehensive at present to answer key questions of Pan-Arctic shrub change. We suggest future directions that could strengthen the evidence, and lead to an understanding of the key mechanisms driving changes in Arctic shrub environments.

Chronic, low intensity herbivory by invertebrates, termed background herbivory, has been understudied in tundra, yet its impacts are likely to increase in a warmer Arctic. The magnitude of these changes is however hard to predict as we know little about the drivers of current levels of invertebrate herbivory in tundra. We assessed the intensity of invertebrate herbivory on a common tundra plant, the dwarf birch (Betula glandulosa-nana complex), and investigated its relationship to latitude and climate across the tundra biome. Leaf damage by defoliating, mining and gall-forming invertebrates was measured in samples collected from 192 sites at 56 locations. Our results indicate that invertebrate herbivory is nearly ubiquitous across the tundra biome but occurs at low intensity. On average, invertebrates damaged 11.2% of the leaves and removed 1.4% of total leaf area. The damage was mainly caused by external leaf feeders, and most damaged leaves were only slightly affected (12% leaf area lost). Foliar damage was consistently positively correlated with mid-summer (July) temperature and, to a lesser extent, precipitation in the year of data collection, irrespective of latitude. Our models predict that, on average, foliar losses to invertebrates on dwarf birch are likely to increase by 6-7% over the current levels with a 1 °C increase in summer temperatures. Our results show that invertebrate herbivory on dwarf birch is small in magnitude but given its prevalence and dependence on climatic variables, background invertebrate herbivory should be included in predictions of climate change impacts on tundra ecosystems.

Shrub densification has been widely reported across the circumpolar arctic and subarctic biomes in recent years.Long-term analyses based on dendrochronological techniques applied to shrubs have linked this phenomenon to cli-mate change. However, the multi-stemmed structure of shrubs makes them difficult to sample and therefore leads tonon-uniform sampling protocols among shrub ecologists, who will favor either root collars or stems to conduct den-drochronological analyses. Through a comparative study of the use of root collars and stems of Betula glandulosa,acommon North American shrub species, we evaluated the relative sensitivity of each plant part to climate variablesand assessed whether this sensitivity is consistent across three different types of environments in northwesternQuebec, Canada (terrace, hilltop and snowbed). We found that root collars had greater sensitivity to climate thanstems and that these differences were maintained across the three types of environments. Growth at the root collarwas best explained by spring precipitation and summer temperature, whereas stem growth showed weak and incon-sistent responses to climate variables. Moreover, sensitivity to climate was not consistent among plant parts, as indi-viduals having climate-sensitive root collars did not tend to have climate-sensitive stems. These differences insensitivity of shrub parts to climate highlight the complexity of resource allocation in multi-stemmed plants. Whereasstem initiation and growth are driven by microenvironmental variables such as light availability and competition, rootcollars integrate the growth of all plant parts instead, rendering them less affected by mechanisms such as competitionand more responsive to signals of global change. Although further investigations are required to determine the degreeto which these findings are generalizable across the tundra biome, our results indicate that consistency and caution inthe choice of plant parts are a key consideration for the success of future dendroclimatological studies on shrubs.

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.

Plant communities have undergone dramatic changes in recent centuries, although not all such changes fit with the dominant biodiversity-crisis nar- rative used to describe them. At the global scale, future declines in plant species diversity are highly likely given habitat conversion in the tropics, although few extinctions have been documented for the Anthropocene to date (<0.1%). Nonnative species introductions have greatly increased plant species richness in many regions of the world at the same time that they have led to the creation of new hybrid polyploid species by bringing pre- viously isolated congeners into close contact. At the local scale, conversion of primary vegetation to agriculture has decreased plant diversity, whereas other drivers of change-e.g., climate warming, habitat fragmentation, and nitrogen deposition-have highly context-dependent effects, resulting in a distribution of temporal trends with a mean close to zero. These results prompt a reassessment of how conservation goals are defined and justified.

We present new data and analyses revealing fundamental flaws in a critique of two recent meta‐analyses of local‐scale temporal biodiversity change. First, the conclusion that short‐term time series lead to biased estimates of long‐term change was based on two errors in the simulations used to support it. Second, the conclusion of negative relationships between temporal biodiversity change and study duration was entirely dependent on unrealistic model assumptions, the use of a subset of data, and inclusion of one outlier data point in one study. Third, the finding of a decline in local biodiversity, after eliminating post‐disturbance studies, is not robust to alternative analyses on the original data set, and is absent in a larger, updated data set. Finally, the undebatable point, noted in both original papers, that studies in the ecological literature are geographically biased, was used to cast doubt on the conclusion that, outside of areas converted to croplands or asphalt, the distribution of biodiversity trends is centered approximately on zero. Future studies may modify conclusions, but at present, alternative conclusions based on the geographic‐bias argument rely on speculation. In sum, the critique raises points of uncertainty typical of all ecological studies, but does not provide an evidence‐based alternative interpretation.

Numerous international scientific assessments and related articles have, during the last decade, described the observed and potential impacts of climate change as well as other related environmental stressors on Arctic ecosystems. There is increasing recognition that observed and projected changes in freshwater sources, fluxes, and storage will have profound implications for the physical, biogeochemical, biological, and ecological processes and properties of Arctic terrestrial and freshwater ecosystems. However, a significant level of uncertainty remains in relation to forecasting the impacts of an intensified hydrological regime and related cryospheric change on ecosystem structure and function. As the terrestrial and freshwater ecology component of the Arctic Freshwater Synthesis, we review these uncertainties and recommend enhanced coordinated circumpolar research and monitoring efforts to improve quantification and prediction of how an altered hydrological regime influences local, regional, and circumpolar‐level responses in terrestrial and freshwater systems. Specifically, we evaluate (i) changes in ecosystem productivity; (ii) alterations in ecosystem‐level biogeochemical cycling and chemical transport; (iii) altered landscapes, successional trajectories, and creation of new habitats; (iv) altered seasonality and phenological mismatches; and (v) gains or losses of species and associated trophic interactions. We emphasize the need for developing a process‐based understanding of interecosystem interactions, along with improved predictive models. We recommend enhanced use of the catchment scale as an integrated unit of study, thereby more explicitly considering the physical, chemical, and ecological processes and fluxes across a full freshwater continuum in a geographic region and spatial range of hydroecological units (e.g., stream‐pond‐lake‐river‐near shore marine environments).

Changing environmental and geomorphological conditions are resulting in vegetation change in ice-wedge polygons in Arctic tundra. However, we do not yet know how microscale vegetation patterns relate to individual environmental and geomorphological parameters. This work aims at examining these relations in polygonal terrain. We analysed composition and cover of vascular plant taxa and surface height, active layer depth, soil temperature, carbon and nitrogen content, pH and electrical conductivity in four polygon mires located on the Yukon coast. We found that vascular plant species composition and cover correlates best with relative surface height. Ridges of low-centred polygons and raised centres of high-centred polygons support the growth of mesic and wetland species (e.g., Betula glandulosa, Salix pulchra, S. reticulata, Rubus chamaemorus, various ericaceous dwarf shrubs, Eriophorum vaginatum, Poa arctica). Wetland and aquatic plant species (e.g., E. angustifolium, Carex aquatilis, C. chordorrhiza, Pedicularis sudetica) grow in low-lying centres of polygons and in troughs between polygons. We also found a relationship between vascular plant species composition and substrate characteristics such as pH, electrical conductivity and total organic carbon, although the individual influence of these parameters could not be determined because of their correlation with relative surface height. Our findings stress the regulatory role of microtopography and substrate in vegetation dynamics of polygonal terrain. Ongoing warming in this region will lead to changes to polygonal terrain through permafrost degradation and subsequent conversion of low-centred into high-centred polygons. Our results indicate that shrubs, particularly Betula glandulosa and heath species, have the potential to expand most.

Permafrost landscapes experience different disturbances and store large amounts of organic matter, which may become a source of greenhouse gases upon permafrost degradation. We analysed the influence of terrain and geomorphic disturbances (e.g. soil creep, active‐layer detachment, gullying, thaw slumping, accumulation of fluvial deposits) on soil organic carbon (SOC) and total nitrogen (TN) storage using 11 permafrost cores from Herschel Island, western Canadian Arctic. Our results indicate a strong correlation between SOC storage and the topographic wetness index. Undisturbed sites stored the majority of SOC and TN in the upper 70 cm of soil. Sites characterised by mass wasting showed significant SOC depletion and soil compaction, whereas sites characterised by the accumulation of peat and fluvial deposits store SOC and TN along the whole core. We upscaled SOC and TN to estimate total stocks using the ecological units determined from vegetation composition, slope angle and the geomorphic disturbance regime. The ecological units were delineated with a supervised classification based on RapidEye multispectral satellite imagery and slope angle. Mean SOC and TN storage for the uppermost 1 m of soil on Herschel Island are 34.8 kg C m‐2 and 3.4 kg N m‐2, respectively.

Rapid climate warming in the tundra biome has been linked to increasing shrub dominance [1,2,3,4]. Shrub expansion can modify climate by altering surface albedo, energy and water balance, and permafrost [2,5,6,7,8], 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 stored [9]. 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.

1. The effects of climate change on Arctic ecosystems can range between various spatiotemporalscales and may include shifts in population distribution, community composition, plant phenology,primary productivity and species biodiversity. The growth rates and age structure of tundra vegeta-tion as well as its response to temperature variation, however, remain poorly understood becausehigh-resolution data are limited in space and time.

2. Anatomical and morphological stem characteristics were recorded to assess the growth behavi ourand age structure of 871 dwarf shrubs from 10 species at 30 sites in coastal East Greenland at ~70°N.Recruitment pulses were linked with changes in mean annual and summer temperature back to the 19thcentury, and a literature review was conducted to place our findings in a pan-Arctic context.

3. Low cambial activity translates into estimated average/maximum plant ages of 59/204 years, sug-gesting relatively small turnover rates and stable community composition. Decade-long changes inthe recruitment intensity were found to lag temperat ure variability by 2 and 6 years during warmerand colder periods, respectively (r = 0.851961-2000 and 1881-1920).

4. Synthesis. Our results reveal a strong temperature dependency of Arctic dwarf shrub reproduction,a high vulnerability of circumpolar tundra ecosystems to climatic changes, and the abil ity of evaluat-ing historical vegetation dynamics well beyond the northern treeline. The combined wood anatomi-cal and plant ecological approach, considering insights from micro-sections to communityassemblages, indicates that model predictions of rapid tundra expansion (i.e. shrub growth) follow-ing intense warming might underestimate plant longevity and persistence but overestimate the sensi-tivity and reaction time of Arctic vegetation.

Methodological constraints can limit our ability to quantify potential impacts of climate warming. We assessed the consistency of three approaches in estimating warming effects on plant community composition: manipulative warming experiments, repeat sampling under ambient temperature change (monitoring), and space-for-time substitution. The three approaches showed agreement in the direction of change (an increase in the relative abundance of species with a warmer thermal niche), but differed in the magnitude of change estimated. Experimental and monitoring approaches were similar in magnitude, whereas space-for-time comparisons indicated a much stronger response. These results suggest that all three approaches are valid, but experimental warming and long-term monitoring are best suited for forecasting impacts over the coming decades.

Shrubs have increased in abundance and dominance in arctic and alpine regions in recent decades. This often dramatic change, likely due to climate warming, has the potential to alter both the structure and function of tundra ecosystems. The analysis of shrub growth is improving our understanding of tundra vegetation dynamics and environmental changes. However, dendrochronological methods developed for trees, need to be adapted for the morphology and growth eccentricity of shrubs. Here, we review current and developing methods to measure radial and axial growth, estimate age, and assess growth dynamics in relation to environmental variables. Recent advances in sampling methods, analysis and applications have improved our ability to investigate growth and recruitment dynamics of shrubs. However, to extrapolate findings to the biome scale, future dendroecological work will require improved approaches that better address variation in growth within parts of the plant, among individuals within populations and between species.

A central current debate in community ecology concerns the relative importance of deterministic versus stochastic processes underlying community structure. However, the concept of stochasticity presents several profound philosophical, theoretical and empirical challenges, which we address here. The philosophical argument that nothing in nature is truly stochastic can be met with the following operational concept of neutral stochasticity in community ecology: change in the composition of a community (i.e. community dynamics) is neutrally stochastic to the degree that individual demographic events – birth, death, immigration, emigration – which cause such changes occur at random with respect to species identities. Empirical methods for identifying the stochastic component of community dynamics or structure include null models and multivariate statistics on observational species‐by‐site data (with or without environmental or trait data), and experimental manipulations of ‘stochastic’ species colonization order or relative densities and frequencies of competing species. We identify the fundamental limitations of each method with respect to its ability to allow inferences about stochastic community processes. Critical future needs include greater precision in articulating the link between results and ecological inferences, a comprehensive theoretical assessment of the interpretation of statistical analyses of observational data, and experiments focusing on community size and on natural variation in species colonization order.

Wetlands are the largest natural source of atmospheric methane. Here, we assess controls on methane flux using a database of approximately 19 000 instantaneous measurements from 71 wetland sites located across subtropical, temperate, and northern high latitude regions. Our analyses confirm general controls on wetland methane emissions from soil temperature, water table, and vegetation, but also show that these relationships are modified depending on wetland type (bog, fen, or swamp), region (subarctic to temperate), and disturbance. Fen methane flux was more sensitive to vegetation and less sensitive to temperature than bog or swamp fluxes. The optimal water table for methane flux was consistently below the peat surface in bogs, close to the peat surface in poor fens, and above the peat surface in rich fens. However, the largest flux in bogs occurred when dry 30‐day averaged antecedent conditions were followed by wet conditions, while in fens and swamps, the largest flux occurred when both 30‐day averaged antecedent and current conditions were wet. Drained wetlands exhibited distinct characteristics, e.g. the absence of large flux following wet and warm conditions, suggesting that the same functional relationships between methane flux and environmental conditions cannot be used across pristine and disturbed wetlands. Together, our results suggest that water table and temperature are dominant controls on methane flux in pristine bogs and swamps, while other processes, such as vascular transport in pristine fens, have the potential to partially override the effect of these controls in other wetland types. Because wetland types vary in methane emissions and have distinct controls, these ecosystems need to be considered separately to yield reliable estimates of global wetland methane release.

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.

Shrubs are the largest plant life form in tundra ecosystems; therefore, any changes in the abundance of shrubs will feedback to influence biodiversity, ecosystem function, and climate. The snow-shrub hypothesis asserts that shrub canopies trap snow and insulate soils in winter, increasing the rates of nutrient cycling to create a positive feedback to shrub expansion. However, previous work has not been able to separate the abiotic from the biotic influences of shrub canopies. We conducted a 3‐year factorial experiment to determine the influences of canopies on soil temperatures and nutrient cycling parameters by removing ~0.5 m high willow (Salix spp.) and birch (Betula glandulosa) shrubs, creating artificial shrub canopies and comparing these manipulations to nearby open tundra and shrub patches. Soil temperatures were 4-5°C warmer in January, and 2°C cooler in July under shrub cover. Natural shrub plots had 14-33 cm more snow in January than adjacent open tundra plots. Snow cover and soil temperatures were similar in the manipulated plots when compared with the respective unmanipulated treatments, indicating that shrub canopy cover was a dominant factor influencing the soil thermal regime. Conversely, we found no strong evidence of increased soil decomposition, CO2 fluxes, or nitrate or ammonia adsorbtion under artificial shrub canopy treatments when compared with unmanipulated open tundra. Our results suggest that the abiotic influences of shrub canopy cover alone on nutrient dynamics are weaker than previously asserted.

Predicting the future ecological impact of global change drivers requires understanding how these same drivers have acted in the past to produce the plant populations and communities we see today. Historical ecological data sources have made contributions of central importance to global change biology, but remain outside the toolkit of most ecologists. Here we review the strengths and weaknesses of four unconventional sources of historical ecological data: land survey records, “legacy” vegetation data, historical maps and photographs, and herbarium specimens. We discuss recent contributions made using these data sources to understanding the impacts of habitat disturbance and climate change on plant populations and communities, and the duration of extinction-colonization time lags in response to landscape change. Historical data frequently support inferences made using conventional ecological studies (e.g., increases in warm‐adapted species as temperature rises), but there are cases when the addition of different data sources leads to different conclusions (e.g., temporal vegetation change not as predicted by chronosequence studies). The explicit combination of historical and contemporary data sources is an especially powerful approach for unraveling long‐term consequences of multiple drivers of global change. Despite the limitations of historical data, which include spotty and potentially biased spatial and temporal coverage, they often represent the only means of characterizing ecological phenomena in the past and have proven indispensable for characterizing the nature, magnitude, and generality of global change impacts on plant populations and communities.

We present a focus issue of Environmental Research Letters on the ‘Recent dynamics of arctic and sub-arctic vegetation’. The focus issue includes three perspective articles (Verbyla 2011 Environ. Res. Lett. 6 041003, Williams et al 2011 Environ. Res. Lett. 6 041004, Loranty and Goetz 2012 Environ. Res. Lett. 7 011005) and 22 research articles. The focus issue arose as a result of heightened interest in the response of high-latitude vegetation to natural and anthropogenic changes in climate and disturbance regimes, and the consequences that these vegetation changes might have for northern ecosystems. A special session at the December 2010 American Geophysical Union Meeting on the ‘Greening of the Arctic’ spurred the call for papers. Many of the resulting articles stem from intensive research efforts stimulated by International Polar Year projects and the growing acknowledgment of ongoing climate change impacts in northern terrestrial ecosystems.

Collaboration and synthesis have become essential parts of research in the fields of ecology and evolution. Some of the most exciting and high‐impact research currently being published is coming from working groups, meta analyses and shared data synthesis activities (Carpenter et al. 2009, Hampton and Parker 2011, Cadotte et al. 2012). If you are an early‐career scientist, this type of collaboration will expand your research network, hopefully advance your career, and gives you the opportunity to do really “cool” science! But getting involved in successful synthesis projects takes some legwork.

Premise of the study : The development of biased sex ratios in dioecious plant species has been ascribed to either (1) factors influencing differential adult mortality of male and female plants or (2) factors acting at an early life stage that determine seed sex ratio or seedling survival.
Methods: To discriminate between these two competing hypotheses, we surveyed sex and age of 379 individuals from five species of the genus Salix across 1 1 alpine valleys in the southwest Yukon.
Key results : We observed uniformly female-biased sex ratios of approximately 2: 1 across all adult age cohorts and patch sizes of the five willow species. No spatial variation in sex ratio occurred that could be associated with site-specific characteristics such as elevation or aspect.
Conclusions: Our results indicate that the female-biased sex ratios in the alpine willow species investigated in this study are not a consequence of ecological processes acting on established adult plants. The sex ratio is instead determined at an early life stage by a mechanism that remains unknown.

First conceptualized in the 1970s, resilience has become a popular term in the ecological literature, used in the title, abstract, or keywords of approximately 1% of papers identified by ISI Web of Science in the field of environmental sciences and ecology in 2011. However, many papers make only passing reference to the term and do not explain what resilience means in the context of their study system, despite there being a number of possible definitions. In an attempt to determine how resilience is being used in ecological studies, we surveyed 234 papers published between 2004 and 2011 that were identified under the topic “resilience” by ISI Web of Science. Of these, 38% used the word resilience fewer than three times (often in the abstract or keyword list), 66% did not define the term, and 71% did not provide a citation to the resilience literature. Studies that defined resilience most often discussed it as pertaining to an entire ecosystem under continuous rather than discrete disturbance. Given the complex nature of this concept, we believe that care should be taken to properly describe what is meant by the term resilience in ecological studies.

Temperature is increasing at unprecedented rates across most of the tundra biome1. Remote-sensing data indicate that contemporary climate warming has already resulted in increased productivity over much of the Arctic2,3, 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.

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.

It is now well documented that Arctic climates and ecosystems are changing at some of the fastest rates on planet Earth. These changes are significant for all Arctic biodiversity, and they are a great challenge for cooperative management boards of Canada’s Arctic national parks, those legislated to maintain or improve the ecological integrity of all national parks. Owing to the inherent complexity of natural ecosystems, it is not at all clear how, nor how rapidly, these ongoing changes will affect park biodiversity and impact the traditional land-based lifestyles of Indigenous park cooperative management partners. In this context, this paper reviews and integrates recent research carried out in Canadian Arctic national parks: (1) geophysical – a reduction in glacial area and volume, active layer thickening, warming soil temperatures, and terrain instability; (2) vegetation – widespread but ecosystem-specific increases in NDVI ‘greenness’, plant biomass, shrub and herb coverage, and growing season lengths; and (3) wildlife – complex changes in small mammals and ungulate populations, very negative effects on some polar bear populations, and relatively stable mammalian predator and raptor populations at this time. This work provides a partial snapshot of ongoing and evolving ecological effects of climate change in Arctic national parks, and provides a strong foundation for prioritising future research and monitoring efforts. These evolving changes also undermine the historical paradigm of place-based conservation and necessitate a new approach for managing protected areas that involves acceptance of ongoing transformational change and adoption of a risk-based, forward looking paradigm in a changing world. It is proposed that Arctic national parks are ideal locations to focus Arctic science, especially as a component of a strategic, coordinated, and pan-Arctic approach to Arctic research that makes the most effective use of limited resources in the vast areas of Canada’s north.

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.

Canopy-forming shrubs are reported to be increasing at sites around the circumpolar Arctic. Our results indicate expansion in canopy cover and height of willows on Herschel Island located at 70° north on the western Arctic coast of the Yukon Territory. We examined historic photographs, repeated vegetation surveys, and conducted monitoring of long-term plots and found evidence of increases of each of the dominant canopy-forming willow species (Salix richardsonii, Salix glauca and Salix pulchra), during the twentieth century. A simple model of patch initiation indicates that the majority of willow patches for each of these species became established between 1910 and 1960, with stem ages and maximum growth rates indicating that some patches could have established as late as the 1980s. Collectively, these results suggest that willow species are increasing in canopy cover and height on Herschel Island. We did not find evidence that expansion of willow patches is currently limited by herbivory, disease, or growing conditions.

Understanding the responses of tundra systems to global change has global implications. Most tundra regions lack sustained environmental monitoring and one of the only ways to document multi-decadal change is to resample historic research sites. The International Polar Year (IPY) provided a unique opportunity for such research through the Back to the Future (BTF) project (IPY project #512). This article synthesizes the results from 13 papers within this Ambio Special Issue. Abiotic changes include glacial recession in the Altai Mountains, Russia; increased snow depth and hardness, permafrost warming, and increased growing season length in sub-arctic Sweden; drying of ponds in Greenland; increased nutrient availability in Alaskan tundra ponds, and warming at most locations studied. Biotic changes ranged from relatively minor plant community change at two sites in Greenland to moderate change in the Yukon, and to dramatic increases in shrub and tree density on Herschel Island, and in subarctic Sweden. The population of geese tripled at one site in northeast Greenland where biomass in non-grazed plots doubled. A model parameterized using results from a BTF study forecasts substantial declines in all snowbeds and increases in shrub tundra on Niwot Ridge, Colorado over the next century. In general, results support and provide improved capacities for validating experimental manipulation, remote sensing, and modeling studies.

To determine the influence of fire and thermokarst in a boreal landscape, we investigated peat cores within and adjacent to a permafrost collapse feature on the Tanana River Floodplain of Interior Alaska. Radioisotope dating, diatom assemblages, plant macrofossils, charcoal fragments, and carbon and nitrogen content of the peat profile indicate ∼600 years of vegetation succession with a transition from a terrestrial forest to a sedge-dominated wetland over 100 years ago, and to a Sphagnum-dominated peatland in approximately 1970. The shift from sedge to Sphagnum, and a de- crease in the detrended tree-ring width index of black spruce trees adjacent to the collapse coincided with an increase in the growing season temperature record from Fairbanks. This concurrent wetland succession and reduced growth of black spruce trees indicates a step-wise ecosystem-level response to a change in regional climate. In 2001, fire was observed coincident with permafrost collapse and resulted in lateral expansion of the peatland. These observations and the peat profile suggest that future warming and/or increased fire dis- turbance could promote permafrost degradation, peatland ex- pansion, and increase carbon storage across this landscape; however, the development of drought conditions could reduce the success of both black spruce and Sphagnum, and po- tentially decrease the long-term ecosystem carbon storage.

Dendroclimatological research is often based on the assumption that the relationship between tree growth and climate is not variable over time. Here we test this assumption by exploring if climate sensitivity of Picea mariana (Mill.) trees growing in open-stand lowland forest and on top of a neighboring peatland in Interior Alaska is stable or changing over time. Climate-growth correlations at the study sites are strongly dependent on microtopography and vary substantially over time. Trees growing in the open forest site generally display stronger climate-growth correlations, especially significantly negative correlations with late summer temperatures (July, August) starting in the period 1920-1970. Trees growing on the peatland site are less climate sensitive, but display positive correlations between annual growth and temperature of October and December in the early 20th century, while in the late 20th century, significant negative correlations exist with January and February temperatures. This study, thus, demonstrates a transient climate-growth response for P. mariana (Mill.) on two sites typical for lowland Interior Alaska. However, due to multiple possible explanations (e.g. changing climate, coupled with aging trees and a growing peatland surface) it is not possible at this time to pinpoint the exact cause for these changes in the climate-growth relationships.

We investigated long‐term and seasonal patterns of N imports and exports, as well as patterns following climate perturbations, across biomes using data from 15 watersheds from nine Long‐Term Ecological Research (LTER) sites in North America. Mean dissolved inorganic nitrogen (DIN) import-export budgets (N import via precipitation-N export via stream flow) for common years across all watersheds was highly variable, ranging from a net loss of − 0·17 ± 0·09 kg N ha−1mo−1 to net retention of 0·68 ± 0·08 kg N ha−1mo−1. The net retention of DIN decreased (smaller import-export budget) with increasing precipitation, as well as with increasing variation in precipitation during the winter, spring, and fall. Averaged across all seasons, net DIN retention decreased as the coefficient of variation (CV) in precipitation increased across all sites (r2 = 0·48, p = 0·005). This trend was made stronger when the disturbed watersheds were withheld from the analysis (r2 = 0·80, p < 0·001, n = 11). Thus, DIN exports were either similar to or exceeded imports in the tropical, boreal, and wet coniferous watersheds, whereas imports exceeded exports in temperate deciduous watersheds. In general, forest harvesting, hurricanes, or floods corresponded with periods of increased DIN exports relative to imports. Periods when water throughput within a watershed was likely to be lower (i.e. low snow pack or El Niño years) corresponded with decreased DIN exports relative to imports. These data provide a basis for ranking diverse sites in terms of their ability to retain DIN in the context of changing precipitation regimes likely to occur in the future.

We measured CO2 and CH4 exchange from the center of a Sphagnum‐dominated permafrost collapse, through an aquatic moat, and into a recently burned black spruce forest on the Tanana River floodplain in interior Alaska. In the anomalously dry growing season of 2004, both the collapse and the surrounding burned area were net sinks for CO2, with a mean daytime net ecosystem exchange of −1.4 μmol CO2 m−2 s−1, while the moat was a CH4 source with a mean flux of 0.013 μmol CH4 m−2 s−1. Regression analyses identified temperature as the dominant factor affecting intragrowing season variation in CO2 exchange and soil moisture as the primary control influencing CH4 emissions. CH4 emissions during the wettest portion of the growing season were four times higher than during the driest periods. If temperatures continue to warm, peatland vegetation will likely expand with permafrost degradation, resulting in greater carbon accumulation and methane emissions for the landscape as a whole.

Tundra ecosystems are sensitive to disturbance and slow to recover. To account for environmental costs of development in the North, cumulative impacts of roads and dust deposition must be quantified. After a previous study, we re-examined tundra adjacent to the 577-km-long Dalton Highway in northern Alaska to assess 13 y of additional calcareous road dust deposition. Dust loading continues to alter substrate properties and community composition. Moist, acidic, tussock-sedge tundra typically has a soil pH of 4. At the road margin the pH of the fibric horizon had increased to pH 5.5 by 1989 and to pH 6.0 by 2002. Plots adjacent to the road have significantly higher graminoid and Rubus chamaemorus biomass and less moss, evergreen shrub, lichen, and forb biomass. Graminoid cover ranges from 30% in undisturbed tundra to over 80% within 5 m of the road. We observed an 80 g·m−2 increase in graminoid biomass and a 130 g·m−2 decline in moss biomass across the study site between 1989 and 2002. Ordinations indicate a broadened zone of dust disturbance in 2002. This evidence of cumulative impacts of dust will improve our evaluation of the ecological costs of future road development in the North.

A removal experiment was conducted to measure how much and by what mechanisms brood parasitic Brown-headed Cowbirds (Molothrus ater) cause nest failures in a commonly used host, the Song Sparrow (Melospiza melodia). When numbers of female cowbirds were reduced experimentally, nest failures fell from 65.0% (n = 663 nests) to 49.9% (n = 331). Cowbird reduction reduced the frequency of nest failure to one-third of control levels in Song Sparrows during the last 80 days of the sparrow’s breeding season, the period when most parasitic laying took place. Cowbird reduction decreased nest failures strongly at the egg stage, and weakly at the nestling stage. Daily nest-failure rates were independent of whether or not a nest was parasitized by cowbirds. Two hypotheses were tested to explain how cowbirds cause host nests to fail: first, egg removal by female cowbirds lowers clutch size below a threshold where the host deserts; second, cowbirds cause host nests to fail by destroying entire clutches or broods. In support of the first hypothesis, desertion following parasitism and egg removal was less frequent when cowbird numbers were reduced (8.9% of n = 158 nests) than for unmanipulated controls (16.5% of n = 424 nests). In support of the second hypothesis, there were fewer cases where young were killed in the nest, or found dead near it, after cowbird numbers were reduced (2.5% of 158 nests) than in controls (4.7% of 424 control nests). In contrast, proportions of nests that failed after the disappearance of all eggs, young, or both, and after unparasitized clutches were deserted, increased when cowbird numbers were reduced. Although our study supports both hypotheses, cowbird-induced desertion had a greater effect on nest failure rates than did cowbird predation. Our study suggests that cowbird removal programs are likely to benefit commonly used and endangered hosts by reducing rates of nest failure.