James D. M. Speed

Local temperatures inferred from plant communities suggest strong spatial buffering of climate warming across Northern Europe

Recent studies from mountainous areas of small spatial extent (<2500 km(2) ) suggest that fine-grained thermal variability over tens or hundreds of metres exceeds much of the climate warming expected for the coming decades. Such variability in temperature provides buffering to mitigate climate-change impacts. Is this local spatial buffering restricted to topographically complex terrains? To answer this, we here study fine-grained thermal variability across a 2500-km wide latitudinal gradient in Northern Europe encompassing a large array of topographic complexities. We first combined plant community data, Ellenberg temperature indicator values, locally measured temperatures (LmT) and globally interpolated temperatures (GiT) in a modelling framework to infer biologically relevant temperature conditions from plant assemblages within <1000-m(2) units (community-inferred temperatures: CiT). We then assessed: (1) CiT range (thermal variability) within 1-km(2) units; (2) the relationship between CiT range and topographically and geographically derived predictors at 1-km resolution; and (3) whether spatial turnover in CiT is greater than spatial turnover in GiT within 100-km(2) units. Ellenberg temperature indicator values in combination with plant assemblages explained 46-72% of variation in LmT and 92-96% of variation in GiT during the growing season (June, July, August). Growing-season CiT range within 1-km(2) units peaked at 60-65°N and increased with terrain roughness, averaging 1.97 °C (SD = 0.84 °C) and 2.68 °C (SD = 1.26 °C) within the flattest and roughest units respectively. Complex interactions between topography-related variables and latitude explained 35% of variation in growing-season CiT range when accounting for sampling effort and residual spatial autocorrelation. Spatial turnover in growing-season CiT within 100-km(2) units was, on average, 1.8 times greater (0.32 °C km(-1) ) than spatial turnover in growing-season GiT (0.18 °C km(-1) ). We conclude that thermal variability within 1-km(2) units strongly increases local spatial buffering of future climate warming across Northern Europe, even in the flattest terrains.

Experimental evidence for herbivore limitation of the treeline

The treeline ecotone divides forest from open alpine or arctic vegetation states. Treelines are generally perceived to be temperature limited. The role of herbivores in limiting the treeline is more controversial, as experimental evidence from relevant large scales is lacking. Here we quantify the impact of different experimentally controlled herbivore densities on the recruitment and survival of birch Betula pubescens tortuosa along an altitudinal gradient in the mountains of southern Norway. After eight years of summer grazing in large-scale enclosures at densities of 0, 25, and 80 sheep/km2, birch recruited within the whole altitudinal range of ungrazed enclosures, but recruitment was rarer in enclosures with low-density sheep and was largely limited to within the treeline in enclosures with high-density sheep. In contrast, the distribution of saplings (birch older than the experiment) did not differ between grazing treatments, suggesting that grazing sheep primarily limit the establishment of new tree recruits rather than decrease the survival of existing individuals. This study provides direct experimental evidence that herbivores can limit the treeline below its potential at the landscape scale and even at low herbivore densities in this climatic zone. Land use changes should thus be considered in addition to climatic changes as potential drivers of ecotone shifts.

What are the impacts of reindeer/caribou ( Rangifer tarandus L.) on arctic and alpine vegetation? A systematic review protocol

BackgroundThe reindeer (or caribou, Rangifer tarandus L.) has a natural range extending over much of Eurasia’s and North America’s arctic, alpine and boreal zones, yet its impact on vegetation is still unclear. This lack of a common understanding hampers both the management of wild and semi-domesticated reindeer populations and the preservation of biodiversity. To achieve a common platform, we have undertaken a systematic review of published studies that compare vegetation at sites with different reindeer densities. Besides biodiversity, we focused on effects on major plant growth forms.MethodsSearches for literature were made using online publication databases, search engines, specialist websites and bibliographies of literature reviews. Search terms were developed in English, Finnish, Norwegian, Russian and Swedish. Identified articles were screened for relevance based on titles, abstracts and full text using inclusion criteria set out in an a priori protocol. Relevant articles were then subject to critical appraisal of susceptibility to bias. Data on outcomes such as abundance, biomass, cover and species richness of vegetation were extracted together with metadata on site properties and other potential effect modifiers.ResultsOur searches identified more than 6,000 articles. After screening for relevance, 100 of them remained. Critical appraisal excluded 60 articles, leaving 40 articles with 41 independent studies. Almost two thirds of these studies had been conducted in Fennoscandia. Meta-analysis could be made of data from 31 of the studies. Overall, effects of reindeer on species richness of vascular plants depended on temperature, ranging from negative at low temperature to positive at high temperature. Effects on forbs, graminoids, woody species, and bryophytes were weak or non-significant, whereas the effect on lichens was negative. However, many individual studies showed clear positive or negative effects, but the available information was insufficient to explain this context dependence.ConclusionsWe see two pressing matters emerging from our study. First, there is a lack of research with which to build a circumpolar understanding of grazing effects, which calls for more studies using a common protocol to quantify reindeer impacts. Secondly, the highly context-dependent outcomes suggest that research and management have to consider local conditions. For instance, predictions of what a management decision would mean for the effects of reindeer on vegetation will have to take the variation of vegetation types and dominant growth forms, productivity, and grazing history into account. Policy and management have to go hand-in-hand with research in individual cases if the dynamics between plants, animals, and humans are to be sufficiently understood.

The Response of Alpine Salix Shrubs to Long-Term Browsing Varies with Elevation and Herbivore Density

Abstract The widespread expansion of shrubs into arctic and alpine regions has frequently been linked to climatic warming, but herbivory can play a role in addition to, or in interaction with, climate. Willow (Salix spp.) shrubs are important constituents of alpine ecosystems, influencing community structure and providing habitat and forage for many species. We investigate the impact of browsing by domestic sheep (Ovis aries), the dominant herbivore in Norwegian mountains, on Salix stem density, height, and radial growth. We used a field experiment, replicated along an elevational gradient, with manipulated densities of sheep (no sheep, low density, and high density at 0, 25, and 80 sheep km-2). We found that Salix shoot density and radial growth were greatest at high sheep density but only at low elevations, indicating that competition from field-layer vegetation at lower sheep densities reduced Salix performance. At higher elevations Salix shoot density and radial growth were lower at high sheep density than at low sheep density and in the absence of sheep. Thus at high elevations sheep browsing is likely to slow the expansion of Salix shrubs, whilst the removal of browsing is likely to constrain Salix expansion at lower elevations.

Sheep grazing in the North Atlantic region: A long-term perspective on environmental sustainability

Sheep grazing is an important part of agriculture in the North Atlantic region, defined here as the Faroe Islands, Greenland, Iceland, Norway and Scotland. This process has played a key role in shaping the landscape and biodiversity of the region, sometimes with major environmental consequences, and has also been instrumental in the development of its rural economy and culture. In this review, we present results of the first interdisciplinary study taking a long-term perspective on sheep management, resource economy and the ecological impacts of sheep grazing, showing that sustainability boundaries are most likely to be exceeded in fragile environments where financial support is linked to the number of sheep produced. The sustainability of sheep grazing can be enhanced by a management regime that promotes grazing densities appropriate to the site and supported by area-based subsidy systems, thus minimizing environmental degradation, encouraging biodiversity and preserving the integrity of ecosystem processes.

What is the influence of a reduction of planktivorous and benthivorous fish on water quality in temperate eutrophic lakes? A systematic review

BackgroundIn recent decades, many attempts have been made to restore eutrophic lakes through biomanipulation. Reducing the populations of planktivorous and benthivorous fish (either directly or through stocking of piscivorous fish) may induce ecosystem changes that increase water transparency and decrease the risk of algal blooms and fish kills, at least in the short term. However, the generality of biomanipulation effects on water quality across lake types and geographical regions is not known. Therefore, we have undertaken a systematic review of such effects in eutrophic lakes in temperate regions throughout the world.MethodsSearches for literature were made using online publication databases, search engines, specialist websites and bibliographies of literature reviews. Search terms were developed in English, Danish, Dutch and Swedish. Identified articles were screened for relevance using inclusion criteria set out in an a priori protocol. To reduce the risk of bias, we then critically appraised the combined evidence found on each biomanipulation. Data were extracted on outcomes such as Secchi depth and chlorophyll a concentration before, during and/or after manipulation, and on effect modifiers such as lake properties and amounts of fish removed or stocked.ResultsOur searches identified more than 14,500 articles. After screening for relevance, 233 of them remained. After exclusions based on critical appraisal, our evidence base included useful data on 128 biomanipulations in 123 lakes. Of these interventions, 85% had been made in Europe and 15% in North America. Meta-analysis showed that removal of planktivores and benthivores (with or without piscivore stocking) leads to increased Secchi depth and decreased chlorophyll a concentration during intervention and the first three years afterwards. Piscivore stocking alone has no significant effect. The response of chlorophyll a levels to biomanipulation is stronger in lakes where fish removal is intense, and in lakes which are small and/or have high pre-manipulation concentrations of total phosphorus.ConclusionsOur review improves on previous reviews of biomanipulation in that we identified a large number of case studies from many parts of the world and used a consistent, repeatable process to screen them for relevance and susceptibility to bias. Our results indicate that removal of planktivorous and benthivorous fish is a useful means of improving water quality in eutrophic lakes. Biomanipulation tends to be particularly successful in relatively small lakes with short retention times and high phosphorus levels. More thorough fish removal increases the efficacy of biomanipulation. Nonetheless successes and failures have occurred across a wide range of conditions.

Browsing of sallow (Salix caprea L.) and rowan (Sorbus aucuparia L.) in the context of life history strategies: a literature review

Sallow (Salix caprea L.) and rowan (Sorbus aucuparia L.) constitute small proportions of the deciduous tree volume in Scandinavia, but are highly preferred winter forage for moose and red deer, which occur at historically high densities. Thus, a possible decline of these tree species has been indicated. Against this background, we have reviewed the life histories of relevance for browsing, as well as the basic biology and genetics of sallow and rowan. The species show similarities with respect to short lifespan, small size and sympodial growth pattern, which are risk factors in a browsing context. They also have high juvenile growth rate, important for growing quickly out of reach of browsers. Sallow depends strongly on disturbance for establishment and is more demanding with respect to soil and light conditions than rowan, possibly important for the substantially lower abundance of sallow on the Norwegian Forest Inventory plots. Similarly, the relative recruitment of small size classes of sallow is less than for rowan. Although recruitment is reported to be hampered in wintering areas with high moose or red deer densities, the inventory data, however, dating only back to 1994, do not suggest a general decrease in any of the species. Sallow and rowan saplings show low mortality in moose and deer dominated areas and the species can be characterised as rather resilient to browsing. Of more concern is that browsing can constrain the development of mature rowan and sallow trees locally, with possible consequences for associated epiphytic biodiversity.

Experimental Effects of Herbivore Density on Aboveground Plant Biomass in an Alpine Grassland Ecosystem

Abstract Herbivores may increase or decrease aboveground plant productivity depending on factors such as herbivore density and habitat productivity. The grazing optimization hypothesis predicts a peak in plant production at intermediate herbivore densities, but has rarely been tested experimentally in an alpine field setting. In an experimental design with three densities of sheep (high, low, and no sheep), we harvested aboveground plant biomass in alpine grasslands prior to treatment and after five years of grazing. Biomass of vascular plants decreased at high sheep density, and marginally increased at low sheep density. The ungrazed treatment was found to be intermediate. Companion studies conducted at the same site suggest, (1) that changes in soil N-mineralization and plant community patterns are contributing to the herbivore-induced effects on plant productivity in alpine grasslands, (2) that herbivore-driven changes in plant productivity feed into the future performance for the herbivore as the marginal increase in productivity at low density corresponds with a temporal increase in lamb growth. Our study provides experimental evidence for a nonlinear effect of increased grazing on plant productivity as predicted by the grazing optimization hypothesis. This has important repercussions for ecosystem function and management, as it demonstrates how herbivore density can either increase or decrease ecosystem productivity over time.

Natural and cultural heritage in mountain landscapes: towards an integrated valuation

Mountain areas of Europe have been managed by humans for a long time, leading to a prevalence of semi-natural habitats in mountain landscapes today. These landscapes contain both natural and cultural heritage; however, natural and cultural heritage are rarely considered together when valuing landscapes and developing management plans in protected areas. Here we present a case study of seven protected areas in the mountains of Great Britain and Norway. We take a long-term perspective on landscape and land-use change and propose an integrated model of landscape valuation on the basis of combined natural and cultural heritage. Our model plots indicators of natural and cultural heritage along a gradient of land-use intensity, allowing simultaneous assessment and highlighting how valuation depends on what type of heritage is considered. We show that while contemporary land-use changes follow similar trajectories in Norway and Britain, different land-use histories mean that the loss of heritage differs between the regions. The model presented here thus allows for the consolidation of valuation based on both cultural and natural heritage in landscapes.

Low Intensities of Red Deer Browsing Constrain Rowan Growth in Mature Boreal Forests of Western Norway

Abstract: Browsing by cervids plays a key role in structuring forest ecosystems and dynamics. Many boreal forest systems are managed for timber resources, and at the same time the wild cervid populations are also harvested. Thus, the determination of sustainable densities of cervids for the purpose of forest and game management is challenging. In this study we report on a red deer (Cervus elaphus) exclosure experiment in the mature forests of Western Norway. Ten pairs of exclosures and browsed plots were initiated in 2008. The rate of browsing and height growth of marked individuals was recorded annually, and the total densities of all tree species assessed over the following 4 y. We found that height growth of rowan (Sorbus aucuparia) saplings (1 m tall), the most numerous tree species at the site, was prevented when 20% of the shoots were browsed. Outside of the exclosures, net height growth of rowan saplings tended to be positive when trees were below 40 cm in height, but growth was constrained in rowan saplings over this height. The density of rowan also increased in both treatments, showing that recruitment was occurring, but the increase was greater where browsed than in the exclosure. The increase in density of rowan, combined with the curtailment of height growth in the presence of red deer, serves to create a carpet of short stature rowan saplings. This has parallels with the browsing lawn concept, but it seems to occur in interaction with snow depth; individuals protruding above the snow layer are likely to be browsed during the winter, whilst smaller individuals are protected during this season, when browsing is at its peak.