Continental-Scale Footprint of Balancing and Positive Selection in a Small Rodent (Microtus arvalis)

Genetic adaptation to different environmental conditions is expected to lead to large differences between populations at selected loci, thus providing a signature of positive selection. Whereas balancing selection can maintain polymorphisms over long evolutionary periods and even geographic scale, thus leads to low levels of divergence between populations at selected loci. However, little is known about the relative importance of these two selective forces in shaping genomic diversity, partly due to difficulties in recognizing balancing selection in species showing low levels of differentiation. Here we address this problem by studying genomic diversity in the European common vole (Microtus arvalis) presenting high levels of differentiation between populations (average FST = 0.31). We studied 3,839 Amplified Fragment Length Polymorphism (AFLP) markers genotyped in 444 individuals from 21 populations distributed across the European continent and hence over different environmental conditions. Our statistical approach to detect markers under selection is based on a Bayesian method specifically developed for AFLP markers, which treats AFLPs as a nearly codominant marker system, and therefore has increased power to detect selection. The high number of screened populations allowed us to detect the signature of balancing selection across a large geographic area. We detected 33 markers potentially under balancing selection, hence strong evidence of stabilizing selection in 21 populations across Europe. However, our analyses identified four-times more markers (138) being under positive selection, and geographical patterns suggest that some of these markers are probably associated with alpine regions, which seem to have environmental conditions that favour adaptation. We conclude that despite favourable conditions in this study for the detection of balancing selection, this evolutionary force seems to play a relatively minor role in shaping the genomic diversity of the common vole, which is more influenced by positive selection and neutral processes like drift and demographic history.

Climatology of Vb cyclones, physical mechanisms and their impact on extreme precipitation over Central Europe

Cyclones, which develop over the western Mediterranean and move northeastward are a major source of extreme weather and known to be responsible for heavy precipitation over the northern side of the Alpine range and Central Europe. As the relevant processes triggering these so-called Vb events and their impact on extreme precipitation are not yet fully understood, this study focuses on gaining insight into the dynamics of past events. For this, a cyclone detection and tracking tool is applied to the ERA-Interim reanalysis (1979–2013) to identify prominent Vb situations. Precipitation in the ERA-Interim and the E-OBS data sets is used to evaluate case-to-case precipitation amounts and to assess consistency between the two data sets. Both data sets exhibit high variability in precipitation amounts among different Vb events. While only 23 % of all Vb events are associated with extreme precipitation, around 15 % of all extreme precipitation days (99 percentile) over the northern Alpine region and Central Europe are induced by Vb events, although Vb cyclones are rare events (2.3 per year). To obtain a better understanding of the variability within Vb events, the analysis of the 10 heaviest and lowest precipitation Vb events reveals noticeable differences in the state of the atmosphere. These differences are most pronounced in the geopotential height and potential vorticity field, indicating a much stronger cyclone for heavy precipitation events. The related differences in wind direction are responsible for the moisture transport around the Alps and the orographical lifting along the northern slopes of the Alps. These effects are the main reasons for a disastrous outcome of Vb events, and consequently are absent in the Vb events associated with low precipitation. Hence, our results point out that heavy precipitation related to Vb events is mainly related to large-scale dynamics rather than to thermodynamic processes.

Pasture degradation modifies the water and carbon cycles of the Tibetan highlands

The Tibetan Plateau has a significant role with regard to atmospheric circulation and the monsoon in particular. Changes between a closed plant cover and open bare soil are one of the striking effects of land use degradation observed with unsustainable range management or climate change, but experiments investigating changes of surface properties and processes together with atmospheric feedbacks are rare and have not been undertaken in the world's two largest alpine ecosystems, the alpine steppe and the Kobresia pygmaea pastures of the Tibetan Plateau. We connected measurements of micro-lysimeter, chamber, 13C labelling, and eddy covariance and combined the observations with land surface and atmospheric models, adapted to the highland conditions. This allowed us to analyse how three degradation stages affect the water and carbon cycle of pastures on the landscape scale within the core region of the Kobresia pygmaea ecosystem. The study revealed that increasing degradation of the Kobresia turf affects carbon allocation and strongly reduces the carbon uptake, compromising the function of Kobresia pastures as a carbon sink. Pasture degradation leads to a shift from transpiration to evaporation while a change in the sum of evapotranspiration over a longer period cannot be confirmed. The results show an earlier onset of convection and cloud generation, likely triggered by a shift in evapotranspiration timing when dominated by evaporation. Consequently, precipitation starts earlier and clouds decrease the incoming solar radiation. In summary, the changes in surface properties by pasture degradation found on the highland have a significant influence on larger scales.

Flood triggering in Switzerland: the role of daily to monthly preceding precipitation

Determining the role of different precipitation periods for peak discharge generation is crucial for both projecting future changes in flood probability and for short- and medium-range flood forecasting. In this study, catchment-averaged daily precipitation time series are analyzed prior to annual peak discharge events (floods) in Switzerland. The high number of floods considered – more than 4000 events from 101 catchments have been analyzed – allows to derive significant information about the role of antecedent precipitation for peak discharge generation. Based on the analysis of precipitation times series, a new separation of flood-related precipitation periods is proposed: (i) the period 0 to 1 day before flood days, when the maximum flood-triggering precipitation rates are generally observed, (ii) the period 2 to 3 days before flood days, when longer-lasting synoptic situations generate "significantly higher than normal" precipitation amounts, and (iii) the period from 4 days to 1 month before flood days when previous wet episodes may have already preconditioned the catchment. The novelty of this study lies in the separation of antecedent precipitation into the precursor antecedent precipitation (4 days before floods or earlier, called PRE-AP) and the short range precipitation (0 to 3 days before floods, a period when precipitation is often driven by one persistent weather situation like e.g., a stationary low-pressure system). A precise separation of "antecedent" and "peak-triggering" precipitation is not attempted. Instead, the strict definition of antecedent precipitation periods permits a direct comparison of all catchments. The precipitation accumulating 0 to 3 days before an event is the most relevant for floods in Switzerland. PRE-AP precipitation has only a weak and region-specific influence on flood probability. Floods were significantly more frequent after wet PRE-AP periods only in the Jura Mountains, in the western and eastern Swiss plateau, and at the outlet of large lakes. As a general rule, wet PRE-AP periods enhance the flood probability in catchments with gentle topography, high infiltration rates, and large storage capacity (karstic cavities, deep soils, large reservoirs). In contrast, floods were significantly less frequent after wet PRE-AP periods in glacial catchments because of reduced melt. For the majority of catchments however, no significant correlation between precipitation amounts and flood occurrences is found when the last 3 days before floods are omitted in the precipitation amounts. Moreover, the PRE-AP was not higher for extreme floods than for annual floods with a high frequency and was very close to climatology for all floods. The fact that floods are not significantly more frequent nor more intense after wet PRE-AP is a clear indicator of a short discharge memory of Pre-Alpine, Alpine and South Alpine Swiss catchments. Our study poses the question whether the impact of long-term precursory precipitation for floods in such catchments is not overestimated in the general perception. The results suggest that the consideration of a 3–4 days precipitation period should be sufficient to represent (understand, reconstruct, model, project) Swiss Alpine floods.

Slab rollback orogeny in the Alps and evolution of the Swiss Molasse basin

The stratigraphies of foreland basins have been related to orogeny, where continent–continent collision causes the construction of topography and the downwarping of the foreland plate. These mechanisms have been inferred for the Molasse basin, stretching along the northern margin of the European Alps. Continuous flexural bending of the subducting European lithosphere as a consequence of topographic loads alone would imply that the Alpine topography would have increased at least between 30 Ma and ca. 5–10 Ma when the basin accumulated the erosional detritus. This, however, is neither consistent with observations nor with isostatic mass balancing models because paleoaltimetry estimates suggest that the topography has not increased since 20 Ma. Here we show that a rollback mechanism for the European plate is capable of explaining the construction of thick sedimentary successions in the Molasse foreland basin where the extra slab load has maintained the Alpine surface at low, but constant, elevations.

Early-Holocene afforestation processes in the lower subalpine belt of the Central Swiss Alps as inferred from macrofossil and pollen records

To reconstruct the vegetation history of the Upper Engadine, continuous sediment cores covering the past 11 800 years from Lej da Champfer and Lej da San Murezzan (Upper Engadine Valley, c. 1800 m a.s.l., southeastern Switzerland) have been analysed for pollen and plant macrofossils. The chronologies of the cores are based on 16 and 22 radiocarbon dates, respectively. The palaeobotanical records of both lakes are in agreement for the Holocene, but remarkable differences exist between the sites during the period 11 100 to 10 500 cal. BP, when Lej da Champfer was affected by re-sedimentation processes. Macrofossil data suggest that Holocene afforestation began at around 11400 cal. BP. A climatic deterioration, the Preboreal Oscillation, stopped and subsequently delayed the establishment of trees until c. 11000 cal. BP, when first Betula, then Pinus sylvestrislmugo, then Larix 300 years later, and finally Pinus cembra expanded within the lake catchment. Treeline was at c. 1500 m during the Younger Dryas (12 542- 11 550 cal. BP) in the Central Alps. Our results, along with other macrofossil studies from the Alps, suggest a nearly simultaneous afforestation (e.g., by Pinus sylvestris in the lower subalpine belt) between 1500 and 2340 m a.s.l. at around 11 400 to 11 300 cal. BP. We suggest that forest-limit species (e.g., Pinus cembra, Larix decidua) could expand faster at today's treeline (c. 2350 m a.s.l.), than 550 m lower. Earlier expansions at higher altitudes probably resulted from reduced competition with low-altitude trees (e.g. Pinus sylvestris) and herbaceous species. Comparison with other proxies such as oxygen isotopes, residual A14C, glacier fluctuations, and alpine climatic cooling phases suggests climatic sensitivity of vegetation during the early Holocene.

Zur Langzeitökologie des Lärchen-Arvengürtels in den südlichen Walliser Alpen

Palaeoecological studies in the "Alpe d'Essertse" area have provided much information about Vegetation changes and timberline fluctuations during the Holocene In this study we repeated previous biostratigraphic investigations using plant macrofossils to improve their temporal and taxonomie resolution and to test their reliability. By analyzing 0.5-cm layers of a lake sediment we reached a temporal resolution of 44 years, and we were able to reconstruct Vegetation changes in the surrounding area at species level. The sedimentary record analyzed extends from the Late-Glacial to the late Holocene Alpine grasslands (12'000-11'000 cal. BP) were afforested by Larix decidua, Juniperus nana, and Pinus cembra (11'000-9'600 cal. B.P). Stable subalpine larch-stone pine-forests (9'600^4'900 cal. BP) were followed by shrublands and meadows as a consequence of the climatically and anthropogenically induced destruction of forest Vegetation (4'900-2'600 cal. BP). Changes in the abundance of P. cembra and L. decidua needles as well as changes of the other taxa were consistent with those found in previous studies from the same lake. Our results demonstrate that plant-macrofossil records can be reproduced spatially and temporally on separate cores with independent 14C chronologies.

Rapid responses of high-mountain vegetation to early Holocene environmental changes in the Swiss Alps

1 The Early Holocene sediment of a lake at tree line (Gouillé Rion, 2343 m a.s.l.) in the Swiss Central Alps was sampled for plant macrofossils. Thin (0.5 cm) slices, representing time intervals of c. 50 years each from 11 800 to 7800 cal. year bp, were analysed and the data compared with independent palaeoclimatic proxies to study vegetational responses to environmental change. 2 Alpine plant communities (e.g. with Salix herbacea) were established at 11 600–11 500 cal. year bp, when oxygen-isotope records showed that temperatures increased by c. 3–4 °C within decades. Larix decidua trees reached the site at c. 11 350 cal. year bp, probably in response to further warming by 1–2 °C. Forests dominated by L. decidua persisted until 9600 cal. year bp, when Pinus cembra became more important. 3 The dominance of Larix decidua for two millennia is explained by dry summer conditions, and possibly low winter temperatures, which favoured it over the late-successional Pinus cembra. Environmental conditions were a result of variations in the earth's orbit, leading to a maximum of summer and a minimum of winter solar radiation. Other heliophilous and drought-adapted species, such as Dryas octopetala and Juniperus nana, could persist in the open L. decidua forests, but were out-competed when the shade-tolerant P. cembra expanded. 4 The relative importance of Larix decidua decreased during periods of diminished solar radiation at 11 100, 10 100 and 9400 cal. year bp. Stable concentrations of L. decidua indicate that these percentage oscillations were caused by temporary increases of Pinus cembra, Dryas octopetala and Juniperus nana that can be explained by increases in moisture and/or decreases in summer temperature. 5 The final collapse of Larix decidua at 8400 cal. year bp was possibly related to abrupt climatic cooling as a consequence of a large meltwater input to the North Atlantic. Similarly, the temporary exclusion of Pinus cembra from tree line at 10 600–10 200 cal. year bp may be related to slowing down of thermohaline circulation at 10 700–10 300 cal. year bp. 6 Our results show that tree line vegetation was in dynamic equilibrium with climate, even during periods of extraordinarily rapid climatic change. They also imply that forecasted global warming may trigger rapid upslope movements of the tree line of up to 800 m within a few decades or centuries at most, probably inducing large-scale displacements of plant species as well as irrecoverable biodiversity losses.

Migration and population expansion of Abies, Fagus, Picea, and Quercus since 15000 years in and across the Alps, based on pollen-percentage threshold values

Aims: The aim of this study is to explore the migration (colonization of new areas) and subsequent population expansion (within an area) since 15 ka cal BP of Abies, Fagus, Picea, and Quercus into and through the Alps solely on the basis of high-quality pollen data. Methods: Chronologies of 101 pollen sequences are improved or created. Data from the area delimited by 45.5–48.1°N and 6–14°E are summarized in three ways: (1) in a selection of pollen-percentage threshold maps (thresholds 0.5%, 1%, 2%, 4%, 8%, 16%, and 32% of land pollen); (2) in graphic summaries of 250-year time slices and geographic segments (lengthwise and transverse in relation to the main axis of the Alps) as pollen-percentage curves, pollen-percentage difference curves, and pollen-percentage threshold ages cal BP graphed against both the length and the transverse Alpine axes; and (3) in tables showing statistical relationships of either pollen-percentage threshold ages cal BP or pollen expansion durations (=time lapse between different pollen-percentage threshold ages cal BP) with latitude, longitude, and elevation; to establish these relationships we used both simple linear regression and multiple linear regression after stepwise-forward selection. Results: The statistical results indicate that (a) the use of pollen-percentage thresholds between 0.5% and 8% yield mostly similar directions of tree migration, so the method is fairly robust, (b) Abies migrated northward, Fagus southward, Picea westward, and Quercus northward; more detail does not emerge due to an extreme scarcity of high-quality data especially along the southern foothills of the Alps and in the eastern Alps. This scarcity allows the reconstruction of one immigration route only of Abies into the southern Alps. The speed of population expansion (following arrival) of Abies increased and of Picea decreased during the Holocene, of Fagus it decreased especially during the later Holocene, and of Quercus it increased especially at the start of the Holocene.

NEENAWA: Network in Eastern European Neolithic and Wetland Archaeology: first steps.

Neolithic and Bronze Age wetland sites around the Alps (so called pile-dwellings, Pfahlbauten or palafittes in German/French) are of outstanding universal value (UNESCO-world heritage since 2011). Typical sites are in lakes, rivers and bogs, dating between 5300 and 800 BC. Of common character is the perfect conservation of wood, textiles from plant fabrics and many other organic materials. Larger quantities of sub-fossilized wood, as in the peri-alpine sites, offer the possibility of high-precision dating by dendrochronology. Research in these wetland sites started in the mid-19th century. Through large scale rescue excavations since the 1970s and the evolution of underwater archaeology in the same period the Swiss accumulated a thorough experience with these specific sites. Research in wetland sites is shared between cantonal institutions and universities and led to a worldwide unique accumulation of knowledge. Comparable sites exist outside of the Alpine area, but in much smaller quantities. Regions like Russia (small lakes in NW-Russia) and Macedonia (medium size lakes in the border zone of Macedonia, Albania and Greece) have a high scientific potential; rivers in Ukraine are supposed to have the same type of sites.

Late Glacial and Holocene environmental history of the Pirin Mountains (SW Bulgaria): a paleolimnological study of Lake Dalgoto (2310 m)

Diatoms, Cladocera, and chironomids preserved in the sediments of Lake Dalgoto were studied to reconstruct the history of the lake ecosystem in the context of the vegetation history as represented by the pollen stratigraphy. Younger Dryas silty sediments at the base of the core are characterized by low diversity of aquatic organisms. The transition to the Holocene is indicated by a sharp change from silt to clay-gyttja. The migration and expansion of trees at lower elevations between 10200 and 8500 14C-yr BP, along with higher diversities and concentrations of aquatic organisms and the decreased proportion of north-alpine diatoms, point to rapidly rising summer temperatures. After 6500 14C-yr BP the expansion of Pinus mugo in the catchment coincides with signs of natural eutrophication as recorded by an increase of planktonic diatoms. In the late Holocene (4000–0 14C-yr BP) Pinus peuce and Abies are reduced and Picea expands. Cereal grains and disturbance indicators suggest late-Holocene human modification of the vegetation.

Climate-pollen relationships AD 1901-1996 in two small mires near the forest limit in the northern and central Swiss Alps

Climatic relationships were established in two 210Pb dated pollen sequences from small mires closely surrounded by forest just below actual forest limits (but about 300 m below potential climatic forest limits) in the northern Swiss Alps (suboceanic in climate; mainly with Picea) and the central Swiss Alps (subcontinental; mainly Pinus cembra and Larix) at annual or near-annual resolution from ad 1901 to 1996. Effects of vegetational succession were removed by splitting the time series into early and late periods and by linear detrending. Both pollen concentrations detrended by the depth-age model and modified percentages (in which counts of dominant pollen types are down-weighted) are correlated by simple linear regression with smoothed climatic parameters with one-and two-year timelags, including average monthly and April/September daylight air temperatures and with seasonal and annual precipitation sums. Results from detrended pollen concentrations suggest that peat accumulation is favoured in the northern-Alpine mire either by early snowmelt or by summer precipitation, but in the central-Alpine mire by increased precipitation and cooler summers, suggesting a position of the northern-Alpine mire near the upper altitudinal limit of peat formation, but of the central-Alpine mire near the lower limit. Results from modified pollen percentages indicate that pollen pro duction by plants growing near their upper altitudinal limit is limited by insufficient warmth in summer, and pollen production by plants growing near their lower altitudinal limit is limited by too-high temperatures. Only weakly significant pollen/climate relationships were found for Pinus cembra and Larix, probably because they experience little climatic stress growing 300 m below the potential climatic forest limit.

Climatic change and contemporaneous land-use phases north and south of the Alps 2300 BC to 800 AD

Fluctuations in the Δ14C curve and subsequent gaps of archaeological findings at 800–650 and 400–100 BC in western and central Europe may indicate major climate-driven land-abandonment phases. To address this hypothesis radiocarbon-dated sediments from four lakes in Switzerland were studied palynologically. Pollen analysis indicates contemporaneous phases of forest clearances and of intensified land-use at 1450–1250 BC, 650–450 BC, 50 BC–100 AD and around 700 AD. These land-use expansions coincided with periods of warm climate as recorded by the Alpine dendroclimatic and Greenland oxygen isotope records. Our results suggest that harvest yields would have increased synchronously over wide areas of central and southern Europe during periods of warm and dry climate. Combined interpretation of palaeoecological and archaeological findings suggests that higher food production led to increased human populations. Positive long-term trends in pollen values of Cerealia and Plantago lanceolata indicate that technical innovations during the Bronze and Iron Age (e.g. metal ploughs, scythes, hay production, fertilising methods) gradually increased agricultural productivity. The successful adoption of yield-increasing advances cannot be explained by climatic determinism alone. Combined with archaeological evidence, our results suggest that despite considerable cycles of spatial and demographic reorganisation (repeated land abandonments and expansions, as well as large-scale migrations and population decreases), human societies were able to shift to lower subsistence levels without dramatic ruptures in material culture. However, our data imply that human societies were not able to compensate rapidly for harvest failures when climate deteriorated. Agriculture in marginal areas was abandoned, and spontaneous reforestations took place on abandoned land south and north of the Alps. Only when the climate changed again to drier and warmer conditions did a new wide-spread phase of forest clearances and field extensions occur, allowing the reoccupation of previously abandoned areas. Spatial distribution of cereal cultivation and growth requirements of Cerealia species suggest that increases in precipitation were far more decisive in driving crop failures over central and southern Europe than temperature decreases.

Wald im Welterbe - Vielfältig, dynamisch und sehr wertvoll

Wälder übernehmen wichtige und vielfältige Funktionen für die Gesellschaft: Sie versorgen uns nicht nur mit Holz, sie schützen auch vor Naturgefahren und säubern Wasser und Luft. Fast die Hälfte der in der Schweiz vorkommenden Tiere und Pflanzen ist auf den Wald als Lebensraum angewiesen. Der Wald – oft Sinnbild von Natur – prägt in seiner Mannigfaltigkeit die Landschaft, in der wir leben. Während auf unserem Planeten täglich grosse Waldflächen verschwinden, breitet sich der Wald im Alpenraum aus und verändert die alpine Landschaft, so auch in der Region unseres Welterbes.