Status and trends in Arctic vegetation: Evidence from experimental warming and long-term monitoring

Changes in Arctic vegetation can have important implications for trophic interactions and ecosystem functioning leading to climate feedbacks. Plot-based vegetation surveys provide detailed insight into vegetation changes at sites around the Arctic and improve our ability to predict the impacts of environmental change on tundra ecosystems. Here, we review studies of changes in plant community composition and phenology from both long-term monitoring and warming experiments in Arctic environments. We find that Arctic plant communities and species are generally sensitive to warming, but trends over a period of time are heterogeneous and complex and do not always mirror expectations based on responses to experimental manipulations. Our findings highlight the need for more geographically widespread, integrated, and comprehensive monitoring efforts that can better resolve the interacting effects of warming and other local and regional ecological factors.     

Tundra in the rain: differential vegetation responses to three years of experimentally doubled summer precipitation in Siberian shrub and Swedish bog tundra

Precipitation amounts and patterns at high latitude sites have been predicted to change as a result of global climatic changes. We addressed vegetation responses to three years of experimentally increased summer precipitation in two previously unaddressed tundra types: Betula nana-dominated shrub tundra (northeast Siberia) and a dry Sphagnum fuscum-dominated bog (northern Sweden). Positive responses to approximately doubled ambient precipitation (an increase of 200 mm year(-1)) were observed at the Siberian site, for B. nana (30 % larger length increments), Salix pulchra (leaf size and length increments) and Arctagrostis latifolia (leaf size and specific leaf area), but none were observed at the Swedish site. Total biomass production did not increase at either of the study sites. This study corroborates studies in other tundra vegetation types and shows that despite regional differences at the plant level, total tundra plant productivity is, at least at the short or medium term, largely irresponsive to experimentally increased summer precipitation.     

Ozone flux to vegetation and its relationship to plant response and ambient air quality standards

The National Ambient Air Quality Standard (NAAQS) for ozone is based on occurrences of the maximum 8 h average ambient ozone concentration. However, biologists have recommended a cumulative ozone exposure parameter to protect vegetation. In this paper we propose a third alternative which uses quantifiable flux-based numerical parameters as a replacement for cumulative ambient parameters. Herein we discuss the concept of ozone flux as it relates to plant response and the NAAQS, and document information needed before a flux-based ozone NAAQS for vegetation can be implemented. Additional research is needed in techniques for determining plant uptake and in the quantification of plant defensive mechanisms to ozone. Models which include feedback mechanisms should be developed to relate ozone flux, loading, and detoxification with photosynthesis and plant productivity.     

Satellite image analysis of human caused changes in the tundra vegetation around the city of Vorkuta,north-European Russia

In this paper we present what kind of human impacted changes can be found in dwarf birch (Betula nana) dominated shrub tundra vegetation around the large industrial complex of Vorkuta in the north-European Russian tundra. Using fieldwork data and Landsat TM satellite image we could identify two impact zones: (1) Pollution zone (150-200 km2). In this zone most of the lichen species are absent. Changes in vegetation communities' species composition in all main plant groups are obvious. Willows especially are more dominant than in the unpolluted sites. (2) Slight pollution/disturbance zone (600-900 km2). Here vegetation changes are mainly similar but less so than the changes in the first zone. Particularly, the amount of herbs and grasses is increased when compared to unpolluted areas. The pollution zones are spatially connected to the main emission sources in the area. Zones spread furthest to the northeast, matching the prevailing winds during winter.     

Modelling the long-term soil response to atmospheric deposition at intensively monitored forest plots in Europe

The dynamic soil chemistry model SMART was applied to 121 intensive forest monitoring plots (mainly located in western and northern Europe) for which both element input (deposition) and element concentrations in the soil solution were available. After calibration of poorly known parameters, the model accurately simulated soil solution concentrations for most plots as indicated by goodness-of-fit measures, although some of the intra-annual variation especially in nitrate and aluminium concentrations could not be reproduced. Model evaluations of two emission-deposition scenarios (current legislation and maximum feasible reductions) for the period 1970-2030 show a strong reduction in sulphate concentrations between 1980 and 2000 in the soil due to the high reductions in sulphur emissions. However, current legislation hardly reduces future nitrogen concentrations, whereas maximum feasible reductions reduces them by more than half. Maximum feasible reductions are also more effective in increasing pH and reducing aluminium concentrations, mostly below ‘critical’ values.     

An analysis of numerical models of air pollutant exposure and vegetation response

A number of empirical (statistical, regression oriented) and mechanistic (process oriented) models are presently available to examine the relationship between air pollution stress and plant response. These models have their strengths and weaknesses. In all these models, a major concern is the numerical definition of the pollutant exposure kinetics (dose). At present there are no numerical definitions of dose which make satisfactory biological sense. A key issue is the existence of a biological time clock where plants respond differently to the pollutant stress at different stages of their growth. On the other hand, policy makers and regulatory personnel prefer a simple approach which would facilitate implementation and administration of ambient air quality standards. Long-term air pollutant averaging techniques create artifacts due to the non-normal distribution of ambient concentrations. A more appropriate approach may be the use of 'median' and 'percentiles' computed from short duration pollutant concentrations. Such an approach would be free of the influence of the non-normal distribution, but would require the development of appropriate exposure-response models. Any transfer of results from unit level models to regional level leads to 'scaling error'. There is no general agreement among researchers on how to deal with the scale problem. While this situation persists, any policy formulated on regional impact assessment must acknowledge the uncertainty.     

General features of the arctic relevant to climate change in freshwater ecosystems

Large variations exist in the size, abundance and biota of the two principal categories of freshwater ecosystems, lotic (flowing water; e.g., rivers, streams, deltas and estuaries) and lentic (standing water; lakes, ponds and wetlands) found across the circumpolar Arctic. Arctic climate, many components of which exhibit strong variations along latitudinal gradients, directly affects a range of physical, chemical and biological processes in these aquatic systems. Furthermore, arctic climate creates additional indirect ecological effects through the control of terrestrial hydrologic systems and processes, particularly those associated with cryospheric components such as permafrost, freshwater ice and snow accumulation/ablation. The ecological structure and function of arctic freshwater systems are also controlled by external processes and conditions, particularly those in the headwaters of the major arctic rivers and in the adjacent marine environment. The movement of physical, chemical and biotic components through the interlinked lentic and lotic freshwater systems are major determinants of arctic freshwater ecology.     

Modelling Tundra Vegetation Response to Recent Arctic Warming

The Arctic land area has warmed by >1 °C in the last 30 years and there is evidence that this has led to increased productivity and stature of tundra vegetation and reduced albedo, effecting a positive (amplifying) feedback to climate warming. We applied an individual-based dynamic vegetation model over the Arctic forced by observed climate and atmospheric CO₂ for 1980–2006. Averaged over the study area, the model simulated increases in primary production and leaf area index, and an increasing representation of shrubs and trees in vegetation. The main underlying mechanism was a warming-driven increase in growing season length, enhancing the production of shrubs and trees to the detriment of shaded ground-level vegetation. The simulated vegetation changes were estimated to correspond to a 1.75 % decline in snow-season albedo. Implications for modelling future climate impacts on Arctic ecosystems and for the incorporation of biogeophysical feedback mechanisms in Arctic system models are discussed.     

A California air standard to protect vegetation from ozone

Evidence shows that the current national primary ambient air quality standard, if attained, would still permit substantial injury to vegetation. Thus, in March 1987, the California Air Resources Board (CARB) began consideration of the evidence for the effects of ozone (O3) on vegetation, and of several possible state ambient air quality standards designed to protect vegetation, especially crops, from O3 injury. In its review, the CARB addressed a number of issues relevant to such a standard. One issue considered by the CARB is the relationship of an ambient air quality standard to natural background levels of O3, which would greatly influence the practicality of attainment. Attainment of a standard close to natural background could entail excessive costs. Another issue considered is the occurrence of oxidants other than O3 that can damage vegetation. Throughout much of California, O3 accounts for over 90% of the oxidant air pollutants, and the CARB considered whether, in keeping with current practice, O3 should be used as a surrogate for total oxidant air pollutants. A major new piece of information presented to the CARB was an assessment of the economic effects of several potential standards. This assessment, produced by University of California scientists at Riverside and Davis, calculated the benefits of the potential standards in comparison to current O3 levels and estimated natural O3 background. This assessment was developed using field chamber response data, local crop data, and local O3 concentration data as inputs to the California Agricultural Resources Model, which accounts for both supply and demand effects. Because of California's varied climate, agricultural production occurs on a year-round basis, with overlapping growing seasons for many crops. Over long periods of time, O3 levels may vary markedly because of the influence of various factors, and a 1-h standard may not be an accurate indicator of growing season O3 exposure. A moving three-month averaging time has been proposed as a way to approximate the growing seasons of California's 200 crops. However, a sufficiently stringent 1-h standard would serve as a surrogate for a growing season standard. The CARB reviewed evidence supporting both long-term and short-term standards. Agriculture dominates the economies of some regions within California but is a minor components of other regional economies. Because the San Joaquin Valley is California's most important agricultural area, the CARB reviewed evidence for a regional standard for this area that would be more stringent than standards for other parts of the state.     

A new flux-orientated concept to derive critical levels for ozone to protect vegetation

The current European critical levels for ozone (O3) to protect crops, natural and semi-natural vegetation and forest trees are based on a relative small number of open-top chamber experiments with a very limited number of plant species. Therefore, the working group "Effects of Ozone on Plants" of the Commission on Air Pollution Prevention of the Association of German Engineers and the German Institute of Standardization reanalysed the literature on O3 effects on European plant species published between 1989 and 1999. An exposure-response relationship for wild plant species and agricultural crops could be derived from 30 experiments with more than 30 species and 90 data points; the relationship for conifer and deciduous trees is based on 20 experiments with nine species and 50 data points. From these relationships maximum O3 concentrations for different risk stages are deduced, below which the vegetation type is protected on the basis of the respective criteria. Because it is assumed that the fumigation concentrations reflect the O3 concentrations at the top of the canopy, i.e. the upper surface boundary of the quasi-laminar layer if the micrometeorological big-leaf approach is applied, the application of these maximum O3 concentrations requires the transformation of O3 concentrations measured at a reference height above the canopy to the effective phytotoxic concentrations at the top of the canopy. Thus, the approach described in this paper is a synthesis of the classical concept of toxicology of air pollutants (critical concentrations) and the more toxicological relevant dose concept.     

Modelling of indoor exposure to nitrogen dioxide in the UK

A dynamic multi-compartment computer model has been developed to describe the physical processes determining indoor pollutant concentrations as a function of outdoor concentrations, indoor emission rates and building characteristics. The model has been parameterised for typical UK homes and workplaces and linked to a time-activity model to calculate exposures for a representative homemaker, schoolchild and office worker, with respect to NO₂. The estimates of population exposures, for selected urban and rural sites, are expressed in terms of annual means and frequency of hours in which air quality standards are exceeded. The annual mean exposures are estimated to fall within the range of 5–21 ppb for homes with no source, and 21–27 ppb for homes with gas cooking, varying across sites and population groups. The contribution of outdoor exposure to annual mean NO₂ exposure varied from 5 to 24%, that of indoor penetration of outdoor air from 17 to 86% and that of gas cooking from 0 to 78%. The frequency of exposure to 1 h mean concentrations above 150 ppb was very low, except for people cooking with gas.     

Modelling annual pasture dynamics: Application to stomatal ozone deposition

Modelling ozone (O₃) deposition for impact risk assessment is still poorly developed for herbaceous vegetation, particularly for Mediterranean annual pastures. High inter-annual climatic variability in the Mediterranean area makes it difficult to develop models characterizing gas exchange behaviour and air pollutant absorption suitable for risk assessment. This paper presents a new model to estimate stomatal conductance (g_s) of Trifolium subterraneum, a characteristic species of dehesa pastures. The MEDPAS (MEDiterranean PAStures) model couples 3 modules estimating soil water content (SWC), vegetation growth and g_s. The g_s module is a reparameterized version of the stomatal component of the EMEP DO₃SE O₃ deposition model. The MEDPAS model was applied to two contrasting years representing typical dry and humid springs respectively and with different O₃ exposures. The MEDPAS model reproduced realistically the g_s seasonal and inter-annual variations observed in the field. SWC was identified as the major driver of differences across years. Despite the higher O₃ exposure in the dry year, meteorological conditions favoured 2.1 times higher g_s and 56 day longer growing season in the humid year compared to the dry year. This resulted in higher ozone fluxes absorbed by T. subterraneum in the humid year. High inter-family variability was found in gas exchange rates, therefore limiting the relevance of single species O₃ deposition flux modelling for dehesa pastures. Stomatal conductance dynamics at the canopy level need to be considered for more accurate O₃ flux modelling for present and future climate scenarios in the Mediterranean area.     

Why geese benefit from the transition from natural vegetation to agriculture

The energy and nutrient content of most agricultural crops are as good as or superior to natural foods for wild geese and they tend to be available in agricultural landscapes in far greater abundance. Artificial grasslands (fertilised native swards and intensively managed reseeds) offer far superior quality forage and higher intake rates than seminatural or natural grasslands. The availability of such abundant artificial food explains the abandonment of traditional habitats for farmland by geese over the last 50–100 years and favours no reduction in current levels of exploitation of agriculture. Continental scale spatial and temporal shifts among geese undergoing spring fattening confirm their flexibility to respond rapidly to broadscale changes in agriculture. These dramatic changes support the hypothesis that use of agricultural landscapes has contributed to elevated reproductive success and that European and North American farmland currently provides unrestricted winter carrying capacity for goose populations formerly limited by wetlands habitats prior to the agrarian revolution of the last century.     

Meta-analysis of the relative sensitivity of semi-natural vegetation species to ozone

This study identified 83 species from existing publications suitable for inclusion in a database of sensitivity of species to ozone (OZOVEG database). An index, the relative sensitivity to ozone, was calculated for each species based on changes in biomass in order to test for species traits associated with ozone sensitivity. Meta-analysis of the ozone sensitivity data showed a wide inter-specific range in response to ozone. Some relationships in comparison to plant physiological and ecological characteristics were identified. Plants of the therophyte lifeform were particularly sensitive to ozone. Species with higher mature leaf N concentration were more sensitive to ozone than those with lower leaf N concentration. Some relationships between relative sensitivity to ozone and Ellenberg habitat requirements were also identified. In contrast, no relationships between relative sensitivity to ozone and mature leaf P concentration, Grime's CSR strategy, leaf longevity, flowering season, stomatal density and maximum altitude were found. The relative sensitivity of species and relationships with plant characteristics identified in this study could be used to predict sensitivity to ozone of untested species and communities.     

Temporal processes that contribute to nonlinearity in vegetation responses to ozone exposure and dose

Ozone interacts with plant tissue through distinct temporal processes. Sequentially, plants are exposed to ambient O₃ that (1) moves through the leaf boundary layer, (2) is taken up into plant tissue primarily through stomata, and (3) undergoes chemical interaction within plant tissue, first by initiating alterations and then as part of plant detoxification and repair. In this paper, we discuss the linkage of the temporal variability of apoplastic ascorbate with the diurnal variability of defense mechanisms in plants and compare this variability with daily maximum O₃ concentration and diurnal uptake and entry of O₃ into the plant through stomata. We describe the quantitative evidence on temporal variability in concentration and uptake and find that the time incidence for maximum defense does not necessarily match diurnal patterns for maximum O₃ concentration or maximum uptake. We suggest that the observed out-of-phase association of the diurnal patterns for the above three processes produces a nonlinear relationship that results in a greater response from the higher hourly average O₃ concentrations than from the lower or mid-level values. The fact that these out-of-phase processes affect the relationship between O₃ exposure/dose and vegetation effects ultimately impact the ability of flux-based indices to predict vegetation effects accurately for purposes of standard setting and critical levels. Based on the quantitative aspect of temporal variability identified in this paper, we suggest that the inclusion of a diurnal pattern for detoxification in effective flux-based models would improve the predictive characteristics of the models. While much of the current information has been obtained using high O₃ exposures, future research results derived from laboratory biochemical experiments that use short but elevated O₃ exposures should be combined with experimental results that use ambient-type exposures over longer periods of time. It is anticipated that improved understanding will come from future research focused on diurnal variability in plant defense mechanisms and their relationship to the diurnal variability in ambient O₃ concentration and stomatal conductance. This should result in more reliable O₃ exposure standards and critical levels.     

Dry deposition of ammonia,nitric acid,ammonium, and nitrate to alpine tundra at Niwot Ridge,Colorado

Micrometeorological measurements and ambient air samples, analyzed for concentrations of NH₃, HNO₃, NH₄⁺, and NO₃⁻, were collected at an alpine tundra site on Niwot Ridge, Colorado. The measured concentrations were extremely low and ranged between 5 and 70 ng N m⁻³. Dry deposition fluxes of these atmospheric species were calculated using the micrometeorological gradient method. The calculated mean flux for NH₃ indicates a net deposition to the surface and indicates that NH₃ contributed significantly to the total N deposition to the tundra during the August–September measurement period. Our pre-measurement estimate of the compensation point for NH₃ in air above the tundra was 100–200 ng N m⁻³; thus, a net emission of NH₃ was expected given the low ambient concentrations of NH₃ observed. Based on our results, however, the NH₃ compensation point at this alpine tundra site appears to have been at or below about 20 ng N m⁻³. Large deposition velocities (>2 cm s⁻¹) were determined for nitrate and ammonium and may result from reactions with surface-derived aerosols.     

Modelling stomatal ozone flux and deposition to grassland communities across Europe

Regional scale modelling of both ozone deposition and the risk of ozone impacts is poorly developed for grassland communities. This paper presents new predictions of stomatal ozone flux to grasslands at five different locations in Europe, using a mechanistic model of canopy development for productive grasslands to generate time series of leaf area index and soil water potential as inputs to the stomatal component of the DO(3)SE ozone deposition model. The parameterisation of both models was based on Lolium perenne, a dominant species of productive pasture in Europe. The modelled seasonal time course of stomatal ozone flux to both the whole canopy and to upper leaves showed large differences between climatic zones, which depended on the timing of the start of the growing season, the effect of soil water potential, and the frequency of hay cuts. Values of modelled accumulated flux indices and the AOT40 index showed a five-fold difference between locations, but the locations with the highest flux differed depending on the index used; the period contributing to the accumulation of AOT40 did not always coincide with the modelled period of active ozone canopy uptake. Use of a fixed seasonal profile of leaf area index in the flux model produced very different estimates of annual accumulated total canopy and leaf ozone flux when compared with the flux model linked to a simulation of canopy growth. Regional scale model estimates of both the risks of ozone impacts and of total ozone deposition will be inaccurate unless the effects of climate and management in modifying grass canopy growth are incorporated.