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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.

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.

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.

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.

Megaherbivores have been lost from most ecosystems world-wide, and current increases in poaching of rhino and elephant spp. threaten their status in the systems where they still occur. Although megaherbivores are said to be key drivers of ecosystem structure and functioning, empirical evidence is strongly biased to studies on African elephant. We urgently need a better understanding of the impact of other megaherbivore species to predict the consequences of megaherbivore loss. We used a unique 'recolonization experiment' to test how a megagrazer, white rhinoceros, is affecting the structure of savanna grasslands in Kruger National Park (KNP). With a 30-year record of rhinoceros distribution, we quantified how they recolonized KNP following their re-introduction. This allowed us to identify landscapes with high rhino densities and long time since recolonization versus landscapes with low rhino densities that were recolonized more recently but were otherwise biophysically similar. We recorded grassland heterogeneity on 40transects covering a total of 30km distributed across both landscapes. We used two proxies of grassland heterogeneity: % short grass cover and number of grazing lawn patches. Grazing lawns are patches with specific communities of prostrate-growing stoloniferous short grass species. Short grass cover was clearly higher in the high rhino impact (17.5%) than low rhino impact landscape (10.7%). Moreover, we encountered ~20 times more grazing lawns in the high rhino impact landscape. The effect of rhino on number of lawns and on short grass cover was similar to the two dominant geologies in KNP, basalt-derived versus granite-derived soils. Synthesis. We provide empirical evidence that white rhinoceros may have started to change the structure and composition of KNP's savanna grasslands. It remains to be tested if these changes lead to other ecological cascading effects. However, our results highlight that the current rhino poaching crisis may not only affect the species, but also threaten the potential key role of this megaherbivore as a driver of savanna functioning.

Biological invasions have increased dramatically in the past centuries and are one of the greatest threats to biodiversity today. Chromolaena odorata, a herbaceous shrub from the Americas, is one of the most widespread and problematic invasive plant species in the tropics and sub-tropics. The plant is a serious problem in South Africa, where invasive species threaten biodiversity and use up water resources. This study combines data on the distribution of C. odorata with ecological and clearing management data to evaluate the efficacy of an invasive plant clearing program over its decade of operation in the Hluhluwe-iMfolozi Park in KwaZulu-Natal, South Africa. Densities and local extent of the C. odorata invasion were significantly reduced during the period of operations of the clearing program. Seasonal effects impacted clearing efficacy, namely a reduction in efficacy during the seed dispersal period. Clearing success was positively associated with clearing effort and fire frequency and negatively associated with rainfall. Management implications drawn from the results include halting clearing during the seed-drop period, giving extra attention to areas with more rainfall and other water availability, and incorporating fire with other clearing methods where possible.

Aim A major challenge for invasion ecology is to identify high-impact invaders to guide prioritization of management interventions. We argue that species with the potential to cause regime shifts (altered states of ecosystem structure and function that are difficult or impossible to reverse) should be prioritized. These are species that modify ecosystems in ways that enhance their own persistence and suppress that of native species through reinforcing feedback processes. Methods Using both systems analysis and meta-analysis approaches, we synthesized changes to ecosystems caused by 173 invasive plant species. For the systems analysis, we examined published studies of impacts of invasive plants to determine which presented evidence consistent with a reinforcement of feedback processes. For the meta-analysis, we calculated the effect size ratio between standardized changes in recipient ecosystem and in the status of introduced species as an indication of a reinforcing feedback in particular species-environment combinations. The systems analysis approach allowed us to conceptualize regime shifts in invader-dominated landscapes and to estimate the likelihood of such changes occurring. The meta-analysis allowed us to quantitatively verify the conceptual model and the key invader-context feedbacks and to detect the strength and direction of feedbacks. Results Most reinforcing feedbacks involve impacts on soil-nutrient cycling by shrub and tree invaders in forests and herbaceous invaders in wetlands. Feedbacks resulting in regime shifts were most likely related to processes associated with seed banks, fire and nutrient cycling. Results were used to derive a key for identifying high-impact invaders. Main conclusions Identifying combinations of plant life-forms and ecosystems most likely to result in regime shifts is a robust approach for predicting high-impact invasions and therefore for prioritizing management interventions. The meta-analysis revealed the need for more quantitative studies, including manipulative experiments, on ecosystem feedbacks.

Introduction: Different climatic regions across the globe are associated with biomes that differ in their cover of woody plants, such as grasslands, savannas, and forests (Whitaker, 1962). For a long time, researchers have assumed that abiotic factors control the spatial distribution of woody plant-dominated ecosystems or biomes. According to this idea, rainfall and temperature determine the transitions from deserts to grasslands to savannas and eventually to forests (e.g., Prentice et al., 1992). However, we increasingly realize that biomes may be far less fixed entities than previously assumed. An alternative view for many regions might be that of “ecosystems uncertain,” which Whittaker (1975) defined as zones “in which either grassland or one of the types dominated by woody plants” may occur under the same climatic conditions. As Bond (2005) discusses, many of these “ecosystems uncertain” may be seen as “consumer-controlled ecosystems” where plant consumers, such as herbivores and fire, prevent a closed forest from developing and are a major determinant of the ecosystem state. Bond (2005) showed that such “ecosystems uncertain” may in fact cover a very large part of the world (Fig. 5.1). More recently, several global analyses confirmed that across a large part of the global land surface, tree cover is indeed bimodal (Staver et al., 2011) or even trimodal (Scheffer et al., 2012). This means that under the same climatic conditions, a system may be in a treeless, savanna, or forest state; this pattern has been described for (sub)tropical (Hirota et al., 2011; Staver et al., 2011), as well as boreal (Scheffer et al., 2012), parts of the world. From this, we can conclude that, rather than being purely controlled from the bottom up, ecosystems, and even biomes, are shaped by interacting bottom-up and top-down factors (Polis and Strong, 1996). There is general agreement that both bottom-up and top-down factors affect plant communities (Polis and Strong, 1996), but the question remains what the relative strengths of such top-down and bottom-up processes are and whether we can find general spatial and temporal patterns in their effects (Gripenberg and Roslin, 2007).

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.

Predicting impacts of global warming requires understanding of the extent to which plant biomass and production are controlled by bottom-up and top-down drivers. By annually monitoring community composition in grazed control plots and herbivore-free exclosures at an Arctic location for 15 years, we detected multiple biotic interactions. Regular rodent cycles acted as pulses driving synchronous fluctuations in the biomass of field-layer vegetation; reindeer influenced the biomass of taller shrubs, and the abundance of plant pathogenic fungi increased when densities of their host plants increased in exclosures. Two outbreaks of geometrid moths occurred during the study period, with contrasting effects on the field layer: one in 2004 had marginal effects, while one in 2012 severely reduced biomass in the control plots and eliminated biomass that had accumulated over 15 years in the exclosures. The latter was followed by a dramatic decline of the dominant understory dwarf-shrub Empetrum hermaphroditum, driven by an interaction between moth herbivory on top buds and leaves, and increased disease severity of a pathogenic fungus. We show that the climate has important direct and indirect effects on all these biotic interactions. We conclude that long time series are essential to identify key biotic interactions in ecosystems, since their importance will be influenced by climatic conditions, and that manipulative treatments are needed in order to obtain the mechanistic understanding needed for robust predictions of future ecosystem changes and their feedback effects.

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.

Herbivores impact nutrient availability and cycling, and the net effect of herbivory on soil nutrients is generally assumed to be positive in nutrient-rich environments and negative in nutrient-poor ones. This is, however, far from a uniform pattern, and there is a recognized need to investigate any interactive effects of herbivory and habitat fertility (i.e., plant C/N ratios) on soil nutrient availabilities. We determined long-term effects of reindeer on soil extractable nitrogen (N) and phosphorus (P) and their net mineralization rates along a fertility gradient of plant carbon (C) to N and P ratios in arctic tundra. Our results showed that reindeer had a positive effect on soil N in the more nutrient-poor sites and a negative effect on soil P in the more nutrient-rich sites, which contrasts from the general consensus. The increase in N availability was linked to a decrease in plant and litter C/N ratios, suggesting that a shift in vegetation composition toward more graminoids favors higher N cycling. Soil P availability was not as closely linked to the vegetation and is likely regulated more by herbivore-induced changes in soil physical and chemical properties. The changes in soil extractable N and P resulted in higher soil N/P ratios, suggesting that reindeer could drive the vegetation toward P-limitation. This research highlights the importance of including both the elements N and P and conducting studies along environmental gradients in order to better understand the interactive effects of herbivory and habitat fertility on nutrient cycling and primary production.

Climatic niche models based on native-range climatic data accurately predict invasive-range distributions in the majority of species. However, these models often do not account for ecological and evolutionary processes, which limit the ability to predict future range expansion. This might be particularly problematic in the case of invaders that occupy environments that would be considered marginal relative to the climatic niche in the native range of the species. Here, we assess the potential for future range expansion in the shrub Chromolaena odorata that is currently invading mesic savannas (>650 mm MAP) in South Africa that are colder and drier than most habitats in its native range. In a greenhouse experiment we tested whether its current distribution in South Africa can be explained by increased competitive ability and/or differentiation in drought tolerance relative to the native population. We compared aboveground biomass, biomass allocation, water use efficiency and relative yields of native and invasive C. odorata and the resident grass Panicum maximum in wet and dry conditions. Surprisingly, we found little differentiation between ranges. Invasive C. odorata showed no increased competitive ability or superior drought tolerance compared to native C. odorata. Moreover we found that P. maximum was a better competitor than either native or invasive C. odorata. These results imply that C. odorata is unlikely to expand its future range towards more extreme, drier, habitats beyond the limits of its current climatic niche and that the species' invasiveness most likely depends on superior light interception when temporarily released from competition by disturbance. Our study highlights the fact that species can successfully invade habitats that are at the extreme end of their ranges and thereby contributes towards a better understanding of range expansion during species invasions.

Our understanding of the link between plant functional traits and ecological impact of invasive alien plant species is fragmentary and the mechanisms leading to impacts are poorly understood. Moreover, current knowledge is heavily biased to the temperate regions of the world and we know much less about traits and impacts of invaders in tropical and subtropical ecosystems. We studied two leaf traits of the invasive alien shrub Chromolaena odorata and the impacts of its invasion on native vegetation in savannas. We compared specific leaf area (SLA) and leaf area index (LAI) between C. odorata and native species and assessed how C. odorata differentially affects canopy light interception, soil moisture, soil nutrients, and litter accumulation compared to native species. We found that C. odorata has higher SLA and LAI than native species, lower light and moisture levels below its canopy, but higher nutrient levels and a higher litter accumulation rate. Because of its higher SLA, C. odorata grows faster, resulting in more biomass, increased litter accumulation and higher nutrient availability. Due to its high SLA and LAI, C. odorata intercepts more light and reduces available moisture more than do native trees due to higher transpiration rates, reducing the biomass of native understory vegetation. This study provides empirical evidence for strong links between plant functional traits and ecological impact of invasive plant species, highlighting the importance of traits in predicting ecosystem-level impacts of invasive plant species.

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.

Question: Resistance of the native community has been identified as an important factor limiting invasion success and invader impact. However, to what extent resistance interacts with disturbance to control invasion success remains unclear. We studied the interaction between biotic resistance, fire and small-scale disturbances mimicking those of large mammalian herbivores (hoof action and grazing) on invasion success of the alien shrub Chromolaena odorata.

Location: Hluhluwe-iMfolozi Park, South Africa.

Methods: We performed a seedling transplant experiment in a savanna grassland. We manipulated the grass layer by clipping (grazing), created small-scale soil disturbances (hoof action) and transplanted seedlings under native tree canopies to assess potential nursing effects. Subsequently, we burned half of our plots. We assessed the effect of fire and small-scale disturbances on seedling growth, survival and biomass allocation.

Results: Seedling survival of C. odorata was 26% in undisturbed savanna grassland, reducing to 5% post-fire. Small-scale disturbances increased seedling biomass and survival and modified biomass allocation, whereas fire greatly reduced seedling survival. Root allocation increased in response to grass clipping, while stem allocation decreased. Tree shading increased seedling survival in the absence of fire, but greatly reduced post-fire survival.

Conclusions: Grass communities in savannas exert a high level of biotic resistance to C. odorata invasion. However, small-scale disturbances, mimicking those of large mammalian herbivores, can facilitate the invasion of C. odorata in savanna grassland both directly, by creating micro-sites for establishment, and indirectly, by reducing the negative effect of fire. Thus, small-scale disturbances may aid the long-term persistence of woody species invading grass-dominated systems.

Previous studies have shown that climate warming is causing shrub cover to increase at high latitudes. Increased shrub cover generally lowers surface albedo, which results in higher energy absorption and further warming. In parts of Fennoscandia, herbivory is known to control vegetation height and abundance, and thus preventing this positive feedback. Here, we combine field measurements of albedo, herbivory and vegetation characteristics in four topographically-defined vegetation types of varying shrub height and abundance with land surface modeling (JULES) to investigate if reindeer grazing can influence the energy balance of an arctic tundra. We find that when reindeer reduces shrub height and abundance, summer albedo increases in both Betula nana-dominated heath vegetation and Salix glauca-dominated willow depressions. Model results reveal associated lower net radiation, and latent and sensible heat fluxes in heavily-grazed sites in all shrub-dominated vegetation types. Our results also suggest that the structural shift from graminoid to shrub tundra drives the difference in summer albedo, rather than shifts from dwarf-shrub to tall-shrub tundra. Reindeer has thus a potential cooling effect on climate by increasing summer albedo and decreasing net radiation, which highlights the importance of mammalian herbivores for the earth system beyond their local grazing impacts. However, the strong effects of reindeer on albedo are probably restricted to areas with high reindeer densities, since a dramatic vegetation change is essential. The importance of these processes across the whole range of reindeer densities found in the arctic tundra needs to be further evaluated.

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.