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JOURNAL OF FOREST SCIENCE, 61, 2015 (11): 496–514

doi: 10.17221/47/2015-JFS

Climate change impacts on the Alpine : an overview with focus on the soil – a review

S. Chersich1, K. Rejšek2, V. Vranová2, M. Bordoni1, C. Meisina1

1University of Pavia, Department of Earth and Environmental Sciences, Pavia, Italy 2Mendel University in Brno, Faculty of Forestry and Wood Technology, Department of Geology and Soil Science, Brno, Czech Republic

ABSTRACT: The Alpine ecosystem is very sensitive to climatic changes, which have an influence on glaciers, , and soils. The aim of this review is to illustrate the effects of global change on the Alpine soil ecosystem, which is an optimal marker to record them. The manuscript enhances our understanding of the global change effect on the Alpine environment: on morphology, on ice, on vegetation and points out how the cycles of soil nutrients equi- librium have been changed with a direct effect on soils that support species. The changes in cryosphere, glacier reduction and periglacial environment as glaciers retreat, decrease in the snow cover extent and earlier snowmelt, determine an effect on soils (on the structure, organic matter and humus forms, soil processes and soil types) from the top of the Alpine horizon to the bottom. The processes induced by change (such as erosion and shifting) have a direct effect on water balance that can be observed on soil profile characters with an effect on upward migration, change in phenology, extensive losses of species. The equilibrium of the biogeochemical cycles has been changed and this has a direct effect on soils that support plant species.

Keywords: Alpine environment; cryosphere; soil science; treeline

The European Alpine belt is a and and to suggest appropriate forest, range stretching along an arc of about 1,200 km farming, environment management and policies (190,000 km2), with about 13 million inhabitants. of the European society (OECD 2007; Gobiet It has not only an environmental value but also et al. 2014). It is known that the climate plays an a social one [e.g. the amount of water delivered important role in every ecosystem, particularly in by the allocates 40% of EU consumption, see the Alps, where meteorological factors, combined EEA (2009)]. The importance of the Alps has been with severe morphological conditions, often show declared officially on March 6th, 1995, when the extreme behaviours. Alpine Convention came into force, signed by five The climate is a principal factor governing the OECD members. It has been declared that the natural environment of on short-time Alps are a territory with a large biodiversity and scales (Beniston 2001), and it characterizes the lo- with a delicate balance that needs protection, and cation and the intensity of biological, physical and then the International Commission for the pro- chemical processes (Vranová et al. 2011; Brad- tection of the Alps (CIPRA) has been established. ley et al. 2015). Mountain are governed Recent studies have shown that it is very impor- by four major factors, namely continentality, lati- tant to define the evidences of the global change tude, altitude and topography. Altitude is certainly to monitor the Alpine , in order to pro- the most distinguishing and fundamental charac- tect and support the Alpine landscapes, habitats teristic of mountain climates (Chimani et al. 2013;

Supported by the Mendel University in Brno, Czech Republic, Grant No. IGA FFWT MENDELU 55/2013

496 J. FOR. SCI., 61, 2015 (11): 496–514 Morán-Tejeda et al. 2013), while topographic fea- vegetational cover had a considerable effect on the tures play a key role in determining local climates, water balance of biotopes (D'Amico et al. 2014). in particular due to slope, aspect and exposure of When the vegetation structure changes, alterations the surface to climatic elements (Beniston 2006; of transpiration flow, accumulation of snow and its Liancourt et al. 2013). melting speed, soil surface evaporation and a depth The singularity of the Alpine climate is a high de- of soil freezing modify those processes that control gree of complexity, due to interactions between the the water balance of a habitat (Beniston, Stoffel mountains and the general circulation of the atmos- 2014; Zongxing et al. 2014). phere (Samec et al. 2006), which result in features The climate in the Alps from the late 19th cen- such as gravity wave breaking, blocking highs, and tury to the end of the 20th century was character- föhn . A further cause of complexity results ised by an increase of the mean annual tempera- from the competing influences of a number of differ- tures by about 2°C, by a more modest increase of ent climatological regimes, namely Mediterranean, maximum and by a trend in the av- Continental, Atlantic and Polar (Beniston 2005). erage data (Beniston 2006; Auer et The complex topography of mountain regions, al. 2007; Solomon et al. 2007). It is estimated that especially in young ranges as the Alps, determines the atmospheric CO2 will rise and it will increase sharp gradients in climatic parameters, in particu- the surface of the Earth by 0.25°C per lar temperature and precipitation, also over very decade throughout the current century (Gobiet et short distances. As a consequence, mountains ex- al. 2014). The aim of this review is to summarize hibit large biodiversity, often with sharp transitions the principal highlights of the con- from vegetation and soil to snow and ice (Benis- sequences in the Alpine environment with a special ton 2006; Schworer et al. 2014). focus on the soil and to emphasize some of the ele- Mountains are susceptible to the impacts of a ments of interactions between the soil, the vegeta- rapidly changing climate and provide interesting tion and the environmental components. locations for early detection and study of the sig- nals of climate change and its impacts on hydro- logical, ecological and societal systems (Cannone Background and aims et al. 2008; Kohler, Maselli 2009; Sugiero et al. 2009; Wolf et al. 2012). Human settlement does We analyse recent literature to describe the effect not explain the marked differences between the po- of the global change on the Alpine land: on morphol- tential and contemporary actual vegetation in the ogy, on ice, on vegetation and on soil, from the top of Alpine environment (Gerlach et al. 2006; Ever- the Alpine horizon (2,400–2,600 m above sea level) shed 2008; Hjulström, Isaksson 2009; Miehe to the bottom. With regard to the response of soils et al. 2014) – the ecological continuity of parent to climate change (Rejšek 2004) dominant soil pro- material and the nowadays vegetation was broken cesses and their distribution are taken into account by both processes occurring after Dryas 3 (the end using the classification of the IUSS Working Group of the Würm period) and the continuing Holocene WRB, 2006. The discussion about the organic hori- global climate changes (Simonneau et al. 2012). zons of soil was referred to in Zanella et al. (2011). Therefore, the Alps represent unique areas for the The effect of climate changes on landscape dy- detection of global change and its environmental namics can be investigated on geomorphic features involvements because of several reasons: 1 – as the deposited or produced during the Late Glacial Max- climate changes rapidly with altitude over relatively imum or Holocene period (Kramer et al. 2010), short horizontal distances, so do vegetation and hy- when a dramatic temperature increase was recorded drology (Scherrer, Körner 2011) and relatively within a relatively short time interval (Stoffel et al. small perturbations in global processes can cascade 2014). That situation was similar to what is happen- down to produce large changes (Helbing 2013); ing nowadays in lands with similar climate and con- 2 – in mountain ecosystems, it is possible to in- textual challenges such as , vestigate impacts of global warming in the absence (Snowy Mountains) and . of direct or significant human interference (Ben- The velocity of landscape dynamics can be assessed iston 2006; You et al. 2014); 3 – because of the by studying the tree line shifting and through a com- abundance of available literature data and monitor- bination of relative (Goudiet 2006) and absolute dat- ing actions about climate and environment in their ing techniques applied to soil and charred material, territory. In addition, the Alps represent unique or/and to rocks (e.g. Favilli et al. 2008, 2009; Hor- areas also from the viewpoint that such changes of mes et al. 2008; Cossarta 2010).

J. FOR. SCI., 61, 2015 (11): 496–514 497 Regional Circulation Models applied to the Euro- possible reason for this in the presence of persistent pean Alps, General Circulation Models and the In- high surface pressure fields over the Alpine region tergovernmental Panel on Climate Change Fourth during autumn and winter, which are characterized Assessment Report represent an effort to charac- by warm temperatures and low precipitation (Boc- terize the Alpine climate in the latter part of the chiola, Diolaiuti 2010; Soncini, Bocchiola 21st century (Solomon et al. 2007). 2011). The Alpine temperature and precipitation We have considered all the cryosphere compo- variability and the fluctuations of high pressure nents: glaciers, and snow. Regarding the episodes are strongly correlated with anomalies of periglacial environment, we investigate the variation the North Atlantic Oscillation index and they are of snow cover, the retreat of glaciers. We analysed controlled by the atmospheric circulation modes the morphological aspects related to slope stabil- (Casty et al. 2005). Other studies confirmed that ity and hydrology like in the case of “rock glaciers”, the days with snow cover on the ground decreased and also the signs within the soil (e.g. stability of ag- in the last twenty years, particularly in sites at mid gregates) that are the “memory of the soil” (Targu- and low altitudes, where the duration of continuous lian, Goryachkin 2011) and indices of soil erod- snow cover and the number of snowfall days de- ibility. The authors state that different acclimation creased and precipitation changed from solid to liq- mechanisms of various floral taxa led by their dis- uid state, with subsequent more rain on snow events tinct water demands and responsiveness to draught in spring time (Laternser, Schneebeli 2003; have been reflected in some soil parameters such Szczypta et al. 2015). The role of snow in the cli- as soil structure, soil organic matter (SOM) and its mate system includes strong positive feedbacks re- mineralization that interact with the nutrient cycle lated to albedo and other weaker feedbacks related and microbial communities and are strictly depen- to moisture storage, latent heat and insulation of the dent on climate change (Bradley et al. 2015). underlying surface (Déry, Brown 2007; Qingbai et al. 2015). The lack of snow cover reduces the de- gree of insulation and results in cold soil tempera- The cryosphere and glacier recession tures, extensive soil freezing and increase in freeze/ thaw cycles, influencing soil microbiological activ- In the climate system, the cryosphere (which con- ity and SOM (Soil Organic Matter) mineralization sists of glaciers, permafrost and snow on the land (Edwards, Cresser 1992; Groffman et al. 2001; and lake and sea ice as well) is intricately linked to Clement et al. 2012; Welc et al. 2014). Glaciers are the surface energy budget, water cycle, sea level among the most important elements of the cryo- change and surface gas exchange (Wojciech 2012). sphere and many works have been written in the last The cryosphere integrates climate variations over a years which studied glacier variations in relation to wide range of time scales; it is a natural sensor of global warming (Le Roy et al. 2015). The first field climate variability and provides a visible expression survey to detect glacier terminus fluctuations start- of climate change. In the past, the cryosphere un- ed in Switzerland at the end of the 19th century and derwent large variations on many time scales as- later in Italy and France (in the Alps). In some cases, sociated with ice ages and with shorter-term varia- data dating back to the 17th century are available but tions like the or the Little Ice Age not from direct field surveys. These long series were (Solomon et al. 2007; Chiri et al. 2015). reconstructed from dendrochronologic evidences Solomon et al. (2007) shows that at the present or are based on other glacier proxies. A method time, snow covers more than 33% of lands north of to evaluate the retreat or advance of glaciers was the equator from November to April, reaching 49% to measure the length variation of glacier tongues coverage in January. Since the early 1920s, and es- (Oerlemans 2005), while recently (mid-20th cen- pecially since the late 1970s, the snow covered area tury) glacier mass balance measurements have been (SCA) in the Northern Hemisphere has declined in performed to assess the gain or loss of ice volume, spring and summer, but not substantially in winter. with particular attention to hydrological purposes Over the 1922 to 2005 period, the linear trend of the [WGMS (ICSI-IAHS), several years]. SCA in March and April has undergone a statisti- In general, during the last decades, after the short cally significant reduction of 7.5 ± 3.5% (Solomon positive impulse at the end of the 1980’s, Alpine gla- et al. 2007). Beniston (1997) showed that since the ciers have been continuously retreating. The Italian mid-1980s, the length of the snow season and snow Alpine glaciers have decreased their area up to 20–22% amount have substantially decreased in the Alps, in in the time window 1990–2003 in Lombardy (Cit- particular at altitudes below 1,500 m and found a terio et al. 2007; Diolaiuti et al. 2012), up to 30%

498 J. FOR. SCI., 61, 2015 (11): 496–514 in the time frame 1975–2005 in Aosta Valley (Dio- soil and the cracks in rocks. The term permafrost laiuti et al. 2012) and a decrease by about 40% “vechnaja merzlota” (eternal frost) was first intro- of their area was found studying Ortles Cevedale duced to describe the condition of earth materials glaciers in the period 1954–2007 (D’Agata et al. that remain below 0°C continuously for two years 2013). From 1850 to 2000, glaciers in the European at least (Muller 1945; Muller et al. 2008). It was Alps lost almost 50% of their area, the largest reduc- observed for the first time in the everlasting thick tion (22%) occurred after 1985 (Zemp et al. 2006). frozen grounds of Central Siberia. It is defined ex- For nine measured glaciers in the Alps, Zemp et al. clusively on the basis of temperature, irrespective (2005) estimated a mean loss of about 600 kg·m–2·yr–1 of the type of earth material, water content, degree from 1980 to 2001, with an exceptional mass loss of of indurations, or lithological characteristics. The 2,500 kg·m–2·yr–1 during 2003. In terms of the vol- distribution and thickness of permafrost is gov- ume, Alpine glaciers have lost on average 1% of their erned by the energy balance between the earth and volume per year since 1975, being reduced from the atmosphere (Gerrard 1992; Gu et al. 2015). 105 × 10 km3 to 75 × 10 km3 at the beginning of the The permafrost is associated with the high C-stocks third millennium (Paul et al. 2004). Zemp et al. (down to the C horizon or rock surface it is 10 to (2005) supposed that the Alpine glacier cover would 15 kg/m2 by Zollinger et al. (2013). decrease by 80% until the middle/end of this mil- Permafrost in the Alps is negatively affected by lennium if there were an increase in temperature the global change, mainly due to warmer air tem- by 3°C, as expected if humans did not start to re- perature, lower snowfall and shorter winter season. act to climate change by adopting mitigation policy According to Solomon et al. (2007) and Li et al. (Solomon et al. 2007). An important aspect is that (2015), the decrease of the thickness and/or the the air temperature has a higher effect on the gla- extent of permafrost is a worldwide ongoing pro- cier mass balance than precipitation; therefore an cess that is increasing the permafrost active layer increase in precipitation by 20% will bring only the (Schuur et al. 2008; Wang et al. 2014). previous value from 80 to 65% (Zemp et al. 2005). Rock glaciers are typical geomorphological fea- From the vegetation point of view, such a decrease in tures of the periglacial environment and are char- the Alpine glacier volumes (Solomina et al. 2015) acteristic of high mountain permafrost. They are led to the ongoing process of re-colonisation and de- made up of debris and ice that creep on slopes at a velopment of new ecosystems (Pichler et al. 2007) typical rate of a few decimetres per year (Serrano in which the fast decline of pioneer vegetation oc- et al. 2006). They are indicators of the present-day curred. Besides, at the elevation threshold of 2600 m climate (especially thermal conditions) and have a.s.l. the resident species may create a migration bar- also been used for reconstructing the palaeocli- rier for the new colonization (Cannone et al. 2008; mate and landscape evolution. Detailed rock gla- Cannone, Pignatti 2014). cier dynamics was investigated in arctic and Alpine The higher temperature causes the contraction of environments in the late 20th century (Serrano snowbeds that shift to higher altitudes where the et al. 2006; Haeberli et al. 2007). The obtained snow-dependent that decrease in coverage displacement rate in the frontal part of the alpine may find the refuge Cannone( et al. 2008). rock glacier is up to 0.6–0.9 m·yr–1 (by geodetic data, Lugon, Stoffel 2010). The relict form and the dynamics of rock glaciers are usually related The periglacial environment to palaeoclimatic changes (e.g. Frauenfelder et al. 2005; Kääb et al. 2007). For instance, speed The deglaciated areas, where soils and rocks are ex- variations change with frequencies in the order of posed to different conditions, are called periglacial en- several decades or centuries and can result from a vironment. The term periglacial was first introduced general spatial-temporal modification in boundary to describe the climatic and geomorphic conditions conditions, such as material and water/ice supply peripheral to the late Pleistocene ice sheets but the or thermal regime, or terrain topography (Frauen- term has undergone a substantial revision and no uni- felder et al. 2005). Photogrammetric and geodetic versally accepted definition exists. The term has been monitoring series of several decades suggest that generalized to include those environments where velocity changes of the mountain permafrost creep climatic conditions result in severe frost action that at pluriannual time-scales may be due to variations dominates geomorphic processes (French 1976). in and/or climate conditions (Paul et al. In the periglacial environment, permafrost has 2004; Roer et al. 2005). At shorter time-scales, the been developed – ice is formed within both the sensitivity of the creep of frozen slopes to external

J. FOR. SCI., 61, 2015 (11): 496–514 499 climate forcing is strongly dependent on individ- Geotechnical changes of frozen slopes and short- ual internal conditions of rock glaciers, as clearly term changes in triggering rainfall have a notable shown by observations of seasonal velocity varia- impact on slope stability, and are thus of interest tions (Kääb et al. 2007). for dealing with applied problems such as con- The question how the weather and the atmo- struction stability, debris-flow hazards, or rockfall spheric warming, which are currently observed in hazards (Noetzli et al. 2007; Kääb et al. 2007; many mountain regions (Kääb et al. 2007), may Bollschweiler, Stoffel 2010). influence the creep of perennially frozen debris Moreover, increasing the thickness of the per- has not been completely solved. Kääb et al. (2007) mafrost active layer and changes in hydrology at showed the potential reaction of perennially frozen higher altitudinal levels will change the ecologi- slopes to ground temperature variations. Boeck- cal conditions on steep mountain slopes, making li et al. (2012) developed a statistical approach to them much more unstable, especially where the modelling permafrost distribution in the Alps and incidence of precipitation events grows, probably they developed a theoretical model of mountain causing a higher frequency and maybe intensity of permafrost reaction to climate change. Davies et avalanches and landslides (Beniston 2006) and al. (2001) found the minimum shear strength of higher erosion rates (Fischer et al. 2013). frozen rock joints at temperatures between –1 and 0°C using centrifuge modelling. Furthermore, some authors have collected various indications that Specificity of soils in Alpine ecosystems the deformation of frozen debris or rock glaciers Soil structure increases with increasing ground temperatures (e.g. Gruber, Haeberli 2007), or have proposed To study the effects of global change on soil struc- creep laws which include a dependence on the ture and physical properties, obviating the complex ground temperature (Kääb et al. 2007). mechanisms that bound climate and structure gen- As above-mentioned, changes in phytocoenoses esis, it is required to investigate aggregate stability. affect the soil evolution. The distinctive points -re This soil property is a good indicator of soil struc- lated to studies of changes affecting the alpine en- ture susceptibility to runoff and erosion Nyberg( vironments (Cannone, Pignatti 2014; Egli et et al. 2001; Barthès, Roose 2002) and, according al. 2014) can be seen especially in uncertainties in to Lavee et al. (1996), of the soil degradation sta- the roles of soil biomass, carbon (Bradshaw et al. tus as a consequence of global change, because it is 2015) and nitrogen (Qi et al. 2011) controlled by sensitive to warming and because it responds sea- local climate oscillations (Xu et al. 2015), interre- sonally to precipitation (Brunetti et al. 2006). lationships between the microrefuges and the local Barthès and Roose (2002) demonstrated that geomorphology (D’Amico et al. 2014; Gentili et al. the topsoil aggregate stability is closely and nega- 2015) and synergic effects that reinforce these pro- tively related to soil susceptibility to erosion by cesses (Smith-McKenna et al. 2014). The useful- runoff, especially in consideration of slope gradient ness of local studies focused on distinctive reasons and climate aggressiveness. Aggregate stability is (Anderson et al. 2011; Bowman et al. 2012) is also expected to decrease with an increase in the fre- indisputable. Díaz-Varela et al. (2010) described a quency of dry periods followed by almost instan- novel procedure to delineate the current and former taneous flash flood phenomena, as a consequence state of ecotones of alpine phytocoenoses which was of an embittered slaking effect. This phenomenon, tested in the Italian Alps for the period 1957–2003 especially in soils with low exchangeable sodium showing a trend of an increase in altitude for the percentage and swelling clay content, like most Al- ecotones scrutinized. The permafrost degradation is pine poorly developed soils, may be considered the associated with the vegetation cover decline and dy- most important mechanism contributing to the soil namics (Cannone et al. 2008). Ecological responses aggregate breakdown. This process is recorded in of plant species and communities to climate change the clay cutans of horizons with the “soil memory” show a range filling processes with downward shifts (Targulian, Goryachkin 2011) of the climate of the persisting species (Cannone, Pignatti change related with a moisture regime change. 2014). Rydgren et al. (2011) and Valese et al. The more humid climate is linked with more illu- (2014) highlighted the role of human disturbances in vial cutans that are records of humid percolating the alpine environments for a direction in vegetation conditions and translocation of clay. Silt and clay succession and reinforced the role of soil properties particles within the soil body (cutans as structur- (Burga et al. 2010), soil texture in particular. ing agents in Bt horizons) are records of increased

500 J. FOR. SCI., 61, 2015 (11): 496–514 aggressiveness of the solution along fissures, which will be most available and it will flow in areas of al- destroy silicates in cutans more intensively (Tar- luvial cones and Fluvisols will increase their range. gulian, Goryachkin 2011). Fluvisols are genetically young soils, azonal soils Aggregate stability has already been used as soil on the more stable portion of alluvial cones, on the erodibility indicator in the Universal Soil Loss high Old Holocene terraces and lacustrine deposits Equation (Wischmeier, Smith 1978) to model (Simonneau et al. 2012). Their profiles show evi- erosion rate by measured losses of soil. dences of stratification, different granulometry is However, the amount of field data available for an indicator of the energy and duration of the past establishing the influence of climate change im- climate events. The organic horizon of Fluvisols is pacts on the multivariate factor of aggregation and primarily a humus form of Mull with humified and on several key processes of water erosion is very biologically very active organic matter. The Mull limited, also considering the multiple effects that starts to be formed when the vernal snow cover climate expresses on ecosystems (Confortola et leaves the soil free. al. 2013). Moreover, despite their importance in Al- In mountain areas higher temperatures ensure a pine Regions, according to Solomon et al. (2007) longer duration of the vegetation season, a reduc- it appears that the analysis of extraordinary rainfall tion of the frequency of frosts and therefore the events and their effects on ecosystems is still un- better coverage of the ground surface, the devel- derdeveloped, like the impacts of global change on opment of humus forms of Mull reducing erosion soil structure and alpine soil susceptibility to ero- rates and increasing the stability of soil. sion (Kirkpatrick et al. 2014). Vice versa, in some upland areas, erosion rates will be reduced as a result of the better soil surface Cryosols, Regosols, Fluvisols and Leptosols cover and topsoil stability arising from higher tem- Strongly related to a decrease in the extent and an peratures that extend the duration of the growing increase in the altitude of permafrost are Cryosols, season and reduce the number of frosts. Cannone i.e. the soils which show evidence of cryogenic pro- et al. (2007) described two patterns for the pioneer cesses (Arkhangelskaya 2014), cryoturbation vegetation: (i) at lower altitudes, the area of pio- and contain water in solid form (permafrost con- neer vegetation increases especially in the vicinity dition). The horizons are cryoturbated, they show of streams, alluvial and debris flow fans, and ii( ) at frost heave, thermal cracking, and higher altitudes, there is a regression in the areas patterned ground microrelief. They occur on top of of pioneer vegetation, on steep slopes in particular. the Alpine horizon (2,400–2,600 m a.s.l.) and are Today Regosols occupy a very large area. Rego- very vulnerable to the change of temperature. They sols are young, undeveloped soils, composed of a are associated with the specific belt where the soil wide variety of textures displaying the Moder hu- temperature is continuously below 0°C for several mus form which relates to the prominent fungi years at least (Kurylyk et al. 2014). Obviously, the and arthropod activities (Chersich 2007). Under global climate changes have an impact on perma- a warmer climate the Regosol area will be reduced frost geothermal regimes in the Alpine mountains because it is very likely that developing Bw hori- (Harris et al. 2010). The raising of the suitable zons will start massively. lower boundary (Bauman et al. 2014) for Cryosols The biological activity will tend to increase and (and their consequent reduction), which is cor- with the incorporation of organic matter a devel- related with the increase of the permafrost lower oped structure is expected to form (with transfor- boundary, has great consequences in the Alpine mation of humus from Moder forms and Mull and environment (Fouché et al. 2014) where the slopes the formation of granular structure and subangular are not stabilized, rockfalls or landslides might oc- polyhedrals; Chersich et al. 2006). Bulk density cur at high elevations, causing serious impacts on and pH will increase while organic acids and the the valleys behind. Many projects are monitoring cation exchange capacity decrease. permafrost (for example, the European projects Histosols and Gleysols such as PermaNET – Long-term Permafrost Moni- toring Network, and PERMOS project – Swiss Per- Organic matter stored in the soil represents one mafrost Monitoring Network). If we correlate the of the greatest reservoirs of organic carbon at a global warming with increasing meteoric events, global scale and the mineralization of its part could we can suppose that the area occupied by Cryosols represent an input of CO2 in the atmosphere with will decrease while that of soils like Fluvisols in- consequences on climate change (Von Lützow et creases. With the melting of permafrost, the water al. 2006; Fontaine et al. 2007).

J. FOR. SCI., 61, 2015 (11): 496–514 501 With regard to the great importance SOM has for tannins or aliphatic compounds, humic polymers and the global balance of CO2 (Lal 2004; Schmidt et al. charred materials (Krull et al. 2003; Von Lützow 2011) the study of its potential biodegradability in et al. 2006). However, the relative contribution of each a warmer environment is currently one of the top- mechanism to the organic matter preservation in soil ics of greater interest in the scientific research (Da- is unknown (Baldock, Skjemstad 2000; Six et al. vidson, Janssens 2006; Uhlířová et al. 2007). 2002; Mikutta et al. 2006; Von Lützow et al. 2006). The soils more related with SOM are Histosols and In Alpine ecosystems SOM may be protected not Gleysols that can occur in the peat. Histosols are only by the above cited mechanisms, but also by soils with a thick organic layer, with redoximorphic the low soil temperature. The processes of SOM features in particular in that part of the profile that stabilization are temperature dependent since bi- has been in contact with water. Gleysols occur in all otic and abiotic reactions of degradation and con- wet sites also near rivers or brooks and ponds. They densation that produce new aromatic structures have an under groundwater influence where there with larger molecular weights are slowed down by are evidences of reduction processes. The layers cold temperatures. Moreover, the formation of or- have generally darker colour with greyish/bluish ganomineral associations is also slowed down, as (redox), reddish, brownish or yellowish mottles on the partial oxidation of organic molecules neces- the ped surfaces. They have a wide range of parent sary to promote adsorption to the mineral surface material: unconsolidated materials, fluvial, lacus- and/or binding with metal ions (Grünewald et al. trine sediments of the Pleistocene or Holocene age. 2006) is also temperature dependent. The severe acidity of some Gleysols and Histosols A major component of SOM of cold environ- (with the humus forms of Hydromoder or Tangel) ments is labile C with a high potential of biodegra- allows the soil memory function. The soil memory dability in a warmer environment (Dai et al. 2002; function is also shown by the Hydromull humus Mikan et al. 2002; Weintraub, Schimel 2003; form that is more common on peat (that started their Shaver et al. 2006). Moreover, the soils of cold en- formation during the Atlantic period: 8000–5000 vironments are characterized by a dense microbial years ago). They are suitable to conserve pollen, community with a high growth potential (Anesio spores, phytoliths, remnants of plants such as old et al. 2009) that is comparable, like total biomass, wood that can testify the presence of different en- with that of temperate soil ecosystems. Therefore, vironment than the present one, so they can give organic matter of alpine soils may be highly sus- evidence of global change in the past (Stepanova ceptible to the actual global warming (Schimel et et al. 2015). The peat soils have a large accumula- al. 2006), which causes permafrost melting (Chris- tion of organic matter. The European Community tensen et al. 2004; Gruber, Haeberli 2007), is very sensitive to the conservation of this habitat increase in soil temperatures (Chudinova et al. because with its deterioration (and peat mineral- 2006) and extension in the length of the growing ization) great amounts of CO2 could be released. season (Euskirchen et al. 2006). The wetland vegetation experienced a regression, Another SOM protection mechanism occurring especially at lower altitudes (Cannone et al. 2008). in the Alpine ecosystem, both in permafrost-affect- ed soils and in soils frozen for many months per Soil organic matter (SOM) and nutrient dynamics year, is the freezing effect (Davidson, Janssens The protection of SOM from degradation is mainly 2006; Uhlířová et al. 2007). In this kind of soils, due to three mechanisms: (1) physical stabilization even if the enzymatic reactions can occur below of organic matter in soil and to its inaccessibility due 0°C (Mikan et al. 2002), the diffusion of substrates to establishing physical barriers between microbes and extracellular enzymes within the soil is ex- and enzymes and their substrates (Six et al. 2004; tremely slow when the soil water is frozen. SOM of Von Lützow et al. 2006); (2) chemical stabiliza- the cold environment, buried in the deep soil layers tion, which consists in intermolecular interactions through cryoturbation, has more low-density phys- between organic substances and minerals with com- ical fractions that are potentially easily degradable plexation of functional groups of organic molecules (Zollinger et al. 2013). In these environments with metal ions that decrease the availability of the SOM has been preserved until today thanks to the organic substrate (Guggenberger, Kaiser 2003; prevailing cold conditions that have constrained its Mikutta et al. 2006); (3) biochemical stabilization, decomposition (Mikan et al. 2002). However, fol- caused by molecular structures inherently stable lowing the warmer conditions the global change is against biochemical decay such as condensed and expected to produce, the easily degradable organic lignin-derived aromatic carbons, melanoidins, some matter will lead to a transformation of the long-term

502 J. FOR. SCI., 61, 2015 (11): 496–514 cryopreserved SOM with the release of large amounts The assessment of the impacts related to global of CO2 and nutrients, and the production of more hu- change on the soil structure and its resistance to mified SOM that will enhance the vegetation growth erosion is not easy to determine, because of the (Zollinger et al. 2013). complexity of interrelated factors to consider (Park In Alpine soils there is a close link between mi- et al. 2014). The relationship has two faces. On the crobial SOM transformations and nutrient availabil- one hand, climate exercises an indirect influence on ity as for P and N elements (Weintraub, Schimel the soil structure affecting factors correlated with 2005) when in this environment these transforma- aggregation such as SOM (Von Lützow, Kögel- tions are an important limiting factor for primary Knabner 2009), soluble salts and amorphous iron production (Kaiser et al. 2005). Therefore, the in- oxides (Mikutta et al. 2006), wetting-drying and tensive SOM mineralization produced by the global freeze-thaw cycles which in turn affect land use change in the Alpine environments could bring a (Ping et al. 2008), vegetation covers and biological higher availability of N and P, which will increase cycles (Six et al. 2004). The structural improvement the assimilation of nutrients by plants but not by mi- of soil, due to the ped aggregation, will lead to a pre- crobes (Schmidt et al. 2002). dominance of humus forms in Mull (Chersich et The greatest part of the knowledge dealing with the al. 2006). Their effects on the soil structure can be effects of global change on SOM and nutrient dynam- positive or negative and structural characteristics ics is due to laboratory incubation researches (Frep- are the dynamic result of their complex interaction. paz et al. 2014). The great importance of snow pre- On the other hand, climate change affects the soil cipitation on nutrient dynamics and also the effects structure directly through the increase of extreme of snow thickness on microbial community and SOM precipitation events and snow-melt, which could dynamics are well known facts (Hiltbrunner et al. raise the frequency and severity of floods (e.g. the 2005; Tan et al. 2014). For these reasons, it should be flood risks are more frequent than in the past;Al - necessary to consider all the factors which could in- lamano et al. 2009) and runoff. The types of soils fluence the climate of Alpine environments. will see a decrease in the area of Cryosols (and probably of Regosols) in favour of an increase in Erosion processes Leptosols and Fluvisols. Bouwer (2011) stated that the global soil loss- es by erosion and landslides have escalated enor- mously over the past few decades. Consequently, The lower Alpine horizon: the Alpine soil erosion under the global climate tree line shifting and podzolization shifting changes becomes one of the most important topics of European research. The upper altitudinal limit of trees is known as According to Christensen and Christensen the tree line, although it is not usually a distinct line (2003) and others (i.e. Beniston 2003), reductions but a steep gradient of increasing stand fragmenta- in average summer precipitation and consequent tion and stuntedness (Körner, Paulsen 2004). The episodes of summer drought may be simultane- tree line has been directly affected by the exploita- ously accompanied by a sharp increase in short but tion of mountain forests which was partly followed potentially-devastating heavy precipitation events either by practices that favoured certain species (Casty et al. 2005; Beniston et al. 2011), such (Pitkänen et al. 2003) or by spontaneous second- as flash floods, more frequent overflow of storm ary successions (Bradshaw 2004). This important drainage facilities, greater soil erosion and higher boundary (Forrest et al. 2012) is observed in all risk of landslides in unstable areas (Bollschweil- parts of the world, and it exhibits common fea- er, Stoffel 2010). Where the climate change is tures (Holtmeier, Broll 2005). Currently there expressed by extreme events, the erosion will be is much interest in the rate at which the tree line stronger, and the extent of Leptosols might be larg- may advance in response to environmental change, er. Leptosols are azonal soils, weakly developed di- especially global warming (Smith et al. 2009). The rectly on the rock (within 25 cm from the soil sur- reason for renewed interest in the tree line is clear: face), or where the erosion has removed the upper an advancing tree line, or a denser forest below the part of the soil profile. Leptosols are shallow, often tree line, would have significant implications for the extremely gravelly soils, so they have severe limita- global carbon cycle (increasing the terrestrial carbon tion to rooting. The forms of humus more frequent sink) and for the biodiversity of the Alpine ecotone also in this case will be those of Mull: Mesomull, as ousting rare species and disrupting plant commu- Dysmull (Chersich 2007). nities (Walther et al. 2005; Berthel et al. 2012;

J. FOR. SCI., 61, 2015 (11): 496–514 503 Prietzel, Christophel, 2014; Vilmundardót- by Körner et al. (2005) was based on the efflux tir et al. 2015). Presently it is really difficult to of pure CO2 into tree canopies through a fine web study such a role of forests in detail; Parviainen et of woven tubes (web-FACE, Free Air CO2 Enrich- al. (2000) stated that natural forests represent only ment) at a concentration of 530 ppm over 4 years. 1.7 % of the total European forest area. Neverthe- The data suggested no lasting growth stimulations less, the transfer of knowledge from virgin forests by CO2 enrichment in mature trees after 4 years, (Brang 2005) allows the frame characterisation of while young trees showed continuous growth stim- ecological conditions caused by an advancing tree ulation. The uptake of carbon was not reduced and line. Therefore, new data on tree-growth dynamics no more carbon was trapped but it seemed that and species adaptation near the treeline (Wieser et the plants pump more carbon through their bodies al. 2014) under global climate changes (Piermattei (Hoffmann et al. 2014). Another possible limiting et al. 2014) can be combined with changes in the dy- factor could be the supply of nitrogen (Guidi et al. namics of forest soil processes. 2014). From the global warming point of view, a ris- There are at least three aspects of environmental ing tree line will be reflected in increasing biomass change to which plants are generally thought to re- (from 15 to 75 t C·ha–1), which would produce a sponse: increasing temperature, rising concentra- warmer microclimate and better decomposabil- tion of carbon dioxide and increasing deposition of ity of produced litter, increasing root growth and nitrogen (Stanhill, Cohen 2001). Several studies root respiration (Pregitzer et al. 2000) and, at the carried out in the Alpine environment (e.g. David end of the process, raising the turnover of carbon 1993 in the French Alps) have demonstrated that compensated by a greater carbon input (Ni et al. during the Boreal period (5000–9000 years BP) the 2015). The global carbon cycle would remain unaf- tree line was placed higher than it is at present. fected and the soil organic matter turnover would Plants may well respond to warming as this will increase (Kitz et al. 2015). increase the rate of cell division and therefore the The timber line is also the boundary for the pod- rate at which the assimilated carbon can be utilized zolization domain because this process requires the (Zhu et al. 2015). Warming will also prolong the presence of coniferous vegetation. The high precipi- growing season and, in the already short growing tation rate in Alpine areas results in a lot of water season at the tree line, this effect will be propor- entering in the soil. Related to the timber line, its tionately greater than in lowland or more south- natural shifting (supported by an anthropogenic ern conditions. In the European Alps, during the pressure) plays a great role in the water balance of last century, the total annual mean temperatures the site. The downward moving of water through the increased by about 2°C (Auer et al. 2007), which soil profile can dissolve large nutrient cations such as is above the observed global increase in tempera- potassium and calcium and stimulate the migration ture by 0.74°C over the last century (Solomon et of organic matter with Al, Fe complexes. The cation al. 2007). Since Alpine trees are climatically de- exchange capacity is dominated by hydrogen ions H+ termined ecotones and they are particularly sen- and Al3+. The more frequent soils under coniferous sible to altered temperature regimes (Theurillat, forests are Podzols developed on aeolian siliceous Guisan 2001), the predicted climate warming is material including loess, and the humus is mainly expected to structurally change the tree composi- an acid one, like Mor or Moder forms (Chersich tion and to raise the tree line. In the Swiss Alps, 2007). The altitudinal rising of podzolization is an more recent results have confirmed these find- evidence of the global change recorded in the soil. ings and reported a strong warming since the The past of the climatic change has already been re- 1980s (Rebetez, Reinhard 2008). Walther et al. corded. In the Alps it is common to find Podzols over (2005) found that the vegetation change in the the timber line. The spodic horizon, which could be south-eastern Swiss Alps has accelerated since considered the “soil memory” (Targulian, Gory- 1985, which was consistent with the increased tem- achkin 2011), points out a warmer climate than the perature regime observed at the same sites. Data present one which allowed the podzolization (going collected from the South Ural region (Kapralov back to 130000–75000 years ago when there was the et al. 2006) showed that in the last 50 years the last interglacial period). In the same way, if the cli- upper boundary of low forest has ascended, with mate is warmer than now, we will find the timberline considerable changes in the composition, density and podzolization at a higher altitude and the hu- and height of the tree layer. The response of forest mus form will be more acid (Pengerud et al. 2014): trees to increasing atmospheric carbon dioxide is from the Mull or Moder of the Alpine grass we will still an open question. The experiment carried out observe the Mor form under coniferous forest.

504 J. FOR. SCI., 61, 2015 (11): 496–514 On the south-facing slopes of the Alps we can find of highly productive soils (loss of organic matter Dystric Cambisols or Podzols over 1,600 m a.s.l. un- in soils, degradation of the ecological functions of der coniferous trees. We find Cambisols in regions soils) and the partial or complete destruction of that were under the influence of glaciations during crops planted in steep terrains. An example of the the Pleistocene. In the brunification, i.e. in the main consequences of landslide development for crops processes for Cambisol formation, it is evident the in steep terrains of the Alpine region is the de- polyhedral subangular soil structure and Bw hori- struction of terraced slopes with vineyards and the zons have mostly brownish coloration with an in- subsequent economic losses for the intense rainfall crease of clay percentage and/or carbonate removal. event of 14–17 November 2000 in the Valtellina Changes in the vegetation cover and the extent of area (Central Alps, Italy) (Crosta et al. 2003). cryosphere can be recorded into the soil profile. Moreover, the increase in landslides and flood Therefore in the Alpine environment we can distin- susceptibility could be responsible for higher risks guish soil processes linked with the altitudinal belt for the human structures, like blockage and de- (e.g. podzolization, brunification, cryoturbation) struction of roads and railways with subsequent and others referred to the particular zone (gleyfica- isolation of towns and buildings and, as a conse- tion and stratification near a water source). quence, for the people living in hazardous areas Below the tree line, due to the effect of tempera- (Einhorn et al. 2015). Thus, better assessment of ture intolerance, the alpine species cannot with- the soil and environmental conditions will become stand the intensity of the competition. Above the fundamental that lead to triggering landslides and tree line, an upward migration of the species on floods in relation to the forecasted climate change mountain summits has been observed with upward effects, for reducing the hazard and the risk linked shrub displacement and an increase of species rich- to these types of phenomena. ness, which may be a transient effect of the habitat fragmentation or a long-term effect of the vegeta- tion succession (Cannone, Pignatti 2014). Comparison of approaches and conclusions

During the 21st century, the most recent climate Impact of soil processes induced by climatic modelling, based on a range of scenarios, shows an changes on socio-economic sphere increase in annual temperature by 0.1–0.4°C per decade (Alcamo et al. 2007) when a rise in pre- In the Alpine region, an increase in the frequency cipitation in the north, its decrease in the south, of intense rainfalls (> 30 mm/day) and permafrost an increase in the seasonality of precipitation and degradation linked to the climatic changes (Ein- prolonged dry periods are expected. Thus the global horn et al. 2015) could determine a change in the warming would bring warmer and more humid con- water regime of soils, promoting an increase of the ditions in winter and much warmer and drier condi- soil moisture and causing the persistence of satu- tions in summer. This would be connected with an rated or close to saturation conditions all the year increase in the frequency of intense precipitation round. These conditions are responsible for trigger- events like rainfalls. Many uncertainties arising from ing erosion processes, debris flows and landslides the ongoing environmental and socio-economic (Collison et al. 2000; Lu, Gsodt 2013) and they processes (Kauppi 1996) have likely been expected. can represent preparatory factors for flood trigger- The aim of this study was to review the papers con- ing (jomelli et al. 2009). Climatic modifications cerning the effects of global change on the Alpine also cause a change in land cover due to different ecosystem, which is very sensitive to climatic chang- rainfall and temperature conditions. In particular, es, therefore an optimal marker to record them. In the shifting of the tree line to a higher altitude can particular, we observed the global change in pedo- cause the development of bare soils or sphere in relation with cryosphere and biosphere. In and shrub lands, which are more susceptible to soil particular, effects on the flora through distributional erosion, shallow landslides and debris flows which changes of species across climate scenarios (Thuil- develop in mountain regions (Begueria 2006). ler 2004) seem to be very presumable. The increases in landslides and flood susceptibil- As for cryosphere, the immediate consequences are ity related to climate change have negative effects the loss of about half of the European Alpine glacier on the landscapes and on the people (Crozier et mass during the past 150 years and a decrease in the al. 2010). Landslides and floods affect the surface snow cover extent. In recent model studies, the snow soils developed on hillslopes, provoking the loss at low to medium elevations will disappear by 2100

J. FOR. SCI., 61, 2015 (11): 496–514 505 (Beniston 2012), a more pronounced decrease in ward shifts and the species of the neighbouring snowfall volumes of the Italian Alps is projected communities filled the range of the same altitu- around 2060 (Soncini, Bocchiola 2011). These dinal belt (Cannone, Pignatti 2014). Concern- evidences are related with the pedosphere. The ing the related soil variability (Rajkai 2008), the particular knowledge of interrelationships be- Cryosol (and Regosol) area will be reduced and tween the soil type formation and the dominat- these soils will be shifted to higher altitudes like ing phytocoenological society related to climate the Podzol area, while the Fluvisol and Leptosol development was described by Pichler et al. area might become larger. It is also fundamental (2011). Concerning the physical conditions of soil, to underline how it is difficult to predict a future the increasing thickness of the active layer can de- scenario because all the different environmental termine the destabilisation of slopes and, together components have a peculiar response to climate with an increase of intensive rainfall events, cause change with a specific reaction time (Soncini, severe water erosion phenomena, or even debris Bocchiola 2011). We recognize the important flows and landslide episodes. It will produce a role of the time as a factor of pedogenesis, “times change in soil structural features, which is also in of changes” but at the moment there are no suf- equilibrium with several factors influenced by cli- ficient bibliographic data to give a reaction time mate change. One of the most important factors for each environmental variable and we can only is SOM. With the increasing soil temperature, the formulate a hypothesis. soil organic matter may not be protected anymore As regards the effects on the socio-economic (e.g. permafrost melting), with the subsequent sphere, a change in the rainfall pattern of the Al- release of large amounts of CO2 and nutrients, pine region could provoke an increase in the fre- producing therefore more humified SOM. The in- quency of extreme rainfall events, which could tense SOM mineralization produced by the global allow for triggering a higher number of erosion change might bring a higher availability of N and processes, landslides and floods. These phenom- P, which, in turn, will lead to greater primary pro- ena could have negative impacts on different as- duction with the subsequent CO2 immobilization pects of the socio-economic sphere of the people in the vegetal structure. However, this fact does living in Alpine regions: (i) partial or complete not seem to be able to balance CO2 emitted from destruction of the crops planted in steep terrains SOM mineralization (Davidson, Janssens 2006). with the subsequent loss of highly productive In biosphere the global warming will cause an soils and severe economic damage; (ii) signifi- upward migration of alpine plants (Parmesan cant damage to human structures and buildings; 2006; Grabherr et al. 2010). In response to the (iii) increase in the number of people living in environmental change, the tree line may advance highly vulnerable areas. upwards. The podzolization line, which is related As highlighted by this review, it is fundamental to acidification by the coniferous tree litter, may to recognize the implications of climate change for also advance to a higher altitude. The warming the Alps, because it will have consequences not will prolong the growing season with subsequent only for the environment but also for the socio- changes in the phenology – e.g. influence on the economic sphere. Therefore, the identification of start of the vegetation period for plant species. An global change evidences is the starting point to increase in the rate of the cell division and there- raise human awareness and to find an appropriate fore in the rate at which the assimilated carbon mitigation strategy. can be utilized are expectable. The snowbeds and species will be reduced, especially con- cerning endemic ones, due to pioneer and early Acknowledgments successional species while the transect effect of an increase in species richness has been observed We thank Prof. F. Previtali (former University that may confirm the forecast of species loss and of Milano, Bicocca), Dr. M. Maggioni (University habitat fragmentation or a longer effect (Can- of Torino, NatRisk Department of Agricultural, none, Pignatti 2014). In this scenario the biodi- Forest and Food Sciences), Dr. F. Favilli (EURAC versity is endangered – based on the relationship Research – Institute for Regional Development and between the productivity and diversity of phyto- Location Management, Bolzano), Dr. Clorino coenoses (Pretzsch 2005). Cioci, Dr. Marco Basili and Dr. Elisa Oberto Regarding the species distribution, it has been for the willingness to discuss and make sugges- observed that the species mainly exhibited down- tions to the article.

506 J. FOR. SCI., 61, 2015 (11): 496–514 References Beniston M. (2001): Impacts of climate change on mountain regions. In: Smith J. (ed.): The Rise of Modern Genomics. Alcamo J., Moreno J.M., Nováky B., Bindi M., Corobov R., New York, Wiley: 193–206. Devoy R.J.N., Giannakopoulos C., Martin E., Olesen J.E., Beniston M. (2003): Climatic change in mountain regions: Shvidenko A. (2007): Chapter 12: Europe. In: Parry M.L., a review of possible impacts. Climatic Change, 59: 5–31. Canziani O.F., Palutikof J.P., van der Linden P.J., Hanson Beniston M. (2005): Mountain climates and climatic change: C.E. (eds): Climate Change 2007: Impacts, Adaptation an overview of processes focusing on the European Alps. and Vulnerability. Contribution of Working Group to the Pure and Applied Geophysics, 162: 1587–1606. 4th Assessment Report of the Intergovernmental Panel Beniston M. (2006): Mountain weather and climate: a general on Climate Change. Cambridge, Cambridge University overview and a focus on climatic change in the Alps. Hyd- Press: 541–580. robiologia, 562: 3–16. Allamano P., Claps P., Laio F. (2009): Global warming in- Beniston M., Stoffel M., Hill M. (2011): Impacts of climatic creases flood risk in mountainous areas. Geophysical change on water and natural hazards in the Alps: can current Research Letters, 36: L24404. water governance cope with future challenges? Examples Anderson R.S., Jiménez-Moreno G., Carrión J.S., Pérez- from the European “ACQWA” project. Environmental Sci- Martínez C. (2011): Postglacial history of alpine vegeta- ence and Policy, 14: 734–743. tion, fire, and climate from Laguna de Río Seco, Sierra Beniston M. (2012): Is snow in the Alps receding or disappear- Nevada, southern . Quaternary Science Reviews, ing? Climatic Change, 3: 349–358. 30: 1615–1629. Beniston M., Stoffel M. (2014): Assessing the impacts of cli- Anesio A.M., Hodson A.J., Fritz A., Psenner R., Sattler B. matic change on mountain water resources. Science of the (2009): High microbial activity on glaciers: importance Total Environment, 493: 1129–1137. to the global carbon cycle. Global Change Biology, 15: Berthel N., Schwörer C., Tinner W. (2012): Impact of Holocene 955–960. climate changes on alpine and treeline vegetation at Sanetsch Arkhangelskaya T.A. (2014): Diversity of thermal conditions Pass, Bernese Alps, Switzerland. Review of Palaeobotany within the paleocryogenic soil complexes of the East Euro- and Palynology, 174: 91–100. pean Plain: The discussion of key factors and mathematical Bocchiola D., Diolaiuti G. (2010): Evidence of climate change modelling. Geoderma, 213: 608–616. within the Adamello Glacier of Italy. Theoretical and Applied Auer I., Böhm R., Jurkovic A., Lipa W., Orlik, A., Potzmann Climatology, 100: 351–369. R., Schöner W., Ungersböck M., Matulla C., Briffa K., Jones Boeckli L., Brenning A., Gruber S., Noetzli J. (2012): A statis- P., Efthymiadis D., Brunetti M., Nanni T., Maugeri M., tical approach to modelling permafrost distribution in the Mercalli L., Mestre O., Moisselin J.M., Begert M., Müller- European Alps or similar mountain ranges. Cryosphere, 6: Westermeier G., Kveton V., Bochnicek O., Stastny P., Lapin 125–140. M., Szalai S., Szentimrey T., Cegnar T., Dolinar M., Gajic- Bollschweiler M., Stoffel M. (2010): Changes and trends in Capka M., Zaninovic K., Majstorovic Z., Nieplova E. (2007): debris-flow frequency since AD 1850: Results from the Swiss HISTALP – historical instrumental climatological surface Alps. Holocene, 20: 907–916. time series of the Greater Alpine Region. International Bouwer L.M. (2011): Have disaster losses increased due to Journal of Climatology, 27: 17–46. anthropogenic climate change? American Meteorological Baldock J.A., Skjemstad J.O. (2000): Role of the soil matrix Society, 92: 39–46. and minerals in protecting natural organic materials against Bowman W.D., Murgel J., Blett T., Porter E. (2012): Nitrogen biological attack. Organic Geochemistry, 31: 697–710. critical loads for alpine vegetation and soils in Rocky Moun- Barthès B., Roose E. (2002): Aggregate stability as an indica- tain National Park. Journal of Environmental Management, tor of soil susceptibility of runoff and erosion; validation 103: 165–171. at several levels. Catena, 47: 133–149. Bradshaw C.J.A., Warkentin I.G. (2015): Global estimates of Baumann F., Schmidt K., Dörfer C., He J.S., Scholten T., boreal forest carbon stocks and flux. Global and Planetary Kühn P. (2014): Pedogenesis, permafrost, substrate and Change, 128: 24–30. topography: plot and landscape scale interrelations of Bradley J.A., Singarayer J.S., Anesio A.M. (2015): Microbial weathering processes on the central-eastern Tibetan Pla- community dynamics in the forefield of glaciers. Proceedings teau. Geoderma, 226–227: 300–316. of the Royal Society B: Biological Sciences, 281: 20140882 Begueria S. (2006): Changes in land cover and shallow Bradshaw R.H.W. (2004): Past anthropogenic influence on landslide activity: a case study in the Spanish . genetic structure and diversity within European forests. Geomorphology, 74: 1–4, 196–206. Forest Ecology and Management, 197: 203–212. Beniston M. (1997): Variations of snow depth and duration Brang P. (2005): Virgin forests as a knowledge source for Cen- in the Swiss Alps over the last 50 years: links to changes in tral European silviculture: reality or myth? Snow Landscape large-scale climatic forcing. Climatic Change, 36: 281–300. Research, 79: 19–31.

J. FOR. SCI., 61, 2015 (11): 496–514 507 Brunetti M., Maugeri M., Nanni T., Auer I., Böhm R., Schöner glaciers during the last century: a contribution to knowl- W. (2006): Precipitation variability and changes in the edge about Alpine glacier changes. Geografiska Annaler, greater Alpine region over the 1800–2003 period. Journal 89 A3: 167–184. of Geophysical Research, 111: D11107. Clement J.C., Robson T.M., Guillemin R., Saccone P., Lochet Burga C.A., Krüsi B., Egli M., Wernli M., Elsener S., Zieflec J., Aubert S., Lavorel S. (2012): The effects of snow-N M., Fischer T., Mavris C. (2010): Plant succession and soil deposition and snowmelt dynamics on soil-N cycling in development on the foreland of the Morteratsch glacier marginal terraced grasslands in the French Alps. Biogeo- (Pontresina, Switzerland): Straight forward or chaotic? chemistry, 108: 297–315. Flora, 205: 561–576. Collison A., Wade S., Griffiths J., Dehn M. (2000): Model- Cannone N., Diolaiuti G., Guglielmin M., Smiraglia C. (2008): ling the impact of predicted climate change on landslide Accelerating climate change impacts on Alpine glacier frequency and magnitude in SE England. Engineering forefield ecosystems in the European Alps. Ecological Ap- Geology, 55: 205–218. plication, 18: 637–648. Confortola G., Soncini A., Bocchiola D. (2013): Climate Cannone N., Pignatti S. (2014): Ecological responses of plant change will affect hydrological regimes in the Alps, A case species and communities to climate warming: upward shift study in Italy. Revue De Geographie Alpine/Journal of or range filling processes? Climatic Change, 123: 201–214. Alpine Research, 101–103. Available at http://rga.revues. Casty C., Wanner H., Luterbacher J., Esper J., Böhm R. (2005): org/2176 (accessed April 22, 2015). Temperature and precipitation variability in the European Cossart E., Fort M., Bourles D., Carcaillet J., Perrier R., Alps since 1500. International Journal of Climatology, 25: Siame L., Braucher R. (2010): Climatic significance of 1855–1880. glacier retreat and rockglaciers re-assessed in the light of Chersich S., Ivetic B., D’Alessio D. (2006): Studio preliminare cosmogenic dating and weathering rind thickness in Clarée della variabilità delle forme di humus studiate secondo valley (Briançonnais, French Alps). Catena, 80: 204–219. due diversi approcci in relazione al tipo di vegetazione Crosta G.B., Dal Negro P., Frattini P. (2003): Soil slips and presente in stazioni campione di aree montane lombarde. debris flows on terraced slopes. Natural Hazards and Earth Available at http://www.sisef.it/forest@/contents/?id=efor System Sciences, 3: 31–42. 0412-0030562 (accessed March 26, 2015). Crozier M.J. (2010): Deciphering the effect of climate change Chersich S. (2007): Approccio morfologico-funzionale nello on landslide activity: A review. Geomorphology, 124: studio dei suoli: relazioni tra gli epipedon umiferi e gli 260–267. orizzonti organo-minerali in alcuni profili pedologici in D’Agata C., Bocchiola D., Maragno D., Smiraglia C., Diolaiuti ambiente alpino. Available at http://www.sisef.it/forest@/ A. (2013): Glacier shrinkage driven by climate change in the pdf/?id=efor0474-0040333 (accessed March 26, 2015). Ortles-Cevedale group (Stelvio National Park, Lombardy, Chimani B., Matulla C., Böhm R., Hofstätter M. (2013): A Italian Alps) during half a century (1954–2007). Theoretical new high resolution absolute temperature grid for the and Applied Climatology, 116: 169–190. Greater Alpine Region back to 1780. International Journal Dai X.Y., Ping C.L., Michaelson G.J. (2002): Characterizing of Climatology, 33: 2129–2141. soil organic matter in arctic soils by different ana- Chiri E., Nauer P.A., Henneberger R., Zeyer J., Schroth lytical approaches. Organic Geochemistry, 33: 407–419. M.H. (2015): Soil-methane sink increases with soil age in D’Amico M.E., Freppaz M., Filippa G., Zanini E. (2014): forefields of Alpine glaciers. Soil Biology and Biochemistry, Vegetation influence on soil formation rate in a proglacial 84: 83–95. chronosequence (Lys Glacier, NW Italian Alps). Catena, Christensen J.H., Christensen O.B. (2003): Climatemodel- 113: 122–137. ling: severe summertime flooding in Europe. Nature, 421: David F. (1993): Evolutions de la limite supérieure des arbres 805–806. dans les Alpes françaises du nord depuis la fin des temps Christensen T.R., Johansson T.R., Akerman H.J., Mastepanov glaciaires. [Ph.D. Thesis.] Marseille, University of Aix- M., Malmer N., Friborg T., Crill P., Svensson B.H. (2004): Marseille III: 94. Thawing subarctic permafrost: effects on vegetation and Davidson E.A., Janssens I.A. (2006): Temperature sensitiv- methane emissions. Geophysical Research Letters, 31: ity of decomposition and feedbacks to climate L04501. change. Nature, 440: 165–173. Chudinova S.M., Frauenfeld O.W., Barry R.G., Zhang T.J., Davies M.C.R., Hamza O., Harris C. (2001): The effect of rise Sorokovikov V.A. (2006): Relationship between air and in mean annual temperature on the stability of rock slopes soil temperature trends and periodicities in the permafrost containing ice-filled discontinuities. Permafrost Periglacial regions of Russia. Journal of Geophysical Research: Earth Process, 12: 137–144. Surface, 111: 1–15. Déry S.J., Brown R.D. (2007): Recent Northern Hemisphere Citterio M., Diolaiuti G., Smiraglia C., D’Agata C., Carnielli snow cover extent trends and implications for the snow- T., Stella G., Siletto G.B. (2007): The fluctuations of Italian albedo feedback. Geophysical Research Letters, 34: L22504

508 J. FOR. SCI., 61, 2015 (11): 496–514 Díaz-Varela R.A., Colombo R., Meroni M., Calvo-Iglesias change: Assessing the vulnerability of habitat M.S., Buffoni A., Tagliaferri A. (2010): Spatio-temporal to treeline shift in the Himalaya. Biological Conservation, analysis of alpine ecotones: A spatial explicit model tar- 150: 129–135. geting altitudinal vegetation shifts. Ecological Modeling, Fouché J., Keller C., Allard M., Ambrosi J.P. (2014): Increased 221: 621–633. CO2 fluxes under warming tests and soil solution chemistry Diolaiuti G., Bocchiola D., D’agata C., Smiraglia C. (2012): in Histic and Turbic Cryosols, Salluit, Nunavik, Canada. Soil Evidence of climate change impact upon glaciers’ reces- Biology and Biochemistry, 68: 185–199. sion within the Italian Alps: the case of Lombardy glaciers. Frauenfelder R., Laustela M., Kääb A. (2005): Relative age Theoretical and Applied Climatology, 109: 429–445. dating of Alpine rock glacier surfaces. Zeitschrift für Geo- Edwards A.C., Cresser M.S. (1992): Freezing and its effect morphologie, NF49: 145–166. on chemical and biological properties of soil. Advance in French H.M. (1976): The Periglacial Environment. London, Soil Science, 18: 59–95. New York, Longman Group Limited: 309. EEA (2009): Regional climate change and adaptation. The Freppaz M., Said-Pullicino D., Filippa G., Curtaz F., Celi L., Alps facing the challenge of changing water resources. Zanini E. (2014): Winterespring transition induces changes EEA, Copenhagen. Available at http://www.eea.europa. in nutrients and microbial biomass in mid-alpine forest soils. eu/publications/alps-climate-change-and-adaptation-2009 Soil Biology and Biochemistry, 78: 54–57. (accessed April 22, 2015). Gentili R., Baroni C., Caccianiga M., Armiraglio S., Ghiani Egli M., Dahms D., Norton K. (2014): Soil formation rates on A., Citterio S. (2015): Potential warm-stage microrefugia silicate parent material in alpine environments: approach- for alpine plants: Feedback between geomorphological es-different results? Geoderma, 213: 320–333. and biological processes. Ecological Complexity, 21: 87–99. Einhorn B., Eckert N., Chaix C., Ravanel L., Deline P., Gardent Gerlach R., Baumewerd-Schmidt H., van den Borg K., Eckmei- M., Boudières V., Richard D., Vengeon J.M., Giraud G., er E., Schmidt M.W.I. (2006): Prehistoric alteration of soil in Schoeneich P. (2015): Climate change and natural hazards the Lower Rhine Basin, Northwest Germany – archaeologi- in the Alps. Journal of Alpine Research, 103: 2–31. cal, 14C and geochemical evidence. Geoderma, 136: 38–50. Euskirchen E.S., McGuire A.D., Kicklighter D.W., Zhuang Q., Gerrard J. (1992): Soils Geomorphology. New York, Chapman Clein J.S., Dargaville R.J., Dye D.G., Kimball J.S., McDonald and Hall: 269. K.C., Melillo J.M., Romanovsky V.E., Smith N.V. (2006): Gobiet A., Kotlarski S., Beniston M., Heinrich G., Rajczak J., Importance of recent shifts in soil thermal dynamics on Stoffel M. (2014): 21st century climate change in the Euro- growing season length, productivity, and carbon sequestra- pean Alps – a review. Science of the Total Environment, tion in terrestrial high-latitude ecosystems. Global Change 493: 1138–1151. Biology, 12: 731–750. Goudiet A.S. (2006): The Schmidt Hammer in geomorphologi- Evershed R.P. (2008): Organic residue analysis in archaeology: cal research. Progress in Physical Geography, 30: 703–718. The archaeological biomarker revolution. Archaeometry, Grabherr G., Gottfried M., Pauli H. (2010): Climate change 50: 895–924. impacts in Alpine environments. Geography Compass, 4: Favilli F., Egli M., Cherubini P., Sartori G., Haeberli W., Delbos 1133–1153. E. (2008): Comparison of different methods of obtaining a Groffman P.M., Driscoll C.T., Fahey T.J., Hardy J.P., Fitzhugh resilient organic matter fraction in Alpine soils. Geoderma, R.D., Tierney G.L. (2001): Colder soils in a warmer world: 145: 355–369. A snow manipulation study in a northern hardwood forest Favilli F., Egli M., Brandova D., Ivy-Ochs S., Kubik P., Cheru- ecosystem. Biogeochemistry, 56: 135–150. bini P., Mirabella A., Sartori G., Giaccai D., Haeberli W. Gruber S., Haeberli W. (2007): Permafrost in steep bedrock (2009): Combined use of relative and absolute dating slopes and its temperature-related destabilization follow- techniques for detecting signals of Alpine landscape ing climate change. Journal of Geophysical Research, 112: evolution during the late Pleistocene and early Holocene. F02–S18. Geomorphology, 112: 48–66. Grünewald G., Kaiser K., Jahn R., Guggenberger G. (2006): Fischer L., Huggel C., Kääb A., Haeberli W. (2013): Slope Organic matter stabilization in young calcareous soils as failures and erosion rates on a glacierized high-mountain revealed by density fractionation and analysis of lignin- face under climatic changes. Earth Surf Process and Land- derived constituents. Organic Geochemistry, 37: 1573–1589. forms, 38: 836–846. Gu L., Yao J., Hu Z., Zhao L. (2015): Comparison of the surface Fontaine S., Barot S., Barré P., Bdioui N., Mary B., Rumpel energy budget between regions of seasonally frozen ground C. (2007): Stability of organic carbon in deep soil layers and permafrost on the . Atmosferic Research, controlled by fresh carbon supply. Nature, 450: 277–280. 153: 553–564. Forrest J.L., Wikramanayake E., Shrestha R., Areendran G., Guggenberger G., Kaiser K. (2003): Dissolved organic matter Gyeltshen K., Maheshwari A., Mazumdar S., Naidoo R., in soils. Challenging the paradigm of sorptive preservation. Thapa G.J., Thapa K. (2012): Conservation and climate Geoderma, 113: 293–310.

J. FOR. SCI., 61, 2015 (11): 496–514 509 Guidi C., Vesterdal L., Gianelle D., Rodeghiero M. (2014): distribution of low forest at the upper limit of their growth Changes in soil organic carbon and nitrogen following in the Northern Ural Mountains. Russian Journal of Ecol- forest expansion on grassland in the Southern Alps. Forest ogy, 37: 367–372. Ecology and Management, 328: 103–116. Kauppi P.E. (1996): What is changing in the global environ- Haeberli W., Hoelzle M., Frank P., Zemp M. (2007): Integrated ment? In: Korpilahti E., Mikkela H., Salonen T. (eds): Caring monitoring of mountain glaciers as key indicators of global for the Forest: Research in a changing world. In: IUFRO XX climate change: the European Alps. Annals of Glaciology, World Congress, Congress Report 11, Tampere, Finland. 46: 150–160. August 6–12, 1996: 29–34. Harris C., Arenson L.U., Christiansen H.H., Etzelmüller B., Kirkpatrick J.B., Green K., Bridle K.L., Venn S.E. (2014): Frauenfelder R., Gruber S., Haeberli W., Hauck C., Hölzle Patterns of variation in Australian alpine soils and their M., Humlum O., Isaksen K., Kääb A., Kern-Lütschg M.A., relationships to parent material, vegetation formation, Lehning M., Matsuoka N., Murton J.B., Nötzli J., Phillips M., climate and topography. Catena, 121: 186–194. Ross N., Seppälä M., Springman S.M., Mühll D.V. (2010): Kitz F., Steinwandter M., Traugott M., Seeber J. (2015): In- Permafrost and climate in Europe: Monitoring and model- creased decomposer diversity accelerates and potentially ling thermal, geomorphological and geotechnical responses. stabilises litter decomposition. Soil Biology and Biochem- Earth Science Review, 92: 117–171. istry, 83: 138–141. Helbing D. (2013): Globally networked risks and how to re- Kohler T., Maselli D. (2009): Mountains and Climate Change spond. Nature, 497: 51–59. – From Understanding to Action. Bern, Geographica Bern- Hiltbrunner E., Schwikowski M., Körner C. (2005): Inorganic ensia with the support of the Swiss Agency for Develop- nitrogen storage in alpine snow pack in Central Alps (Swit- ment and Cooperation (SDC), and an international team of zerland). Atmospheric Environment, 39: 2249–2259. contributors. Available at http://www.fao.org/docrep/017/ Hjulström B., Isaksson S. (2009): Identification of activity area i2869e/i2869e00.pdf (accessed April 24, 2015). signatures in a reconstructed Iron Age house by combining Körner C., Paulsen J. (2004): A world-wide study of high element and lipid analyses of sediments. Journal of Archaeol- altitude treeline temperatures. Journal of Biogeography, ogy Science, 36: 174–183. 31: 713–732. Hoffmann U., Hoffmann T., Johnson E.A., Kuhn N.J. (2014): Körner C., Asshoff R., Bignucolo O., Hättenschwiler S., Keel Assessment of variability and uncertainty of soil organic S.G., Peláez-Riedl S., Pepin S., Siegwolf R.T.W., Zotz G. carbon in a mountainous boreal forest (Canadian Rocky (2005): Carbon flux and growth in mature deciduous forest

Mountains, Alberta). Catena, 113: 107–121. trees exposed to elevated CO2. Science, 309: 1360–1362. Holtmeier F.K., Broll G. (2005): Sensitivity and response of Kramer A., Herzschuh U., Mischke S., Zhang C. (2010): northern hemisphere altitudinal and polar treelines to en- Holocene treeline shifts and monsoon variability in the vironmental change at landscape and local scales. Global Hengduan Mountains (southeastern Tibetan Plateau), Ecology and Biogeography, 14: 395–410. implications from palynological investigations. Palaeo- Hormes A., Ivy-Ochs S., Kubik P.W., Ferreli L., Michetti A.M. geography Palaeoclimatology Palaeoecology, 286: 23–41. (2008): 10Be exposure ages of a rock avalanche and a late Krull E.S., Baldock J.A., Skjemstad J.O. (2003): Importance glacial moraine in Alta Valtellina, Italian Alps. Quaternary of mechanisms and processes of the stabilization of soil International, 190: 136–145. organic matter for modelling carbon turnover. Functional IUSS Working Group WRB (2006): World Reference Base for Plant Biology, 30: 207–222. Soil Resources 2006. World Soil Resources Reports No 103. Kurylyk B.L., MacQuarrie K.T.B., McKenzie J.M. (2014): Rome, FAO: 145. Climate change impacts on groundwater and soil tem- Jomelli V., Brunstein D., Déqué M., Vrac M., Grancher D. peratures in cold and temperate regions: Implications, (2009): Impacts of future climatic change (2070–2099) on the mathematical theory, and emerging simulation tools. Earth potential occurrence of debris flows: a case study in the Mas- Science Review, 138: 313–334. sif des Ecrins (French Alps). Climatic Change, 97: 171–191. Lal R. (2004): Soil carbon sequestration to mitigate climate Kääb A., Frauenfelder R., Roer I. (2007): On the response of change. Geoderma, 123: 1–22. rock glacier creep to surface temperature increase. Climate Laternser M., Schneebeli M. (2003): Long-term snow climate change impacts on mountain glaciers and permafrost. Global trends of the Swiss Alps (1931–99). International Journal and Planetary Change, 56: 172–187. of Climatology, 23: 733–750. Kaiser C., Meyer H., Biasi C., Rusalimova O., Barsukov P., Lavee H., Sarah T., Imeson A.C. (1996): Aggregate stability Richter A. (2005): Storage and mineralization of carbon and dynamics as affected by soil temperature and moisture nitrogen in soils of a frost-boil tundra ecosystem in Siberia. regimes. Geographical Analysis, 78: 73–82. Applied Soil Ecology, 29: 173–183. Liancourt P., Spence L.A., Song D.S., Lkhagva A., Sharkhuu Kapralov D.S., Shiyatov S.G., Moiseev P.A., Fomin V. (2006): A., Boldgiv B., Helliker B.R., Petraitis P.S., Casper B.B. Changes in the composition, structure, and altitudinal (2013): Plant response to climate change varies with topog-

510 J. FOR. SCI., 61, 2015 (11): 496–514 raphy, interactions with neighbors, and ecotype. Ecology, crobiological properties in Swiss Alpine grasslands. Soil 94: 444–453. Biology and Biochemistry, 69: 382–392. Lu N., Godt J.W. (2013): Hillslope Hydrology and Stability. Parmesan C. (2006): Ecological and evolutionary responses New York, Cambridge University Press: 458. to recent climate change. Annual Review of Ecology, Evolu- Lugon R., Stoffel M. (2010): Rock-glacier dynamics and magni- tion, and Systematics, 37: 637–669. tude–frequency relations of debris flows in a high-elevation Parviainen J., Bücking W., Vandekerkhove K., Schuck A., watershed: Ritigraben, Swiss Alps. Global and Planetary Päivinen R. (2000): Strict forest reserves in Europe: efforts Change, 73: 202–210. to enhance biodiversity and research on forests left for free Miehe G., Sabine Miehe S., Böhner J., Kaiser K., Hensen I., development in Europe (EU-COST-Action A4). Forestry, Madsen D., Liu J.Q., Opgenoorth L. (2014): How old is the 73: 107–118. human footprint in the world’s largest alpine ecosystem? Paul F., Kääb A., Maisch M., Kellenberger T., Haeberli W. A review of multiproxy records from the Tibetan Plateau (2004): Rapid disintegration of Alpine glaciers observed from the ecologists’ viewpoint. Quaternary Science Review, with satellite data. Geophysical Research Letters, 31: 86: 190–209. L21402. Mikan C.J., Schimel J.P., Doyle A.P. (2002): Temperature con- Pengerud A., Johnsen L.K., Mulder J., Strand L.T. (2014): trols of microbial respiration in arctic tundra soils above Potential adsorption of dissolved organic matter in poorly and below freezing. Soil Biology and Biochemistry, 34: podzolised, high-latitude soils. Geoderma, 226–227: 1785–1795. 39–46. Mikutta R., Kleber M., Torn M.S., Jahn R. (2006): Stabilization Pichler V., Codinho-Ferreira P., Zlatanov T., Pichlerová M., of soil organic matter: association with minerals or chemical Gregor J. (2011): Changes in forest cover and its diversity. recalcitrance? Biogeochemistry, 77: 25–56. In: Bredemeier M., Cohen S., Godbold D.L., Lode E., Morán-Tejeda E., López-Moreno J.I., Beniston M. (2013): Pichler V., Schleppi P. (eds): Forest Management and Water The changing roles of temperature and precipitation on Cycle. Berlin, Springer Verlag: 209–224. snowpack variability in Switzerland as a function of altitude. Pichler V., Hamor F., Vološčuk J., Sukharyuk D. (2007): Out- Geophysical Research Letters, 40: 2131–2136. standing Universal Value of the Ecological Processes in Muller S.W. (1945): Permafrost or permanently frozen ground the Primeval Beech Forests on the Carpathians and their and related engineering problems. Strategic Engineering Management as World Heritage Sites. Bratislava, VEDA Study, 62: 231. Publishing: 63. Muller S.W., French H., Nelson F. (2008): Frozen in time: Piermattei A., Garbarino M., Urbinati C. (2014): Struc- Permafrost and engineering problems. Reston, American tural attributes, tree-ring growth and climate sensitivity Society of Civil Engineers: 280. of Pinusnigra Arn. at high altitude: common patterns of Ni X., Yang W., Tan B., He J., Xu L., Li H., Wu F. (2015): Acceler- a possible treeline shift in the central Apennines (Italy). ated foliar litter humification in forest gaps: Dual feedbacks Dendrochronologia, 32: 210–219. of carbon sequestration during winter and the growing Ping C.L., Michaelson G.J., Kimble J.M., Romanovsky V.E., Shur season in an alpine forest. Geoderma, 241–242: 136–144. Y.L., Swanson D.K., Walker D.A. (2008): Cryogenesis and Noetzli J., Gruber S., Kohl T., Salzmann N., Haeberli W. (2007): soil formation along a bioclimate gradient in Arctic North Three-dimensional distribution and evolution of permafrost America. Journal of Geophysical Research, 113: G03–S12. temperatures in idealized high-mountain topography. Jour- Pitkänen A., Huttunen P., Jungner H., Merilainen J., Tolonen nal of Geophysical Research, 112: F02–S13, K. (2003): Holocene fire history of middle boreal pine forest Nyberg L., Stähli M., Mellander P.E., Bishop K.H. (2001): sites in eastern Finland. Annales Botanici Fennici, 40: 15–33. Soil frost effects on soil water and runoff dynamics along a Pregitzer K.S., King J.S., Burton A.J., Brown S.E. (2000): Re- boreal forest transect: 1. Field investigations. Hydrological sponses of tree fine roots to temperature. New Phytologists, Processes, 15: 909–926. 147: 105–115. OECD (2007): Climate Change in the European Alps: Adapt- Pretzsch H. (2005): Diversity and productivity in forests: ing Winter Tourism and Natural Hazards Management. evidence from long-term experimental plots. In: Scherer- Paris, OECD Publishing: 131. Available at http://www. Lorenzen M., Kőrner C., Schulze E.D. (eds): Forest Diversity oecd-ilibrary.org/docserver/download/9707061e.pdf?expi and Function: Temperate and Boreal Systems. Ecology Vol. res=1429871645&id=id&accname=ocid72027156&check 176. Berlin, Springer Verlag: 41–64. sum=8740231BC1624278AA042A84F0D1EF32 (accessed Prietzel J., Christophel D. (2014): Organic carbon stocks in for- 24 April, 2015). est soils of the German Alps. Geoderma, 221–222: 28–39. Oerlemans J. (2005): Extracting a climate signal from 169 Qi G., Wang Q., Zhou W., Ding H.,Wang X., Qi L., Wang Y., glacier records. Science, 308: 675–677. Li S., Dai L. (2011): Moisture effect on carbon and nitrogen Park J.H., Meusburger K., Jang I., Kang H., Alewell C. (2014): mineralization in topsoil of Changbai Mountain, Northeast Erosion-induced changes in soil biogeochemical and mi- China. Journal of Forest Science, 57: 340–348.

J. FOR. SCI., 61, 2015 (11): 496–514 511 Qingbai W., Yandong H., Hanbo Y., Yongzhi L. (2015): Schworer C., Henne P.D., Tinner W. (2014): A model-data Changes in active-layer thickness and near-surface per- comparison of Holocene timberline changes in the Swiss mafrost between 2002 and 2012 in alpine ecosystems, Alps reveals past and future drivers of mountain forest Qinghai-Xizang (Tibet) Plateau, China. Global and Plan- dynamics. Global Change Biology, 20: 1512–1526. etary Change, 124: 149–155. Serrano E., San Jose J.J., Agudo C. (2006): Rock glacier dynam- Rajkai K. (2008): The role of soil in bioclimatology – a review. ics in a marginal periglacial high mountain environment: Soil and Water Research, 3: 30–41. Flow, movement (1991–2000) and structure of the Argualas Rebetez M., Reinhard M. (2008): Monthly air temperature rock glacier, the Pyrenees. Geomorphology, 74: 285–296. trends in Switzerland 1901–2000 and 1975–2004. Theoreti- Shaver G.R., Giblin A., Nadelhoffer K.J., Thieler K.K., Downs cal and Applied Climatology, 91: 27–34. M.R., Laundre J.A., Rastetter E.B. (2006): Carbon turnover Rejšek K. (2004): The HadCH2 climate change model, elevat- in Alaskan tundra soils: effects of organic matter quality, ed CO2 model (SRES A2) and assessments on their likely temperature, moisture and fertilizer. Journal of Ecology, impacts on forest soils in The Czech Republic. Ekológia 94: 740–753. (Bratislava), 23: 80–87. Simonneau A., Chapron E., Vannière B., Wirth S.B., Gilli A., Di Roer I., Kääb A., Dikau R. (2005): Rockglacier acceleration Giovanni C., Anselmetti F.S., Desmet M., Magny M. (2012): in the Turtmann valley (Swiss Alps) probable controls. Multidisciplinary distinction of mass-movement and flood- Norwegian Journal of Geography, 59: 157–163. induced deposits in lacustrine environments: implications Le Roy M., Nicolussi K., Deline P., Astrade L., Edouard J.L., for Holocene palaeohydrology and natural hazards (Lake Miramont C., Arnaud F. (2015): Calendar-dated glacier Ledro, Southern Alps, Italy). Climate Past Discussion, 8: variations in the western European Alps during the Ne- 3205–3249. oglacial: the Mer de Glace record, Mont Blanc massif. Six J., Conant R.T., Paul E.A., Paustian K. (2002): Stabiliza- Quaternary Science Reviews, 108: 1–22. tion mechanisms of soil organic matter: implications for Rydgren K., Halvorsen R., Odland A., Skjerdal G. (2011): C-saturation of soils. Plant and Soil, 241: 155–176. Restoration of alpine spoil heaps: Successional rates predict Six J., Bossuyt H., Degryze S., Denef K. (2004): A history of vegetation recovery in 50 years. Ecological Engineering, research on the link between (micro) aggregates, soil biota, 37: 294–301. and soil organic matter dynamics. Soil Tillage Research, Samec P., Boublík K., Rejšek K. (2006): Climate change in 79: 7–31. problems of the submontane to subalpine altitudinal zones Smith-McKenna E.K., Malanson G.P., Resler L.M., Carstensen conditions variability. Meteorologický časopis, 9: 3–11. L.W., Prisley S.P., Tomback D.F. (2014): Cascading effects of Scherrer D., Körner C. (2011): Topographically controlled feedbacks, disease, and climate change on alpine treeline dy- thermal-habitat differentiation buffers diver- namics. Environmental Modeling and Software, 62: 85–96. sity against climate warming. Journal of Biogeography, Smith W.K., Germino M.J., Johnson D.M., Reinhardt K. (2009): 38: 406–416. The altitude of Alpine treeline: a bellwether of climate change Schimel P., Fahnestock J., Michaelson G., Mikan C., Ping C.L., effects. Botanical Reviews, 75: 163–190. Romanovsky V.E., Welker J. (2006): Cold-season produc- Solomina O.N., Bradley R.S., Hodgson D.A., Ivy-Ochs S.,

tion of CO2 in arctic soils: can laboratory and field estimates Jomelli V., Mackintosh A.N., Nesje A., Owen L.A., Wan- be reconciled through a simple modelling approach? Arc- ner H., Wiles G.C., Young N.E. (2015): Holocene glacier tic, Antarctic, and Alpine Research, 38: 249–256. fluctuations. Quaternary Science Reviews, 111: 9–34. Schmidt I.K., Jonasson S., Shaver G.R., Michelsen A., Nordin Solomon S., Qin D., Manning M., Chen Z., Marquis M., Averyt A. (2002): Mineralization and distribution of nutrients in K.B., Tignor M., Miller H.L. (2007): Contribution of Working plants and microbes in four arctic ecosystems: responses Group I to the 4th Assessment Report of the Intergovern- to warming. Plant and Soil, 242: 93–106. mental Panel on Climate Change. Cambridge, Cambridge Schmidt M.W.I., Torn M.S., Abiven S., Dittmar T., Guggen- University Press: 339. berger G., Janssens I.A., Kleber M., Kögel-Knabner I., Soncini A., Bocchiola D. (2011): Assessment of future snowfall Lehmann J., Manning D.A.C., Nannipieri P., Rasse D.P., regimes within the Italian Alps using general circulation Weiner S., Trumbore S.E. (2011): Persistence of soil organic models. Cold Regions Science and Technology, 68: 113–123. matter as an ecosystem property. Nature, 478: 49–56. Stanhill G., Cohen S. (2001): Global dimming: a review of Schuur E.A.G., Bockheim J,, Canadell J.G., Euskirchen E., the evidence for a widespread and significant reduction in Field C.B., Goryachkin S.V., Hagemann S., Kuhry P., Laf- global with discussion of its probable causes and leur P.M., Lee H., Mazhitova G., Nelson F.E., Rinke A., possible agricultural consequences. Agricultural Forest and Romanovsky, V.E., Shiklomanov N., Tarnocai C., Venevsky Meteorology, 107: 255–278. S., Vogel J.G., Zimov S.A. (2008): Vulnerability of perma- Stepanova V.A., Pokrovsky O.S., Viers J., Mironycheva-Toka- frost carbon to climate change: Implications for the global reva N.P., Kosykh N.P., Vishnyakova E.K. (2015): Elemental carbon cycle. Bioscience, 58: 701–714. composition of peat profiles in western Siberia: Effect of

512 J. FOR. SCI., 61, 2015 (11): 496–514 the micro-landscape, latitude position and permafrost Wang G., Mao T., Chang J., Du J. (2014): Impacts of surface coverage. Applied Geochemistry, 53: 53–70. soil organic content on the soil thermal dynamics of alpine Stoffel M., Tiranti D., Huggel C. (2014): Climate change impacts in permafrost regions: data from field observations. on mass movements – case studies from the European Alps. Geoderma, 232–234: 414–425. Science of the Total Environment, 493: 1255–1266. Weintraub M.N., Schimel J.P. (2003): Interactions between Sugiero D., Jaszczak R., Rączka G., Strzeliński P., Węgiel A., carbon and nitrogen mineralization and soil organic Wierzbicka A. (2009): Species composition in low mountain matter chemistry in arctic tundra soils. Ecosystems, 6: beech (Fagus sylvatica L.) stands in the Bieszczady National 129–143. Park under the global warming. Journal of Forest Science, Weintraub M.N., Schimel J.P. (2005): The seasonal dynam- 55: 244–250. ics of amino acids and other nutrients in Alaskan Arctic Szczypta C., Gascoin S., Houet T., Hagolle O., Dejoux J.F., Vi- tundra soils. Biogeochemistry, 73: 359–380. gneau C., Fanise P. (2015): Impact of climate and land cover Welc M., Frossard E., Egli S., Bünemann E.K., Jansa J. (2014): changes on snow cover in a small Pyrenean catchment. Journal Rhizosphere fungal assemblages and soil enzymatic activi- of Hydrology, 521: 84–99. ties in a 110-years alpine chronosequence. Soil Biology Tan B., Wu F.Z., Yang W.Q., He X.H. (2014): Snow removal alters and Biochemistry, 74: 21–30. soil microbial biomass and enzyme activity in a Tibetan alpine WGMS (ICSI-IAHS) (various years). Mass Balance Bul- forest. Applied Soil Ecology, 76: 34–41. letin. World Glacier Monitoring Service (International Targulian V.O., Goryachkin S.V. (2011): Soil memory and Commission on Snow and Ice), Zurich. Available at environmental reconstructions. Eurasian Soil Science, 44: http://www.wgms.ch/pub_wgms.html (accessed March 464–465. 26, 2015). Theurillat J.P., Guisan A. (2001): Potential impact of climate Wieser G., Leo M., Oberhuber W. (2014): Transpiration and change on vegetation in the European Alps: a review. Climatic canopy conductance in an inner alpine Scots pine (Pinus Change, 50: 77–109. sylvestris L.) forest. Flora, 209: 491–498. Thuiller W. (2004): Patterns and uncertainties of speceis´range Wischmeier W.H., Smith D.D. (1978): Predicting Rainfall shifts under climate change. Global Change Biology, 10: Erosion Losses. A Guide to Erosion Planning. Agriculture 2020–2027. Handbook Vol. 537. Washington, USDA: 62. Uhlířová E., Šantrůčková H., Davidov S.P. (2007): Quality and Wolf A., Lazzarotto P., Bugmann H. (2012): The relative potential biodegradability of soil organic matter preserved importance of land use and climatic change in Alpine in permafrost of Siberian tussock tundra. Soil Biology and catchments. Climatic Change, 111: 279–300. Biochemistry, 39: 1978–1989. Wojciech D. (2012): The cryosphere and glacial permafrost Valese E., Conedera M., Held A.C., Ascoli D. (2014): Fire, hu- as its integral component. Central European Journal of mans and landscape in the European Alpine region during Geoscience, 4: 623–640. the Holocene. Anthropocene, 6: 63–74. Xu M., Peng F., You Q., Guo J., Tian X., Xue X., Liu M. Vilmundardóttir O.K., Gísladóttir G., Lal R. (2015): Soil carbon (2015): Year-round warming and autumnal clipping lead accretion along an age chronosequence formed by the retreat to downward transport of root biomass, carbon and total of the Skaftafellsjökull glacier, SE-Iceland. Geomorphology, nitrogen in soil of an alpine . Environmental and 228: 124–133. Experimental , 109: 54–62. Von Lützow M., Kögel-Knabner I., Ekschmitt K., Matzner E., You Q.G., Xue X., Peng F., Xu M., Duan H., Dong S. (2014): Guggenberger G., Marschner B., Flessa H. (2006): Stabilization Comparison of ecosystem characteristics between de- of organic matter in temperate soils: mechanisms and their graded and intact alpine meadow in the Qinghai-Tibetan relevance under different soil conditions – a review. European Plateau. China Ecological Engineering, 71: 133–143. Journal of Soil Science, 57: 426–467. Zanella A., Jabiol B., Ponge J.F., Sartori G., De Waal R., Van Von Lützow M., Kögel-Knabner I. (2009): Temperature sensitiv- Delft B., Graefe U., Cools N., Katzensteiner K., Hager H., ity of soil organic matter decomposition-what do we know? Englisch M., Brethes A., Broll G., Gobat J.M., Brun J.J, Biology and Fertility of Soils, 46: 1–15. Milbert G., Kolb E., Wolf U., Frizzera L., Galvan P., Kolli Vranová V., Pavelka M., Rejšek K., Formánek P. (2011): Impact R., Baritz R., Kemmers R., Vacca A., Serra G., Banas D., of land-use change on seasonal dynamics of total protein Garlato A., Chersich S., Klimo E., Langohr R. (2011): flow from roots of mountain meadow plant communities. European Humus Forms Reference Base. Available at In: Richards K.E. (ed.): Mountain Ecosystems: Dynamics, http://hal.archives-ouvertes.fr/docs/00/56/17/95/PDF/ Management and Conservation. New York, Nova Science Humus_Forms_ERB_31_01_2011.pdf (accessed March Publishers: 93–100 26, 2015). Walther G.R., Beissner S., Burga C.A. (2005): Trends in the Zollinger B., Alewell C., Kneisel C., Meusburger K., Gärtner upward shift of alpine plants. Journal of Vegetation Science, H., Brandová D., Ivy-Ochs S., Schmidt M.W.I., Egli M. 16: 541–548. (2013): Effect of permafrost on the formation of soil or-

J. FOR. SCI., 61, 2015 (11): 496–514 513 ganic carbon pools and their physical-chemical properties between an alpine ecosystem and the atmosphere in the Eastern Swiss Alps. Catena, 110:70–85. on the Nam Co area of the Tibetan Plateau. Agricultural Zemp M., Frauenfelder R., Haeberli W., Hoelzle M. (2005): and Forest Meteorology, 203: 169–179. Worldwide glaciers mass balance measurements: General Zongxing L., Qi F., Wei L., Tingting W., Aifang C., Yan G., trends and first results of the extraordinary year 2003 in Xiaoyan G., Yanhui P., Jianguo L., Rui G., Bing J. (2014): Central Europe. Data Glaciological Studies, 99: 3–12. Study on the contribution of cryosphere to runoff in the Zemp M., Haeberli W., Hoelze M., Paul F. (2006): Alpine cold alpine basin: A case study of Hulugou River Basin in glaciers to disappear within decades? Geophysical Research the Qilian Mountains. Global Planet Change, 122: 345–361. Letters, 33: L13504. Zhu Z., Ma Y., Li M., Zeyong Hu Z., Xu C., Zhang L., Han Received for publication May 11, 2015 C., Wang Y., Ichiro T. (2015): Carbon dioxide exchange Accepted after corrections October 30, 2015

Corresponding author: Silvia Chersich, Ph.D., University of Pavia, Department of Earth and Environmental Sciences, Via Ferrata 1, 27100 Pavia, Italy. e-mail: [email protected];

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