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Spatial variation in plant chemical defence towards herbivores can help us understand variation in herbivore top–down control of shrubs in the Arctic and possibly also shrub responses to global warming. Less defended, non-resinous shrubs could be more influenced by herbivores than more defended, resinous shrubs. However, sparse field measurements limit our current understanding of how much of the circum-Arctic variation in defence compounds is explained by taxa or defence functional groups (resinous/non-resinous). We measured circum-Arctic chemical defence and leaf digestibility in resinous (Betula glandulosa, B. nana ssp. exilis) and non-resinous (B. nana ssp. nana, B. pumila) shrub birches to see how they vary among and within taxa and functional groups. Using liquid chromatography–mass spectrometry (LC–MS) metabolomic analyses and in vitro leaf digestibility via incubation in cattle rumen fluid, we analysed defence composition and leaf digestibility in 128 samples from 44 tundra locations.

We found biogeographical patterns in anti-herbivore defence where mean leaf triterpene concentrations and twig resin gland density were greater in resinous taxa and mean concentrations of condensing tannins were greater in non-resinous taxa. This indicates a biome-wide trade-off between triterpene- or tannin-dominated defences. However, we also found variations in chemical defence composition and resin gland density both within and among functional groups (resinous/non-resinous) and taxa, suggesting these categorisations only partly predict chemical herbivore defence. Complex tannins were the only defence compounds negatively related to in vitro digestibility, identifying this previously neglected tannin group as having a potential key role in birch anti-herbivore defence.

We conclude that circum-Arctic variation in birch anti-herbivore defence can be partly derived from biogeographical distributions of birch taxa, although our detailed mapping of plant defence provides more information on this variation and can be used for better predictions of herbivore effects on Arctic vegetation.

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.

Aim: Plant functional groups are widely used in community ecology and earth system modelling to describe trait variation within and across plant communities. However, this approach rests on the assumption that functional groups explain a large proportion of trait variation among species. We test whether four commonly used plant functional groups represent variation in six ecologically important plant traits.

Location: Tundra biome.

Time period: Data collected between 1964 and 2016.

Major taxa studied: 295 tundra vascular plant species.

Methods: We compiled a database of six plant traits (plant height, leaf area, specific leaf area, leaf dry matter content, leaf nitrogen, seed mass) for tundra species. We examined the variation in species-level trait expression explained by four traditional functional groups (evergreen shrubs, deciduous shrubs, graminoids, forbs), and whether variation explained was dependent upon the traits included in analysis. We further compared the explanatory power and species composition of functional groups to alternative classifications generated using post hoc clustering of species-level traits.

Results: Traditional functional groups explained significant differences in trait expression, particularly amongst traits associated with resource economics, which were consistent across sites and at the biome scale. However, functional groups explained 19% of overall trait variation and poorly represented differences in traits associated with plant size. Post hoc classification of species did not correspond well with traditional functional groups, and explained twice as much variation in species-level trait expression.

Main conclusions: Traditional functional groups only coarsely represent variation in well-measured traits within tundra plant communities, and better explain resource economic traits than size-related traits. We recommend caution when using functional group approaches to predict tundra vegetation change, or ecosystem functions relating to plant size, such as albedo or carbon storage. We argue that alternative classifications or direct use of specific plant traits could provide new insights for ecological prediction and modelling.

Question: Biotic resistance is defined as the reduction in invasion success caused by the native community through competition, herbivory and/or pathogens. Biotic resistance has mostly been studied during the initial stages of invasion. However, to what extent biotic resistance hampers survival, or persistence, of invaders in the longer term is often not known. We studied how native grassland communities affected growth, reproduction and survival during the adult life stage of the high-impact woody invader Chromolaena odorata under different water availability treatments.

Location: Hluhluwe-iMfolozi Park, South Africa.

Methods: We performed a 2-years full-factorial field experiment in a savanna grassland where we manipulated water availability and neighbouring vegetation; in one-third of the plots vegetation was cleared and planted with C.odorata monocultures, in one-third C.odorata shrubs were planted in grasslands, and one-third were unplanted grassland controls.

Results: Growth and reproduction of adult C.odorata were greatly reduced due to competition with native grasses, but not survival of C.odorata. Shrub survival was high and did not differ in plots with and without grass. Water availability did not affect growth, reproduction and competitive ability of C.odorata, but higher water availability did increase the competitive ability of the grasses. Soil moisture levels were lower in grasslands planted with C.odorata compared to unplanted controls, independent of the water treatment, suggesting higher water use of C.odorata compared to native grasses.

Conclusions: Savanna grasslands have a strong competitive effect on invasion by the exotic woody shrub C.odorata, reducing growth and reproduction, but not survival of the shrub. We found no evidence that biotic resistance was stronger under more unfavourable abiotic conditions, as C.odorata was equally impacted in all water treatments. The high survival rates of C.odorata suggest that competitive interactions are not likely to prevent invader persistence in the landscape. Invader persistence is important in determining longer-term invasion success as well as invader impact, and the concept of persistence should not be overlooked in studies on invasive species.

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 above mentioned article was originally scheduled for publication in the special issue on Ecology of Tundra Arthropods with guest editors Toke T. Hoye . Lauren E. Culler. Erroneously, the article was published in Polar Biology, Volume 40, Issue 11, November, 2017. The publisher sincerely apologizes to the guest editors and the authors for the inconvenience caused.

Motivation: The Tundra Trait Team (TTT) database includes field‐based measurements of key traits related to plant form and function at multiple sites across the tundra biome. This dataset can be used to address theoretical questions about plant strategy and trade‐offs, trait–environment relationships and environmental filtering, and trait variation across spatial scales, to validate satellite data, and to inform Earth system model parameters.

Main types of variable contained: The database contains 91,970 measurements of 18 plant traits. The most frequently measured traits (> 1,000 observations each) include plant height, leaf area, specific leaf area, leaf fresh and dry mass, leaf dry matter content, leaf nitrogen, carbon and phosphorus content, leaf C:N and N:P, seed mass, and stem specific density.

Spatial location and grain: Measurements were collected in tundra habitats in both the Northern and Southern Hemispheres, including Arctic sites in Alaska, Canada, Greenland, Fennoscandia and Siberia, alpine sites in the European Alps, Colorado Rockies, Caucasus, Ural Mountains, Pyrenees, Australian Alps, and Central Otago Mountains (New Zealand), and sub‐Antarctic Marion Island. More than 99% of observations are georeferenced.

Time period and grain: All data were collected between 1964 and 2018. A small number of sites have repeated trait measurements at two or more time periods.

Major taxa and level of measurement: Trait measurements were made on 978 terrestrial vascular plant species growing in tundra habitats. Most observations are on individuals (86%), while the remainder represent plot or site means or maximums per species.

Software format: csv file and GitHub repository with data cleaning scripts in R; contribution to TRY plant trait database (www.try-db.org) to be included in the next version release.

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 degrees 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.

Aim: Habitat loss and fragmentation is one of the main drivers of defaunation, that is the loss of large mammals. Biological invasions could be drivers of such phenomenon. However, their impact on large herbivore communities has not been studied to our knowledge. We made use of a landscape-scale control programme of one of the world's worst invaders, the shrub Chromolaena odorata, as a natural experiment to assess how this alien invader affects habitat use by 14 species of ungulates in an African savanna. Location: Hluhluwe-iMfolozi Park, KwaZulu-Natal, South Africa. Methods: At the height of the invasion in 2004, a large-scale control programme was initiated that successfully reduced densities of C.odorata across the park. We estimated mammalian herbivore habitat and patch use by dung counts and the presence and density of C.odorata along 24 line transects with a total length of 190km during the peak of the invasion (2004) and a decade after the initiation of a successful control programme (2014). To account for differences in herbivore assemblies between habitats and the preferential invasion of closed savanna woodlands, we analysed the recolonization of previously invaded patches by herbivores based on the change in dung abundance. Results: Herbivore species differed in how they responded to invaded patches of this non-native shrub. Grazers were the most negatively affected, especially those that avoid predators by running. Browsers were negatively impacted only at the highest invasive shrub densities. Some species, especially bushpig, positively selected invaded patches. Main conclusions: Large herbivores varied in their response to invasion with differences in impact depending on feeding strategy and predator avoidance strategy, but the majority of ungulates responded positively to the removal of this invasive shrub.