Global environmental change effects on ecosystems: the importance of land‐use legacies

One of the major challenges in ecology is to predict how multiple global environmental changes will affect future ecosystem patterns (e.g. plant community composition) and processes (e.g. nutrient cycling). Here, we highlight arguments for the necessary inclusion of land‐use legacies in this endeavour. Alterations in resources and conditions engendered by previous land use, together with influences on plant community processes such as dispersal, selection, drift and speciation, have steered communities and ecosystem functions onto trajectories of change. These trajectories may be modulated by contemporary environmental changes such as climate warming and nitrogen deposition. We performed a literature review which suggests that these potential interactions have rarely been investigated. This crucial oversight is potentially due to an assumption that knowledge of the contemporary state allows accurate projection into the future. Lessons from other complex dynamic systems, and the recent recognition of the importance of previous conditions in explaining contemporary and future ecosystem properties, demand the testing of this assumption. Vegetation resurvey databases across gradients of land use and environmental change, complemented by rigorous experiments, offer a means to test for interactions between land‐use legacies and multiple environmental changes. Implementing these tests in the context of a trait‐based framework will allow biologists to synthesize compositional and functional ecosystem responses. This will further our understanding of the importance of land‐use legacies in determining future ecosystem properties, and soundly inform conservation and restoration management actions.     

Four decades of plant community change along a continental gradient of warming

Many studies of individual sites have revealed biotic changes consistent with climate warming (e.g., upward elevational distribution shifts), but our understanding of the tremendous variation among studies in the magnitude of such biotic changes is minimal. In this study, we resurveyed forest vegetation plots 40 years after the initial surveys in three protected areas along a west‐to‐east gradient of increasingly steep recent warming trends in eastern Canada (Québec). Consistent with the hypothesis that climate warming has been an important driver of vegetation change, we found an increasing magnitude of changes in species richness and composition from west to east among the three parks. For the two mountainous parks, we found no significant changes in elevational species’ distributions in the easternmost park (raw mean = +11.4 m at Forillon Park) where warming has been minimal, and significant upward distribution shifts in the centrally located park (+38.9 m at Mont‐Mégantic), where the recent warming trend has been marked. Community Temperature Indices (CTI), reflecting the average affinities of locally co‐occurring species to temperature conditions across their geographic ranges (“Species Temperature Indices”), did not change over time as predicted. However, close examination of the underpinnings of CTI values suggested a high sensitivity to uncertainty in individual species’ temperature indices, and so a potentially limited responsiveness to warming. Overall, by testing a priori predictions concerning variation among parks in the direction and magnitude of vegetation changes, we have provided stronger evidence for a link between climate warming and biotic responses than otherwise possible and provided a potential explanation for large variation among studies in warming‐related biotic changes.     

Global change pressures on soils from land use and management

Soils are subject to varying degrees of direct or indirect human disturbance, constituting a major global change driver. Factoring out natural from direct and indirect human influence is not always straightforward, but some human activities have clear impacts. These include land‐use change, land management and land degradation (erosion, compaction, sealing and salinization). The intensity of land use also exerts a great impact on soils, and soils are also subject to indirect impacts arising from human activity, such as acid deposition (sulphur and nitrogen) and heavy metal pollution. In this critical review, we report the state‐of‐the‐art understanding of these global change pressures on soils, identify knowledge gaps and research challenges and highlight actions and policies to minimize adverse environmental impacts arising from these global change drivers. Soils are central to considerations of what constitutes sustainable intensification. Therefore, ensuring that vulnerable and high environmental value soils are considered when protecting important habitats and ecosystems, will help to reduce the pressure on land from global change drivers. To ensure that soils are protected as part of wider environmental efforts, a global soil resilience programme should be considered, to monitor, recover or sustain soil fertility and function, and to enhance the ecosystem services provided by soils. Soils cannot, and should not, be considered in isolation of the ecosystems that they underpin and vice versa. The role of soils in supporting ecosystems and natural capital needs greater recognition. The lasting legacy of the International Year of Soils in 2015 should be to put soils at the centre of policy supporting environmental protection and sustainable development.     

Trait variations of ground flora species disentangle the effects of global change and altered land‐use in Swedish forests during 20 years

Northern forest ecosystems are exposed to a range of anthropogenic processes including global warming, atmospheric deposition, and changing land‐use. The vegetation of northern forests is composed of species with several functional traits related to these processes, whose effects may be difficult to disentangle. Here, we combined analyses of spatio‐temporal dynamics and functional traits of ground flora species, including morphological characteristics, responses to macro‐ and microclimate, soil conditions, and disturbance. Based on data from the Swedish National Forest Inventory, we compared changes in occurrence of a large number of ground flora species during a 20‐year period (1994–2013) in boreal and temperate Sweden respectively. Our results show that a majority of the common ground flora species have changed their overall frequency. Comparisons of functional traits between increasing and declining species, and of trends in mean trait values of sample plots, indicate that current floristic changes are caused by combined effects of climate warming, nitrogen deposition and changing land‐use. Changes and their relations with plant traits were generally larger in temperate southern Sweden. Nutrient‐demanding species with mesotrophic morphology were favored by ongoing eutrophication due to nitrogen deposition in the temperate zone, while dwarf shrubs with low demands on nitrogen decreased in frequency. An increase of species with less northern and less eastern distribution limits was also restricted to temperate Sweden, and indicates effects of a moister and milder macroclimate. A trend toward dense plantation forests is mirrored by a decrease of light‐demanding species in both vegetation zones, and a decrease of grassland species in the temperate zone. Although denser tree canopies may buffer effects of a warmer climate and of nitrogen deposition to some extent, traits related to these processes were weakly correlated in the group of species with changing frequency. Hence, our results indicate specific effects of these often confounded anthropogenic processes.     

Future climate change effects on US forest composition may offset benefits of reduced atmospheric deposition of N and S

Climate change and atmospheric deposition of nitrogen (N) and sulfur (S) are important drivers of forest demography. Here we apply previously derived growth and survival responses for 94 tree species, representing >90% of the contiguous US forest basal area, to project how changes in mean annual temperature, precipitation, and N and S deposition from 20 different future scenarios may affect forest composition to 2100. We find that under the low climate change scenario (RCP 4.5), reductions in aboveground tree biomass from higher temperatures are roughly offset by increases in aboveground tree biomass from reductions in N and S deposition. However, under the higher climate change scenario (RCP 8.5) the decreases from climate change overwhelm increases from reductions in N and S deposition. These broad trends underlie wide variation among species. We found averaged across temperature scenarios the relative abundance of 60 species were projected to decrease more than 5% and 20 species were projected to increase more than 5%; and reductions of N and S deposition led to a decrease for 13 species and an increase for 40 species. This suggests large shifts in the composition of US forests in the future. Negative climate effects were mostly from elevated temperature and were not offset by scenarios with wetter conditions. We found that by 2100 an estimated 1 billion trees under the RCP 4.5 scenario and 20 billion trees under the RCP 8.5 scenario may be pushed outside the temperature record upon which these relationships were derived. These results may not fully capture future changes in forest composition as several other factors were not included. Overall efforts to reduce atmospheric deposition of N and S will likely be insufficient to overcome climate change impacts on forest demography across much of the United States unless we adhere to the low climate change scenario.     

The impact of herbivore exclusion on forb diversity: Comparing species and functional responses during a drought

Rainfall and herbivory shape savannah herbaceous communities, but these disturbances are being altered globally. To assess potential consequences of such alterations, we evaluated herbivore effects on species and functional diversity during an episodic drought in a sodic savannah using data collected from long-term herbivore exclosures in the Kruger National Park, South Africa. Herbaceous life forms are rarely acknowledged as distinct functional entities. Moreover, the functional ecology of forbs remains elusive. Here, we present disturbances–responses by forbs separately from grasses. We hypothesised that combinations of intense utilisation and drought would be associated with low diversity and high dominance at species and functional levels for both life forms. Contrary to our hypothesis, low forb and grass diversity was associated with long-term herbivore exclusion, which exceeded expected undesirable effects of intense utilisation and drought. Grasses responded less sensitively, suggesting that forbs respond dynamically to changes in herbivore assemblage when these alterations are combined with drought. Consistent with patterns in savannah systems, forbs contributed significantly to species and functional trait diversity. High forb diversity is suggested to enhance resilience of this nutrient-rich ecosystem against declines in its functioning when subjected to drought and alterations in herbivory.     

Potential effects of warming and drying on peatland plant community composition

Boreal peatlands may be particularly vulnerable to climate change, because temperature regimes that currently constrain biological activity in these regions are predicted to increase substantially within the next century. Changes in peatland plant community composition in response to climate change may alter nutrient availability, energy budgets, trace gas fluxes, and carbon storage. We investigated plant community response to warming and drying in a field mesocosm experiment in northern Minnesota, USA. Large intact soil monoliths removed from a bog and a fen received three infrared warming treatments crossed with three water‐table treatments (n = 3) for five years. Foliar cover of each species was estimated annually. In the bog, increases in soil temperature and decreases in water‐table elevation increased cover of shrubs by 50% and decreased cover of graminoids by 50%. The response of shrubs to warming was distinctly species‐specific, and ranged from increases (for Andromeda glaucophylla) to decreases (for Kalmia polifolia). In the fens, changes in plant cover were driven primarily by changes in water‐table elevation, and responses were species‐ and lifeform‐specific: increases in water‐table elevation increased cover of graminoids – in particular Carex lasiocarpa and Carex livida– as well as mosses. In contrast, decreases in water‐table elevation increased cover of shrubs, in particular A. glaucophylla and Chamaedaphne calyculata. The differential and sometimes opposite response of species and lifeforms to the treatments suggest that the structure and function of both bog and fen plant communities will change – in different directions or at different magnitudes – in response to warming and/or changes in water‐table elevation that may accompany regional or global climate change.     

Individualistic responses of forest herb traits to environmental change

• Intraspecific trait variation (ITV; i.e. variability in mean and/or distribution of plant attribute values within species) can occur in response to multiple drivers. Environmental change and land‐use legacies could directly alter trait values within species but could also affect them indirectly through changes in vegetation cover. Increasing variability in environmental conditions could lead to more ITV, but responses might differ among species. Disentangling these drivers on ITV is necessary to accurately predict plant community responses to global change.
• We planted herb communities into forest soils with and without a recent history of agriculture. Soils were collected across temperate European regions, while the 15 selected herb species had different colonizing abilities and affinities to forest habitat. These mesocosms (384) were exposed to two‐level full‐factorial treatments of warming, nitrogen addition and illumination. We measured plant height and specific leaf area (SLA).
• For the majority of species, mean plant height increased as vegetation cover increased in response to light addition, warming and agricultural legacy. The coefficient of variation (CV) for height was larger in fast‐colonizing species. Mean SLA for vernal species increased with warming, while light addition generally decreased mean SLA for shade‐tolerant species. Interactions between treatments were not important predictors.
• Environmental change treatments influenced ITV, either via increasing vegetation cover or by affecting trait values directly. Species’ ITV was individualistic, i.e. species responded to different single resource and condition manipulations that benefited their growth in the short term. These individual responses could be important for altered community organization after a prolonged period.

     

Permafrost condition determines plant community composition and community‐level foliar functional traits in a boreal peatland

Boreal peatlands are critical ecosystems globally because they house 30%–40% of terrestrial carbon (C), much of which is stored in permafrost soil vulnerable to climate warming‐induced thaw. Permafrost thaw leads to thickening of the active (seasonally thawed) layer and alters nutrient and light availability. These physical changes may influence community‐level plant functional traits through intraspecific trait variation and/or species turnover. As permafrost thaw is expected to cause an efflux of carbon dioxide (CO₂) and methane (CH₄) from the soil to the atmosphere, it is important to understand thaw‐induced changes in plant community productivity to evaluate whether these changes may offset some of the anticipated increases in C emissions. To this end, we collected vascular plant community composition and foliar functional trait data along gradients in aboveground tree biomass and active layer thickness (ALT) in a rapidly thawing boreal peatland, with the expectation that changes in above‐ and belowground conditions are indicative of altered resource availability. We aimed to determine whether community‐level traits vary across these gradients, and whether these changes are dominated by intraspecific trait variation, species turnover, or both. Our results highlight that variability in community‐level traits was largely attributable to species turnover and that both community composition and traits were predominantly driven by ALT. Specifically, thicker active layers associated with permafrost‐free peatlands (i.e., bogs and fens) shifted community composition from slower‐growing evergreen shrubs to faster‐growing graminoids and forbs with a corresponding shift toward more productive trait values. The results from this rapidly thawing peatland suggest that continued warming‐induced permafrost thaw and thermokarst development alter plant community composition and community‐level traits and thus ecosystem productivity. Increased productivity may help to mitigate anticipated CO₂ efflux from thawing permafrost, at least in the short term, though this response may be swamped by increase CH₄ release.     

Patterns and drivers of plant functional group dominance across the Western Hemisphere: a macroecological re‐assessment based on a massive botanical dataset

Plant functional group dominance has been linked to climate, topography and anthropogenic factors. Here, we assess existing theory linking functional group dominance patterns to their drivers by quantifying the spatial distribution of plant functional groups at a 100‐km grid scale. We use a standardized plant species occurrence dataset of unprecedented size covering the entire New World. Functional group distributions were estimated from 3 648 533 standardized occurrence records for a total of 83 854 vascular plant species, extracted from the Botanical Information and Ecology Network (BIEN) database. Seven plant functional groups were considered, describing major differences in structure and function: epiphytes; climbers; ferns; herbs; shrubs; coniferous trees; and angiosperm trees. Two measures of dominance (relative number of occurrences and relative species richness) were analysed against a range of hypothesized predictors. The functional groups showed distinct geographical patterns of dominance across the New World. Temperature seasonality and annual precipitation were most frequently selected, supporting existing hypotheses for the geographical dominance of each functional group. Human influence and topography were secondarily important. Our results support the prediction that future climate change and anthropogenic pressures could shift geographical patterns in dominance of plant functional groups, with probable consequences for ecosystem functioning. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 180 , 141–160.     

Effects of plant traits on ecosystem and regional processes: a conceptual framework for predicting the consequences of global change

Human activities are causing widespread changes in the species composition of natural and managed ecosystems, but the consequences of these changes are poorly understood. This paper presents a conceptual framework for predicting the ecosystem and regional consequences of changes in plant species composition. Changes in species composition have greatest ecological effects when they modify the ecological factors that directly control (and respond to) ecosystem processes. These interactive controls include: functional types of organisms present in the ecosystem; soil resources used by organisms to grow and reproduce; modulators such as microclimate that influence the activity of organisms; disturbance regime; and human activities. Plant traits related to size and growth rate are particularly important because they determine the productive capacity of vegetation and the rates of decomposition and nitrogen mineralization. Because the same plant traits affect most key processes in the cycling of carbon and nutrients, changes in plant traits tend to affect most biogeochemical cycling processes in parallel. Plant traits also have landscape and regional effects through their effects on water and energy exchange and disturbance regime.     

Disentangling the effects of plant species invasion and urban development on arthropod community composition

Urban development and species invasion are two major global threats to biodiversity. These threats often co‐occur, as developed areas are more prone to species invasion. However, few empirical studies have tested if both factors affect biodiversity in similar ways. Here we study the individual and combined effects of urban development and plant invasion on the composition of arthropod communities. We assessed 36 paired invaded and non‐invaded sample plots, invaded by the plant Antigonon leptopus, with half of these pairs located in natural and the other half in developed land‐use types on the Caribbean island of St. Eustatius. We used several taxonomic and functional variables to describe community composition and diversity. Our results show that both urban development and A. leptopus invasion affected community composition, albeit in different ways. Development significantly increased species richness and exponential Shannon diversity, while invasion had no effect on these variables. However, invasion significantly increased arthropod abundance and caused biotic homogenization. Specifically, uninvaded arthropod communities were distinctly different in species composition between developed and natural sites, while they became undistinguishable after A. leptopus invasion. Moreover, functional variables were significantly affected by species invasion, but not by urban development. Invaded communities had higher community‐weighted mean body size and the feeding guild composition of invaded arthropod communities was characterized by the exceptional numbers of nectarivores, herbivores, and detritivores. With the exception of species richness and exponential Shannon diversity, invasion influenced four out of six response variables to a greater degree than urban development did. Hence, we can conclude that species invasion is not just a passenger of urban development but also a driver of change.     

Assessing the effects of land-use change on plant traits, communities and ecosystem functioning in grasslands: a standardized methodology and lessons from an application to 11 European sites

BACKGROUND AND AIMS: A standardized methodology to assess the impacts of land-use changes on vegetation and ecosystem functioning is presented. It assumes that species traits are central to these impacts, and is designed to be applicable in different historical, climatic contexts and local settings. Preliminary results are presented to show its applicability. METHODS: Eleven sites, representative of various types of land-use changes occurring in marginal agro-ecosystems across Europe and Israel, were selected. Climatic data were obtained at the site level; soil data, disturbance and nutrition indices were described at the plot level within sites. Sixteen traits describing plant stature, leaf characteristics and reproductive phase were recorded on the most abundant species of each treatment. These data were combined with species abundance to calculate trait values weighed by the abundance of species in the communities. The ecosystem properties selected were components of above-ground net primary productivity and decomposition of litter. KEY RESULTS: The wide variety of land-use systems that characterize marginal landscapes across Europe was reflected by the different disturbance indices, and were also reflected in soil and/or nutrient availability gradients. The trait toolkit allowed us to describe adequately the functional response of vegetation to land-use changes, but we suggest that some traits (vegetative plant height, stem dry matter content) should be omitted in studies involving mainly herbaceous species. Using the example of the relationship between leaf dry matter content and above-ground dead material, we demonstrate how the data collected may be used to analyse direct effects of climate and land use on ecosystem properties vs. indirect effects via changes in plant traits. CONCLUSIONS: This work shows the applicability of a set of protocols that can be widely applied to assess the impacts of global change drivers on species, communities and ecosystems.     

Vegetation change at high elevation: scale dependence and interactive effects on Niwot Ridge

Background: High-elevation mountain systems may be particularly responsive to climate change.

Aims: Here we investigate how changes along elevation gradients in mountain systems can aid in predicting vegetation distributional changes in time, focusing on how changing climatic controls affect meso-scale transitions at the lower and upper boundaries of alpine vegetation (with forest and subnival zones, respectively) as well as micro-scale transitions among plant communities within the alpine belt. We focus on climate-related drivers, particularly in relation to climate change, but also consider how species interactions, dispersal and responses to disturbance may influence plant responses to these abiotic drivers.

Results: Empirical observations and experimental studies indicate that changing climatic controls influence both meso-scale transitions at the upper and lower boundaries of alpine vegetation and micro-scale transitions among plant communities within tundra. Micro-scale heterogeneity appears to buffer response in many cases, while interactions between climate and other changes may often accelerate change.

Conclusions: Interactions with microtopography and larger edaphic gradients have the capacity to both facilitate rapid changes and reinforce stability, and that these interactions will affect the responsiveness of vegetation to climate change at different spatial scales.     

Interactive effects between plant functional types and soil factors on tundra species diversity and community composition

Plant communities are coupled with abiotic factors, as species diversity and community composition both respond to and influence climate and soil characteristics. Interactions between vegetation and abiotic factors depend on plant functional types (PFT) as different growth forms will have differential responses to and effects on site characteristics. However, despite the importance of different PFT for community assembly and ecosystem functioning, research has mainly focused on vascular plants. Here, we established a set of observational plots in two contrasting habitats in northeastern Siberia in order to assess the relationship between species diversity and community composition with soil variables, as well as the relationship between vegetation cover and species diversity for two PFT (nonvascular and vascular). We found that nonvascular species diversity decreased with soil acidity and moisture and, to a lesser extent, with soil temperature and active layer thickness. In contrast, no such correlation was found for vascular species diversity. Differences in community composition were found mainly along soil acidity and moisture gradients. However, the proportion of variation in composition explained by the measured soil variables was much lower for nonvascular than for vascular species when considering the PFT separately. We also found different relationships between vegetation cover and species diversity according the PFT and habitat. In support of niche differentiation theory, species diversity and community composition were related to edaphic factors. The distinct relationships found for nonvascular and vascular species suggest the importance of considering multiple PFT when assessing species diversity and composition and their interaction with edaphic factors. Synthesis: Identifying vegetation responses to edaphic factors is a first step toward a better understanding of vegetation–soil feedbacks under climate change. Our results suggest that incorporating differential responses of PFT is important for predicting vegetation shifts, primary productivity, and in turn, ecosystem functioning in a changing climate.     

The effect of biochar management on soil and plant community properties in a boreal forest

Biochar management has been proposed as a possible tool to mitigate anthropogenic CO₂ emissions, and thus far its impacts in forested environments remain poorly understood. We conducted a large‐scale, replicated field experiment using 0.05‐ha plots in the boreal region in northern Sweden to evaluate how soil and vegetation properties and processes responded to biochar application and the disturbance associated with burying biochar in the soil. We employed a randomized block design, where biochar and soil mixing treatments were established in factorial combination (i.e., control, soil mixing only, biochar only, and biochar and soil mixing; n = 6 plots of each). After two growing seasons, we found that biochar application enhanced net soil N mineralization rates and soil concentrations regardless of the soil mixing treatment, but had no impact on the availability of , the majority of soil microbial community parameters, or soil respiration. Meanwhile, soil mixing enhanced soil concentrations, but had negative impacts on net N mineralization rates and several soil microbial community variables. Many of the effects of soil mixing on soil nutrient and microbial community properties were less extreme when biochar was also added. Biochar addition had almost no effects on vegetation properties (except for a small reduction in species richness of the ground layer vegetation), while soil mixing caused significant reductions in graminoid and total ground layer vegetation cover, and enhanced seedling survival rates of P. sylvestris, and seed germination rates for four tree species. Our results suggest that biochar application can serve as an effective tool to store soil C in boreal forests while enhancing availability. They also suggest that biochar may serve as a useful complement to site preparation techniques that are frequently used in the boreal region, by enhancing soil fertility and reducing nutrient losses when soils are scarified during site preparation.