Among‐tree variability and feedback effects result in different growth responses to climate change at the upper treeline in the Swiss Alps

Upper treeline ecotones are important life form boundaries and particularly sensitive to a warming climate. Changes in growth conditions at these ecotones have wide‐ranging implications for the provision of ecosystem services in densely populated mountain regions like the European Alps. We quantify climate effects on short‐ and long‐term tree growth responses, focusing on among‐tree variability and potential feedback effects. Although among‐tree variability is thought to be substantial, it has not been considered systematically yet in studies on growth–climate relationships. We compiled tree‐ring data including almost 600 trees of major treeline species (Larix decidua, Picea abies, Pinus cembra, and Pinus mugo) from three climate regions of the Swiss Alps. We further acquired tree size distribution data using unmanned aerial vehicles. To account for among‐tree variability, we employed information‐theoretic model selections based on linear mixed‐effects models (LMMs) with flexible choice of monthly temperature effects on growth. We isolated long‐term trends in ring‐width indices (RWI) in interaction with elevation. The LMMs revealed substantial amounts of previously unquantified among‐tree variability, indicating different strategies of single trees regarding when and to what extent to invest assimilates into growth. Furthermore, the LMMs indicated strongly positive temperature effects on growth during short summer periods across all species, and significant contributions of fall (L. decidua) and current year's spring (L. decidua, P. abies). In the longer term, all species showed consistently positive RWI trends at highest elevations, but different patterns with decreasing elevation. L. decidua exhibited even negative RWI trends compared to the highest treeline sites, whereas P. abies, P. cembra, and P. mugo showed steeper or flatter trends with decreasing elevation. This does not only reflect effects of ameliorated climate conditions on tree growth over time, but also reveals first signs of long‐suspected negative and positive feedback of climate change on stand dynamics at treeline.     

Mosses modify effects of warmer and wetter conditions on tree seedlings at the alpine treeline

Climate warming enables tree seedling establishment beyond the current alpine treeline, but to achieve this, seedlings have to establish within existing tundra vegetation. In tundra, mosses are a prominent feature, known to regulate soil temperature and moisture through their physical structure and associated water retention capacity. Moss presence and species identity might therefore modify the impact of increases in temperature and precipitation on tree seedling establishment at the arctic‐alpine treeline. We followed Betula pubescens and Pinus sylvestris seedling survival and growth during three growing seasons in the field. Tree seedlings were transplanted along a natural precipitation gradient at the subarctic‐alpine treeline in northern Sweden, into plots dominated by each of three common moss species and exposed to combinations of moss removal and experimental warming by open‐top chambers (OTCs). Independent of climate, the presence of feather moss, but not Sphagnum, strongly supressed survival of both tree species. Positive effects of warming and precipitation on survival and growth of B. pubescens seedlings occurred in the absence of mosses and as expected, this was partly dependent on moss species. P. sylvestris survival was greatest at high precipitation, and this effect was more pronounced in Sphagnum than in feather moss plots irrespective of whether the mosses had been removed or not. Moss presence did not reduce the effects of OTCs on soil temperature. Mosses therefore modified seedling response to climate through other mechanisms, such as altered competition or nutrient availability. We conclude that both moss presence and species identity pose a strong control on seedling establishment at the alpine treeline, and that in some cases mosses weaken climate‐change effects on seedling establishment. Changes in moss abundance and species composition therefore have the potential to hamper treeline expansion induced by climate warming.     

High and dry: post‐fire tree seedling establishment in subalpine forests decreases with post‐fire drought and large stand‐replacing burn patches

Aim Climate warming and increased wildfire activity are hypothesized to catalyse biogeographical shifts, reducing the resilience of fire‐prone forests world‐wide. Two key mechanisms underpinning hypotheses are: (1) reduced seed availability in large stand‐replacing burn patches, and (2) reduced seedling establishment/survival after post‐fire drought. We tested for regional evidence consistent with these mechanisms in an extensive fire‐prone forest biome by assessing post‐fire tree seedling establishment, a key indicator of forest resilience. Location Subalpine forests, US Rocky Mountains. Methods We analysed post‐fire tree seedling establishment from 184 field plots where stand‐replacing forest fires were followed by varying post‐fire climate conditions. Generalized linear mixed models tested how establishment rates varied with post‐fire drought severity and distance to seed source (among other relevant factors) for tree species with contrasting post‐fire regeneration adaptations. Results Total post‐fire tree seedling establishment (all species combined) declined sharply with greater post‐fire drought severity and with greater distance to seed sources (i.e. the interior of burn patches). Effects varied among key species groups. For conifers that dominate present‐day subalpine forests (Picea engelmannii, Abies lasiocarpa), post‐fire seedling establishment declined sharply with both factors. One exception was serotinous Pinus contorta, which did not vary with either factor. For montane species expected to move upslope under future climate change (Larix occidentalis, Pseudotsuga menziesii, Populus tremuloides) and upper treeline species (Pinus albicaulis), establishment was unrelated to either factor. Greater post‐fire tree seedling establishment on cooler/wetter aspects suggested local topographic refugia during post‐fire droughts. Main conclusions If future drought and wildfire patterns manifest as expected, post‐fire tree seedling establishment of species that currently characterize subalpine forests could be substantially reduced. Compensatory increases from lower montane and upper treeline species may partially offset these reductions, but our data suggest important near‐ to mid‐term shifts in the composition and structure of high‐elevation forests under continued climate warming and increased wildfire activity.     

Isolation and characterization of polymorphic nuclear microsatellite loci in Pinus cembra L

We developed eight polymorphic nuclear microsatellite markers for the Swiss stone pine (Pinus cembra L.), of which seven may be amplified in a multiplex polymerase chain reaction. Allelic polymorphism across all loci and 40 individuals representing two populations in the Swiss Alps was high (mean = 7.6 alleles). No significant linkage disequlibrium was displayed between pairs of loci. Significant deviation from Hardy–Weinberg equilibrium was revealed at three loci in one population. Cross–amplification was achieved in two related species within the genus (P. sibirica and P. pumila). Thus, the markers may be useful for population genetic studies in these three pine species. They will be applied in ongoing projects on genetic diversity and patterns of gene flow in P. cembra.     

Short-term signals of climate change along an altitudinal gradient in the South Alps

Short-term changes in plant species number, frequency and composition were studied along an altitudinal gradient crossing four summits from the treeline ecotone to the subnival zone in the South Alps (Dolomites, Italy). Large-scale (summit areas) and small-scale patterns (16 plots of 1 m2/summit) were monitored. After 5 years, a re-visitation of the summit areas revealed a considerable increase of species richness at the upper alpine and subnival zone (10% and 9%, respectively) and relatively modest increases at the lower alpine zone and the treeline ecotone (3% and 1%, respectively). At the small scale, the results were partly different, with species richness decreasing at the lower summits and increasing at the higher summits. The changes can most likely be attributed to climate warming effects and to competitive interactions. The main newcomers at the lower three summits were species from the treeline and the lower altitudinal zones. Only at the highest summit, the newcomers came from the alpine species pool. At the treeline ecotone, the abundance of Pinus cembra, of dwarf shrubs and clonal graminoid species increased. Here, displacements of alpine species may be predicted for the near future. At the higher summits, expansions of the established alpine species and further invasions of species from lower altitudes are forecasted.     

A latitudinal gradient in tree growth response to climate warming in the Siberian taiga

We investigated the climate response of three Siberian taiga species, Larix cajanderi, Picea obovata, and Pinus sylvestris, across a latitudinal gradient in central Siberia. We hypothesized that warming is more frequently associated with increased growth for evergreen conifers (P. obovata and P. sylvestris) than for L. cajanderi, and for northern than for southern sites; we also hypothesized that increased growth is associated with a positive trend in normalized difference vegetation index (NDVI). In mixed stands, growth of L. cajanderi and P. obovata increased over time, but the larger growth increases in P. obovata may presage a shift in competitive balance between these species. Climate response varied among and within populations of all species, and positive responses to temperature prevailed at northern sites, where trees grew faster in years with warm early summers. Negative responses to warming declined along the south to north latitudinal gradient. We observed considerable variability in climate response within populations which even exceeded that among species or sites. Tree response to climate was correlated with NDVI trends, indicating that patterns of tree‐growth response to climate were indicative of a coherent landscape‐scale response to warming. Our findings suggest that increased productivity with warming is likely only in the northern reaches of the Siberian taiga. An increased prevalence of evergreen conifers in areas currently dominated by deciduous Larix species also seems likely.     

Temperature as a predictor of survival of the pine processionary moth in the Italian Alps

1 The pine processionary moth Thaumetopoea pityocampa is expanding its geographical range in Europe, as a consequence of enhanced winter survival under a warmer climate. A combination of daytime nest temperatures and night air temperatures determines the number of hours larvae are able to feed (hours above realized feeding threshold, RFT). 2 We tested the RFT‐based model for survival across multiple areas of the insect’s range in the Italian Alps over a 2‐year period. In a series of translocation experiments using natural temperature gradients as spatial analogues for global warming, we transferred colonies of T. pityocampa larvae to sites within zones of historical distribution, recent distribution, and outside the present range. The sites included traditional (Pinus nigra, Pinus sylvestris) as well as novel (Pinus mugo, Pinus uncinata, Pinus cembra) hosts. 3 Survival during precold (August to November), cold (December to February) and postcold (March to May) periods were analysed against climatic variables (temperature and rainfall) and predictors developed by the model. 4 Host species did not significantly affect final survival, with the exception of slower larval development, and resulting lower cold tolerance, on P. cembra than on P. mugo at the same site. 5 Across all the sites and hosts, final survival of colonies depended on the number of feeding hours during the cold period (RFT), which explained 82% of the variance in a regression model. We recommend using RFT, or its surrogate daily mean minimum temperature when nest temperature is not available, in predictive models of range expansion of T. pityocampa under climate change scenarios.     

Holocene timber-line dynamics at Bachalpsee, a lake at 2265 m a.s.l. in the northern Swiss Alps

Palaeobotanical analysis and radiocarbon dating of lake sediments from Bachalpsee (2265 m a.s.l.), a small lake above the present-day timber-line in the northern Swiss Alps reveals that the region was already deglaciated during the Younger Dryas. The sediment record is dominated by long-distance transported pollen that originates from lowland vegetation but the plant macrofossils give evidence of the local vegetation development. Comparison with palaeobotanical results from three sites along an altitudinal transect permits the reconstruction of the regional timber-line history. Throughout the entire Holocene the catchment of Bachalpsee consisted of a mosaic of open meadows and dwarf shrubs (Salix, Vaccinium, Rhododendron). Chironomid and cladoceran assemblages suggest that the early to mid-Holocene was the warmest interval at Bachalpsee. Comparison of the palaeobotanical results with those from the mire “Feld” (2130 m a.s.l.) in the vicinity of Bachalpsee showed that during the mid-Holocene the timber-line was formed by Pinus cembra and Picea abies with some scattered Abies alba trees and was situated close to Bachalpsee but never reached its catchment. The maximum timber-line in the Northern Alps was reached between 6000 and 3000 cal b.p. which is several millennia later than in the Central Alps. The species composition of the tree-line (Abies alba, Pinus cembra and the absence of Larix decidua) points to less continental and moister climatic conditions compared with the central Alps during the early to mid-Holocene. From 3000 cal b.p. onwards the timber-line was lowered by human deforestation with the most intense pulses of human impact occurring since the Middle Ages. The catchment of Bachalpsee has been used as alpine pasture since the Bronze Age.     

Competitor or facilitator? The ambiguous role of alpine grassland for the early establishment of tree seedlings at treeline

Alpine treelines are expected to move upslope with a warming climate. However, so far treelines have responded inconsistently and future shifts remain difficult to predict since many factors unrelated to temperature, such as biotic interactions, affect responses at the local scale. Especially during the earliest regeneration stages, trees can be strongly influenced by alpine vegetation via both competition and facilitation. We aimed to understand the relative importance of these two types of interaction in different vegetation structures for treeline regeneration dynamics. Effects of herbaceous alpine vegetation on seedling emergence and first‐year performance were studied in a field experiment in the French Alps (2100 m a.s.l.) with five important European treeline tree species: Larix decidua, Picea abies, Pinus cembra, Pinus uncinata and Sorbus aucuparia. Total emergence and locally‐germinated seedling survival were not affected, but for seedlings planted at two months of age, negative vegetation impacts dominated for all response parameters: first‐year survival, growth and carbohydrate accumulation. However, in the winter half‐year, evergreen tree seedlings increased carbohydrate reserves under the protection of senescent herbs. Also, responses of locally‐germinated seedlings suggest facilitative vegetation effects in the first two months after emergence. Thus, the interaction switched between competition and facilitation according to ontogenetic stage and seasons. Still, the net outcome after one year was negative, but species differed in their susceptibilities. Because initial establishment is the first bottleneck determining whether treelines remain stable or move upslope, understanding establishment, including site‐, life‐stage and species‐specific processes, is essential for understanding observed treeline spatial patterns and dynamics. When developing predictive models of treeline dynamics, all these ‘local’ aspects should be incorporated in addition to more global drivers like changes in temperature.     

A Pinus cembra L. tree-ring record for late spring to late summer temperature in the Rhaetian Alps,Italy

Ongoing climate change strongly affects high-elevation environments in the European Alps, influencing the cryosphere and the biosphere and causing widespread retreat of glaciers and changes in biomes. Nevertheless, high-elevation areas often lack long meteorological series, and global datasets cannot represent local variations well. Thus, proxy data, such as tree rings, provide information on past climatic variations from these remote sites. Although maximum latewood density (MXD) chronologies provide better temperature information than those based on tree-ring width (TRW), MXD series from the European Alps are lacking. To derive high-quality temperature information for the Rhaetian Alps, Pinus cembra L. trees sampled at approximately 2000 m a.s.l. were used to build one MXD chronology spanning from 1647 to 2015. The MXD data were significantly and highly correlated with seasonal May-September mean temperatures. The MXD chronology showed a generally positive trend since the middle of the 19^th century, interrupted by short phases of climatic deterioration in the beginning of the 20^th century and in the 1970s, conforming with the temperature trends. Our results underline the potential for using Pinus cembra L. MXD to reconstruct mean temperature variations, especially during the onset and latter part of the growing season, providing additional information on parts of the growing season not inferred from TRW. Future studies on MXD for this species will increase the availability of temporal and spatial data, allowing detailed climate reconstructions.     

A model‐data comparison of Holocene timberline changes in the Swiss Alps reveals past and future drivers of mountain forest dynamics

Mountain vegetation is strongly affected by temperature and is expected to shift upwards with climate change. Dynamic vegetation models are often used to assess the impact of climate on vegetation and model output can be compared with paleobotanical data as a reality check. Recent paleoecological studies have revealed regional variation in the upward shift of timberlines in the Northern and Central European Alps in response to rapid warming at the Younger Dryas/Preboreal transition ca. 11 700 years ago, probably caused by a climatic gradient across the Alps. This contrasts with previous studies that successfully simulated the early Holocene afforestation in the (warmer) Central Alps with a chironomid‐inferred temperature reconstruction from the (colder) Northern Alps. We use LandClim , a dynamic landscape vegetation model to simulate mountain forests under different temperature, soil and precipitation scenarios around Iffigsee (2065 m a.s.l.) a lake in the Northwestern Swiss Alps, and compare the model output with the paleobotanical records. The model clearly overestimates the upward shift of timberline in a climate scenario that applies chironomid‐inferred July‐temperature anomalies to all months. However, forest establishment at 9800 cal. BP at Iffigsee is successfully simulated with lower moisture availability and monthly temperatures corrected for stronger seasonality during the early Holocene. The model‐data comparison reveals a contraction in the realized niche of Abies alba due to the prominent role of anthropogenic disturbance after ca. 5000 cal. BP, which has important implications for species distribution models (SDMs) that rely on equilibrium with climate and niche stability. Under future climate projections, LandClim indicates a rapid upward shift of mountain vegetation belts by ca. 500 m and treeline positions of ca. 2500 m a.s.l. by the end of this century. Resulting biodiversity losses in the alpine vegetation belt might be mitigated with low‐impact pastoralism to preserve species‐rich alpine meadows.     

Recent treeline dynamics are similar between dry and mesic areas of Nepal,central Himalaya

Aims We investigated the treeline dynamics of two environmentally contrasting areas in the Nepalese Himalaya to address the following questions: (i) Does the timing of establishment of the current treeline differ between the two study areas, and can area-specific treeline developments be identified? (ii) Do recruitment patterns and height growth indicate recent climate-driven treeline advance, following the general prediction for the central Himalayan region, in the two study areas?Methods A dry-climate treeline dominated by Pinus wallichiana and a mesic-climate treeline with Abies spectabilis were selected for study. In each area, we sampled the size and age structure of the study species along three elevational transects (20-m wide) from the forest line to the tree species line crossing the treeline. We also sampled treeline trees from within and outside transects to reconstruct past treeline establishment dynamics.Important findings Despite differences in moisture regimes, tree species and recent climate trends, our two study areas showed very similar treeline dynamics over the past six decades. In both areas, the recruitment of treeline trees indicates stationary treelines over the past six decades with the current treelines being dominated by trees that were established around 1990. The mesic area has experienced an overall climatic warming trend, and the stationary Abies treeline is hypothesized to be regulated by non-climatic factors, notably grazing. The dry area has not experienced warming but increased climatic variability and some very cool summers in the recent decades may explain the stationary to weakly receding Pinus treeline, which appears more climatically controlled with decreased recruitment over the past decades and decreased growth towards higher elevations. In both areas, there is a potential for treeline advance, depending on future land use and climate change. Our results highlight the importance of conducting treeline ecotone analyses for several sites or areas, and considering both climatic and non-climatic drivers of the treeline dynamics within each of these areas, for understanding regional treeline dynamics.     

Evidence of enhanced freezing damage in treeline plants during six years of CO2 enrichment and soil warming

Climate change and elevated atmospheric CO₂ levels could increase the vulnerability of plants to freezing. We analyzed tissue damage resulting from naturally occurring freezing events in plants from a long–term in situ CO₂ enrichment (+ 200 ppm, 2001–2009) and soil warming (+ 4°C since 2007) experiment at treeline in the Swiss Alps (Stillberg, Davos). Summer freezing events caused damage in several abundant subalpine and alpine plant species in four out of six years between 2005 and 2010. Most freezing damage occurred when temperatures dropped below –1.5°C two to three weeks after snow melt. The tree Larix decidua and the dwarf shrubs Vaccinium myrtillus and Empetrum hermaphroditum showed more freezing damage under experimentally elevated CO₂ and/or temperatures than under control conditions. Soil warming induced a 50% die‐back of E. hermaphroditum during a single freezing event due to melting of the protective snow cover. Although we could not identify a clear mechanism, we relate greater freezing susceptibility to a combination of advanced plant phenology in spring and changes in plant physiology. The climate record since 1975 at the treeline site indicated a summer warming by 0.58°C/decade and a 3.5 days/decade earlier snow melt, but no significant decrease in freezing events during the vegetation period. Therefore, in a warmer climate with higher CO₂ levels but constant likelihood of extreme weather events, subalpine and alpine plants may be more susceptible to freezing events, which may partially offset expected enhanced growth with global change. Hence, freezing damage should be considered when predicting changes in growth of alpine plants or changes in community composition under future atmospheric and climate conditions.     

Temporal dynamic of wood formation in Pinus cembra along the alpine treeline ecotone and the effect of climate variables

We determined the temporal dynamic of cambial activity and xylem development of stone pine (Pinus cembra L.) throughout the treeline ecotone. Repeated micro-sampling of the developing tree ring was carried out during the growing seasons 2006 and 2007 at the timberline (1,950 m a.s.l.), treeline (2,110 m a.s.l.) and within the krummholz belt (2,180 m a.s.l.) and the influence of climate variables on intra-annual wood formation was determined. At the beginning of both growing seasons, highest numbers of cambial and enlarging cells were observed at the treeline. Soil temperatures at time of initiation of cambial activity were c. 1.5°C higher at treeline (open canopy) compared to timberline (closed canopy), suggesting that a threshold root-zone temperature is involved in triggering onset of above ground stem growth. The rate of xylem cell production determined in two weekly intervals during June through August 2006–2007 was significantly correlated with air temperature (temperature sums expressed as degree-days and mean daily maximum temperature) at the timberline only. Lack of significant relationships between tracheid production and temperature variables at the treeline and within the krummholz belt support past dendroclimatological studies that more extreme environmental conditions (e.g., wind exposure, frost desiccation, late frost) increasingly control tree growth above timberline. Results of this study revealed that spatial and temporal (i.e., year-to-year) variability in timing and dynamic of wood formation of P. cembra is strongly influenced by local site factors within the treeline ecotone and the dynamics of seasonal temperature variation, respectively.     

The effects of wind and temperature on cuticular transpiration of Picea abies and Pinus cembra and their significance in dessication damage at the alpine treeline

Summary The importance of high winter winds and plant temperatures as causes of winter desiccation damage at the alpine treeline were studied in the Austrian Alps. Samples of 1- and 2-year twigs of Picea abies and Pinus cembra were collected from the valley bottom (1,000 m a.s.l.), forestline (1,940 m a.s.l.), kampfzone (2.090 m a.s.l.), wind-protected treeline (2,140 m a.s.l.), and wind-exposed treeline (2,140 m a.s.l.). Cuticular transpiration was measured at three different levels of wind speed (4, 10, and 15 ms^-1) and temperature (15°, 20°, and 25° C). At elevated wind speeds slight increases in water loss were observed, whereas at higher temperatures much greater increases occurred. Studies on winter water relations show a significant decline in the actual moisture content and osmotic potentials of twigs, especially in the kampfzone and at treeline. The roles of high winds and temperatures in depleting the winter water economy and causing desiccation damage in the alpine treeline environment are discussed.     

Early season temperature controls cambial activity and total tree ring width at the alpine treeline

Background: Temperature directly affects xylogenesis at high-elevation treelines. The low-temperature limitation of meristematic processes is thus key to understand treeline formation.

Aims: We aimed to experimentally test in situ the direct low-temperature effect on wood tissue formation at the alpine treeline.

Methods: We applied controlled Peltier-mediated cooling and warming (±3 K) to branch segments in Pinus uncinata at the treeline in the Swiss Alps. In addition, we studied xylogenesis in untreated trees during the growing season by sequential micro-coring.

Results: Micro-cores indicated that the cambial zone was fully developed by the time the cooling and warming treatment started, shortly after snowmelt. Presumably, because of this, experimental cooling of branches did not significantly reduce the number of cells produced per season. Warming extended the formation of early wood into the late season, and thus reduced the fraction of late wood.

Conclusions: We conclude that temperatures very early in the season determine the width of the cambial zone which, in turn, strongly controls the number of tracheids produced during the remaining growing season. Temperatures later in the season mainly determine the early wood to late wood ratio. These data provide an empirical basis for the mechanistic understanding of tree growth at the treeline in response to temperature.     

Tree-rings,genetics and the environment: Complex interactions at the rear edge of species distribution range

Under climate change, modifications on plants’ growth are expected to be the strongest at species margins. Therein, tree acclimation could play a key role as migration is predicted to be too slow to track shifts of bioclimatic envelops. A requirement is, however, that intra-population genetic diversity be high enough for allowing such adaptation of tree populations to climate change. In this study, we tested for the existence of relationships between genetic diversity, site environmental conditions, and the response of annual tree growth to climate of Pinus cembra at its southern limit in the Alps. Site-specific climatic and environmental factors predominantly determined the response of trees along the precipitation gradient. The growth-climate interactions were chiefly linked to mean annual precipitation and temperature, slope and tree-size, and less to genetic diversity. We show that genetic background of Pinus cembra has exclusively indirect modulating power with limited effects on tree-ring formation, and within the southern limit in the Alps, genetic variability is not necessarily well expressed in the patterns of annual tree growth. Our results may imply little adaptive capacity of these populations to future changes in the water balance.     

The Early Holocene treeline in the southern French Alps: new evidence from travertine formations

Aims The present paper concerns the analysis of macro‐remains (plant imprints) from high altitude travertine deposits dating back to the early Holocene (9800 BP ). Our results allow us to discuss treeline location and ecology, and to supplement previous data based on pollen from natural sediments and charcoal from natural soil. Location The travertine under study is located in the Queyras massif, in the southern French Alps, along the Italian border. The site is situated on a south‐facing slope. Methods The travertine deposit has been dated by ²³⁰Th/²³⁴U. Taxonomic identification of plant imprints was achieved by comparison of the morphology of fossil specimens with extant material. Results Imprints of Pinus uncinata (cones and needles), broad‐leaved trees (Betula cf. pubescens, Populus tremula and Salix spp.), and unidentified trunks have been found. The plant assemblage records the existence at a high altitude (2200 m) of shade‐intolerant vegetation at around 9800 BP. Our data indicate that the treeline limit was at least 100 m higher than previously thought. The morphological features of the travertine, the plant assemblage and trunk diameters indicate that during the travertinization process, vegetation around the site was probably dense with tall upright trees. Main conclusions Our data show that Pinus uncinata and broad‐leaved trees developed at 2200 m, while regional pollen analyses locate the treeline at lower altitudes. Different research methods appear to provide different results. Around 9800 BP tree regeneration and growth were made possible up to at least 2200 m a.s.l., probably as a result of warmer and wetter summers. Warmer conditions are inferred from the predicted increase in incoming solar radiation based on calculated orbital variations, and wetter conditions are inferred from the build‐up of the travertine.     

Variation in the chloroplast DNA of Swiss stone pine (Pinus cembra L.) reflects contrasting post-glacial history of populations from the Carpathians and the Alps

Aim To characterize the genetic structure and diversity of Pinus cembra L. populations native to two disjunct geographical areas, the Alps and the Carpathians, and to evaluate the rate of genetic differentiation among populations. Location The Swiss Alps and the Carpathians. Methods We screened 28 populations at three paternally inherited chloroplast simple sequence repeats (cpSSRs) for length variation in their mononucleotide repeats. Statistical analysis assessed haplotypic variation and fixation indices. Hierarchical analysis of molecular variance (AMOVA), Mantel test, spatial analysis of molecular variance (SAMOVA) and barrier analyses were applied to evaluate the geographical partitioning of genetic diversity across the species’ range. Results Haplotypic diversity was generally high throughout the natural range of P. cembra, with the mean value substantially higher in the Carpathians (H = 0.53) than in the Alps (H = 0.35). The isolated Carpathian populations showed the highest haplotype diversity among the populations originating from the High Tatras (Velka Studena Dolina) and South Carpathians (Retezat Mountains). AMOVA revealed that only 3% of the total genetic variation derived from genetic differentiation between the two mountain ranges. Differentiation among Carpathian populations was higher (F_ST = 0.19) than among Alpine populations (F_ST = 0.04). Low, but significant, correlation was found between the geographical and genetic distances among pairs of populations (r = 0.286, P < 0.001). SAMOVA results revealed no evident geographical structure of populations. barrier analysis showed the strongest differentiation in the eastern part of the species’ range, i.e. in the Carpathians. Main conclusions The populations of P. cembra within the two parts of the species’ range still share many cpDNA haplotypes, suggesting a common gene pool conserved from a previously large, continuous distribution range. Carpathian populations have maintained high haplotypic variation, even higher than Alpine populations, despite their small population sizes and spatial isolation. Based on our results, we emphasize the importance of the Carpathian populations of Swiss stone pine for conservation. These populations comprise private haplotypes and they may represent a particular legacy of the species’ evolutionary history.     

Phenotypic plasticity facilitates resistance to climate change in a highly variable environment

Increased summer drought will exacerbate the regeneration of many tree species at their lower latitudinal and altitudinal distribution limits. In vulnerable habitats, introduction of more drought-tolerant provenances or species is currently considered to accelerate tree species migration and facilitate forest persistence. Trade-offs between drought adaptation and growth plasticity might, however, limit the effectiveness of assisted migration, especially if introductions focus on provenances or species from different climatic regions. We tested in a common garden experiment the performance of Pinus sylvestris seedlings from the continental Central Alps under increased temperatures and extended spring and/or summer drought, and compared seedling emergence, survival and biomass allocation to that of P. sylvestris and closely related Pinus nigra from a Mediterranean seed source. Soil heating had only minor effects on seedling performance but high spring precipitation doubled the number of continental P. sylvestris seedlings present after the summer drought. At the same time, twice as many seedlings of the Mediterranean than the continental P. sylvestris provenance were present, which was due to both higher emergence and lower mortality under dry conditions. Both P. sylvestris provenances allocated similar amounts of biomass to roots when grown under low summer precipitation. Mediterranean seedlings, however, revealed lower phenotypic plasticity than continental seedlings under high precipitation, which might limit their competitive ability in continental Alpine forests in non-drought years. By contrast, high variability in the response of individual seedlings to summer drought indicates the potential of continental P. sylvestris provenances to adapt to changing environmental conditions.