Holocene climate,fire and vegetation dynamics at the treeline in the Northwestern Swiss Alps

Treelines are expected to rise to higher elevations with climate warming; the rate and extent however are still largely unknown. Here we present the first multi-proxy palaeoecological study from the treeline in the Northwestern Swiss Alps that covers the entire Holocene. We reconstructed climate, fire and vegetation dynamics at Iffigsee, an alpine lake at 2,065 m a.s.l., by using seismic sedimentary surveys, loss on ignition, visible spectrum reflectance spectroscopy, pollen, spore, macrofossil and charcoal analyses. Afforestation with Larix decidua and tree Betula (probably B. pendula) started at ~9,800 cal. b.p., more than 1,000 years later than at similar elevations in the Central and Southern Alps, indicating cooler temperatures and/or a high seasonality. Highest biomass production and forest position of ~2,100–2,300 m a.s.l. are inferred during the Holocene Thermal Maximum from 7,000 to 5,000 cal. b.p. With the onset of pastoralism and transhumance at 6,800–6,500 cal. b.p., human impact became an important factor in the vegetation dynamics at Iffigsee. This early evidence of pastoralism is documented by the presence of grazing indicators (pollen, spores), as well as a wealth of archaeological finds at the nearby mountain pass of Schnidejoch. Human and fire impact during the Neolithic and Bronze Ages led to the establishment of pastures and facilitated the expansion of Picea abies and Alnus viridis. We expect that in mountain areas with land abandonment, the treeline will react quickly to future climate warming by shifting to higher elevations, causing drastic changes in species distribution and composition as well as severe biodiversity losses.     

Debris-flow activity along a torrent in the Swiss Alps: Minimum frequency of events and implications for forest dynamics

This study reports on a tree-ring-based reconstruction of geomorphic activity and illustrates impacts of such processes on tree germination along a debris-flow torrent in the Swiss Alps. Analysis included the identification of growth disturbances and the assessment of germination dates for 28 trees along the channel of the Geisstriftbach torrent (Valais, Swiss Alps). Provided that recolonizing trees indicate the minimum time elapsed since the last deposition, germination dates suggest that a devastating debris-flow event in the 1880s had cleared the surface and scoured the currently active channel. This interpretation is supported by two topographic maps showing a dislocation of the channel. Analyzing the age structure of trees along the channel in more detail, we observe higher tree ages with increasing distance from the cone apex. In addition, dendrogeomorphic methods allowed for the reconstruction of 13 debris-flow events between AD 1913 and 2006. In combination with geomorphic mapping, the spatial distribution of trees affected by individual events was assessed and a minimum frequency of previous debris-flow events reconstructed. Although the present study was based on a limited set of tree-ring records, it illustrates that tree-ring analysis in combination with cartographic methods holds much promise for dating minimum ages of surfaces cleared by destructive events as well as for determining the spatio-temporal impacts of past debris-flow activity.     

Inbreeding depression and male fitness in black grouse

The male lifetime lekking performance was studied, and related to inbreeding-outbreeding in a wild population of black grouse (Tetrao tetrix) in central Finland between 1989 and 1995. Inbreeding was measured as the mean heterozygosity and mean d(2) of 15 microsatellite loci. We found a significantly positive relationship between mean d(2) and lifetime copulation success (LCS), while the relationship between heterozygosity and LCS was close to significant. We also found that males that never obtained a lek territory had significantly lower mean heterozygosity than males that were observed on a territory at least during one mating season in their life. Furthermore, among males that were successful in obtaining a lek territory, LCS and mean d(2) were highest for those males that held central territories. We suggest that inbred males have a disadvantage (or outbred males have an advantage) in the competition for territories that may explain the relationships with LCS and inbreeding. Furthermore, the fact that mean d(2) was positively correlated with LCS whereas heterozygosity was not when we restricted the analysis to territorial males, suggests that mean d(2) provides more information about levels of inbreeding-outbreeding than heterozygosity alone, and potentially highlights the effects of heterosis. To our knowledge, this is the first time that measures of inbreeding and lifetime fitness have been linked in a non-isolated population. This is important in establishing that the relationships found in previous studies are not artefacts of low gene flow created by limited dispersal but a general feature of wild vertebrate populations.     

Altitudinal distribution of alien plant species in the Swiss Alps

In summer 2003 we recorded the presence and abundance of alien plant species at 232 sites (107 railway stations and 125 road sites) along mountain passes in the Swiss Alps. The altitudinal distribution of species was related to the current abundance of the species in Switzerland and time since introduction. A total of 155 alien taxa were recorded. Numbers of species per site declined exponentially with altitude, and only a few species were found in the alpine zone (>2000 m). In contrast, species richness among comparable native taxa appeared to be nearly independent of altitude over the range investigated. Maximum altitude reached by alien species was related positively to both total area occupied in Switzerland and to time since introduction. A comparison of the results with earlier records suggests that many species, particularly those previously restricted to low or intermediate altitudes, have advanced their altitudinal limits over the past few decades. Various hypotheses are presented to explain the declining abundance of alien species with altitude: low-altitude filter effects, low propagule pressure, and genetic swamping of peripheral populations at higher altitudes. However, at present we do not have sufficient evidence to determine the relative importance of these effects. We conclude that invasion into mountain areas such as the Swiss Alps tends to proceed rather slowly, though the process may be accelerated by climatic warming. For this reason, further research to investigate the processes determining how plants invade mountain areas is urgently needed. And more generally, investigations into the distribution of alien species along strong altitudinal gradients may provide valuable insights into the mechanisms driving the spread of alien organisms.     

Spatial synchrony in red grouse population dynamics

Red grouse Lagopus lagopus scoticus populations exhibit unstable dynamics that are often characterised by regular periodic fluctuations in abundance. Time-series' of grouse harvesting records collected from 287 management units (moors) across Scotland, England and Wales were analysed to investigate the broad scale patterns of synchrony in these fluctuations. Estimation of the spatial autocorrelation of grouse population dynamics across moors indicates relatively high levels of synchrony between populations on adjacent moors, but that this synchrony declines sharply with increasing inter-moor distance. At distances of greater than 100  km, grouse population time-series exhibit only weakly positive cross-correlation coefficients. Twenty-eight geographical, environmental and other candidate variables were examined to construct a general linear model to explain variation in local synchrony. Grouse moor productivity (average size of shooting bag), distance from the Atlantic coast moving in a north-easterly direction, April and June temperatures, and June rainfall significantly increased the explanatory power of this model. An understanding of the processes underlying synchrony in red grouse population dynamics is a prerequisite to anticipating the effects of large-scale environmental change on regional patterns of grouse distribution and abundance.     

Prevalence of acute mountain sickness in the Swiss Alps.

OBJECTIVE--To assess the prevalence of symptoms and signs of acute mountain sickness of the Swiss Alps. DESIGN--A study using an interview and clinical examination in a representative population of mountaineers. Positive symptoms and signs were assigned scores to quantify the severity of acute mountain sickness. SETTING--Four huts in the Swiss Alps at 2850 m, 3050 m, 3650 m, and 4559 m. SUBJECTS--466 Climbers, mostly recreational: 47 at 2850 m, 128 at 3050 m, 82 at 3650, and 209 at 4559 m. RESULTS--In all, 117 of the subjects were entirely free of symptoms and clinical signs of acute mountain sickness; 191 had one or two symptoms and signs; and 158 had more than two. Those with more than two symptoms and signs were defined as suffering from acute mountain sickness. At 4559 m 11 climbers presented with high altitude pulmonary oedema or cerebral oedema, or both. Men and women were equally affected. The prevalence of acute mountain sickness correlated with altitude: it was 9% at 2850 m, 13% at 3050 m, 34% at 3650 m, and 53% at 4559 m. The most frequent symptoms and signs were insomnia, headache, peripheral oedema, and scanty pulmonary rales. Severe headache, vomiting, dizziness, tachypnoea, and pronounced pulmonary rales were associated with other symptoms and signs and therefore characteristic of acute mountain sickness. CONCLUSION--Acute mountain sickness is not an uncommon disease at moderately high altitude--that is, above 2800 m. Severe headache, vomiting, dizziness, tachypnoea, and pronounced pulmonary rales indicate severe acute mountain sickness, and subjects who suffer these should immediately descend to lower altitudes.     

Diversity change of mountain hay meadows in the Swiss Alps

We investigated changes in plant diversity in the traditionally fertilised and species-rich Golden Oat meadows by repeating historic grassland relevés from the 1940s in two agricultural areas in the Swiss Alps. The effects of time on alpha and beta diversity at plot level were analysed within the study areas. The specificity of the two study areas was investigated by comparing the variability of species composition within and between study areas.In both regions, species richness at plot level (alpha diversity) was found to be lower in the recent relevés than in the historic ones. However, the variability of species composition within the study areas (beta diversity) was higher between the recent relevés than between the historic relevés. There was a significant floristic differentiation between the two study areas sixty years ago but not today.Plant diversity in Golden Oat meadows has changed significantly during the last sixty years. The decrease in species richness at plot level was anticipated, but not the increase in beta diversity between plots, which may indicate individualised management regimes of modern farmers. The loss of specificity between the study areas revealed an important and as yet hardly considered further dimension of biodiversity change, which leads to a homogenisation of plant diversity at a regional level.     

Soil protist function varies with elevation in the Swiss Alps

Protists are abundant and play key trophic functions in soil. Documenting how their trophic contributions vary across large environmental gradients is essential to understand and predict how biogeochemical cycles will be impacted by global changes. Here, using amplicon sequencing of environmental DNA in open habitat soil from 161 locations spanning 2600 m of elevation in the Swiss Alps (from 400 to 3000 m), we found that, over the whole study area, soils are dominated by consumers, followed by parasites and phototrophs. In contrast, the proportion of these groups in local communities shows large variations in relation to elevation. While there is, on average, three times more consumers than parasites at low elevation (400–1000 m), this ratio increases to 12 at high elevation (2000–3000 m). This suggests that the decrease in protist host biomass and diversity toward mountains tops impact protist functional composition. Furthermore, the taxonomic composition of protists that infect animals was related to elevation while that of protists that infect plants or of protist consumers was related to soil pH. This study provides a first step to document and understand how soil protist functions vary along the elevational gradient.     

Uncertainty in predicting range dynamics of endemic alpine plants under climate warming

Correlative species distribution models have long been the predominant approach to predict species’ range responses to climate change. Recently, the use of dynamic models is increasingly advocated for because these models better represent the main processes involved in range shifts and also simulate transient dynamics. A well‐known problem with the application of these models is the lack of data for estimating necessary parameters of demographic and dispersal processes. However, what has been hardly considered so far is the fact that simulating transient dynamics potentially implies additional uncertainty arising from our ignorance of short‐term climate variability in future climatic trends. Here, we use endemic mountain plants of Austria as a case study to assess how the integration of decadal variability in future climate affects outcomes of dynamic range models as compared to projected long‐term trends and uncertainty in demographic and dispersal parameters. We do so by contrasting simulations of a so‐called hybrid model run under fluctuating climatic conditions with those based on a linear interpolation of climatic conditions between current values and those predicted for the end of the 21st century. We find that accounting for short‐term climate variability modifies model results nearly as differences in projected long‐term trends and much more than uncertainty in demographic/dispersal parameters. In particular, range loss and extinction rates are much higher when simulations are run under fluctuating conditions. These results highlight the importance of considering the appropriate temporal resolution when parameterizing and applying range‐dynamic models, and hybrid models in particular. In case of our endemic mountain plants, we hypothesize that smoothed linear time series deliver more reliable results because these long‐lived species are primarily responsive to long‐term climate averages.     

Flammability dynamics in the Australian Alps

Forests of the Australian Alps (SE Australia) are considered some of the most vulnerable to climate change in the country, with ecosystem collapse considered likely for some due to frequent fire. It is not yet known, however, whether increasing fire frequency may stabilize due to reductions in flammability related to reduced time for fuel accumulation, show no trend, or increase due to positive feedbacks related to vegetation changes. To determine what these trends have been historically, dynamics were measured for 58 years of mapped fire history. The 1.4 million ha forested area was divided into broad formations based on structure and dominant canopy trees, and dynamics were measured for each using flammability ratio, a modification of probability of ignition at a point. Crown fire likelihood was measured for each formation, based on satellite‐derived measurements of the 2003 fire effects across a large part of the area. Contrary to popular perception but consistent with mechanistic expectations, all forests exhibited pronounced positive feedbacks. The strongest response was observed in tall, wet forests dominated by Ash‐type eucalypts, where, despite a short period of low flammability following fire, post‐disturbance stands have been more than eight times as likely to burn than have mature stands. The weakest feedbacks occurred in open forest, although post‐disturbance forests were still 1.5 times as likely to burn as mature forests. Apart from low, dry open woodland where there was insufficient data to detect a trend, all forests were most likely to experience crown fire during their period of regeneration. The implications of this are significant for the Alps, as increasing fire frequency has the potential to accelerate by producing an increasingly flammable landscape. These effects may be semi‐permanent in tall, wet forest, where frequent fire promotes ecosystem collapse into either the more flammable open forest formation, or to heathland.     

The relative importance of climatic effects, wildfires and management for future forest landscape dynamics in the Swiss Alps

Forest landscape dynamics result from the complex interaction of driving forces and ecological processes operating on various scales. Projected climate change for the 21st century will alter climate‐sensitive processes, causing shifts in species composition and also bringing about changes in disturbance regimes, particularly regarding wildfires. Previous studies of the impact of climate change on forests have focused mainly on the direct effects of climate. In the present study, we assessed the interactions among forest dynamics, climate change and large‐scale disturbances such as fire, wind and forest management. We used the Land Clim model to investigate the influence, interactions and the relative importance of these different drivers of landscape dynamics in two case study areas of the European Alps. The simulations revealed that projected future climate change would cause extensive forest cover changes, beginning in the coming decades. Fire is likely to become almost as important for shaping the landscape as the direct effects of climate change, even in areas where major wildfires do not occur under current climatic conditions. The effects of variable wind disturbances and harvesting regimes, however, are less likely to have a considerable impact on forest development compared with the direct effects of climate change coupled with the indirect effects of increased fire activity. We conclude that the joint direct and indirect effects of climate change are likely to have major consequences for mountain forests in the European Alps, including their ability to provide protection against natural hazards.     

Small-scale habitat use of black grouse (Tetrao tetrix L.) and rock ptarmigan (Lagopus muta helvetica Thienemann) in the Austrian Alps

We investigated the small-scale habitat use of two grouse species, black grouse (Tetrao tetrix L.) and rock ptarmigan (Lagopus muta helvetica Thienemann) in a study area in the Austrian Central Alps in summer. To build habitat suitability models, we applied multiple logistic regression using presence–absence data from fieldwork as the response variable and a set of habitat characteristics as explanatory variables, respectively. To gain a better understanding of the mechanisms that drive habitat selection, we tested for two-way interaction terms before excluding any variables from the initial variable set. Four explanatory variables significantly contributed to the black grouse model: dwarf shrub cover, dwarf shrub height, patchiness and ant hills. The final model for rock ptarmigan contained three explanatory variables: dwarf shrub cover, rock cover and dwarf shrub height. Most notably, the interaction terms dwarf shrub cover × patchiness in the black grouse model and dwarf shrub cover × dwarf shrub height, rock cover × dwarf shrub height in the rock ptarmigan model point out trade-off mechanisms between food, cover and overview providing features. Thus, our models do not only identify the parameters that mainly drive habitat selection, but also deepen our understanding about the causal relationships between these factors. Therefore, the information gained in this study allows for a deduction of appropriate habitat management strategies and supports conservation efforts of local stakeholders.     

Modelling range dynamics of terricolous lichens of the genus Peltigera in the Alps under a climate change scenario

Climate change is expected to strongly impact biodiversity in Alpine ecosystems and species distribution modelling is increasingly used to provide anticipatory information to guide conservation. In this study, (1) we quantified the range loss, range gain, range change and range turnover caused by climate change in the genus Peltigera a group of terricolous lichens widespread across the Alps, and then (2) we evaluated the relationships between the predictors of range dynamics and functional traits. Our results indicate moderate range dynamics for species of the genus Peltigera across the Alps under a climate change scenario. This would imply a relative stability and resistance of these lichens to climate change that may reflect the local persistence of the species under sub-optimal conditions. Our results also suggest that range dynamics could be associated with functional traits mainly related to water-use strategies and to a trade-off between dispersal and establishment ability. This finding suggests that functional traits may strongly modulate the lichen response to climate change and that species with similar functional traits are prone to similar selective pressures.     

Conservation and restoration of Nardus grasslands in the Swiss northern Alps

Aim: Species‐rich Nardus grasslands are high nature‐value habitats. In Switzerland, many of these grasslands are degraded even though they have been under protection since the 1980s. Degradation shows two divergent trends: Nardus grasslands are either dominated by Nardus stricta or by eutrophic plants, both trends leading to the disappearance of typical Nardus grassland species. With this study, we aim to identify the factors that could be adjusted to conserve the integrity of this habitat. Location: Bernese Alps, Switzerland. Methods: In 2016, we investigated the underlying causes of this degradation process by assessing vegetation composition in 48 Nardus grasslands located in the Swiss northern Alps of canton Bern and linking it to soil, management and environmental variables. To explore the effect of the degradation on higher trophic levels, orthopteran species richness and densities were assessed. Results: Results show that Nardus meadows (mown) are rarely degraded compared to Nardus pastures (grazed). Within pastures, eutrophic plants are most abundant on small pastures with low soil carbon/nitrogen ratio, indicating high nutrient availability. Nardus stricta dominance is most problematic on north‐exposed slopes and in summer pastures. A plausible driver of both degradation trends is the grazing management regime: within small pastures at low elevation where the grazing periods are short but intense, soil carbon/nitrogen ratio is low because of high dung deposition, thus the eutrophic species become dominant. Contrastingly, on large summer pastures with low‐intensity and long‐term grazing, N. stricta becomes dominant due to selective grazing. Both degradation trends show a negative impact on the orthopteran density. Conclusion: Species‐rich Nardus grasslands are a precious alpine habitat for specialised plant species and orthopterans. With an extensive mowing regime or a more controlled grazing regime that homogenises intensity in time and space, species‐rich Nardus grasslands can be conserved in Switzerland.     

Contemporary carbon stocks of mineral forest soils in the Swiss Alps

Soil organic carbon (SOC) has been identified as the main globalterrestrial carbon reservoir, but considerable uncertainty remains as toregional SOC variability and the distribution of C between vegetationand soil. We used gridded forest soil data (8–km × 8–km)representative of Swiss forests in terms of climate and forest typedistribution to analyse spatial patterns of mineral SOC stocks alonggradients in the European Alps for the year 1993. At stand level, meanSOC stocks of 98 t C ha^–1 (N = 168,coefficient of variation: 70%) were obtained for the entiremineral soil profile, 76 t C ha^–1 (N =137, CV: 50%) in 0–30 cm topsoil, and 62 t Cha^–1 (N = 156, CV: 46%) in0–20 cm topsoil. Extrapolating to national scale, we calculatedcontemporary SOC stocks of 110 Tg C (entire mineral soil, standarderror: 6 Tg C), 87 Tg C (0–30 cm topsoil, standarderror: 3.5 Tg C) and 70 Tg C (0–20 cm topsoil, standarderror: 2.5 Tg C) for mineral soils of accessible Swiss forests(1.1399 Mha). According to our estimate, the 0–20 cm layers ofmineral forest soils in Switzerland store about half of the Csequestered by forest trees (136 Tg C) and more than five times morethan organic horizons (13.2 Tg C).At stand level, regression analyses on the entire data set yielded nostrong climatic or topographic signature for forest SOC stocks in top(0–20 cm) and entire mineral soils across the Alps, despite thewide range of values of site parameters. Similarly, geostatisticalanalyses revealed no clear spatial trends for SOC in Switzerland at thescale of sampling. Using subsets, biotic, abiotic controls andcategorial variables (forest type, region) explained nearly 60%of the SOC variability in topsoil mineral layers (0–20 cm) forbroadleaf stands (N = 56), but only little of thevariability in needleleaf stands (N = 91,R ² = 0.23 for topsoil layers).Considerable uncertainties remain in assessments of SOC stocks, due tounquantified errors in soil density and rock fraction, lack of data onwithin-site SOC variability and missing or poorly quantifiedenvironmental control parameters. Considering further spatial SOCvariability, replicate pointwise soil sampling at 8–km × 8–kmresolution without organic horizons will thus hardly allow to detectchanges in SOC stocks in strongly heterogeneous mountain landscapes.