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Seed dispersal is a key component in a plant's life cycle, allowing the colonization of new habitat and eventually enabling range shifts. For animal‐dispersed plants, the behaviour of seed‐dispersing animals has direct consequences for the fate of a seed. However, there is little empirical evidence on how individual behaviour of animals affects seed dispersal.Here we investigated the effects of individual behaviour on seed dispersal distance and location by analysing movement data of spotted nutcrackers (Nucifraga caryocatactes) during harvest and scatter‐hoarding of Swiss stone pine (Pinus cembra) seeds. To assess individual variation in seed dispersal patterns, we compared daytime hourly displacement patterns across individuals. We further identified seed caching locations of individual birds and analysed their potential for pine regeneration.We found that nutcrackers showed one of two distinct behaviours: short‐distance seed dispersal and long‐distance seed dispersal. The caching locations resulting from the two behaviours differed markedly in the regeneration potential of Swiss stone pine: seeds transported over short distances were cached at high elevations with high regeneration potential within the pine's current habitat, while seeds transported over long distances were cached at low elevations with low regeneration potential outside of the pine's habitat. The behaviours were not associated with bird body mass, relative wing length or sex.Our results emphasize that intraspecific individual behaviours of seed‐dispersing animals can have far‐reaching consequences for ecosystem functions. As a major determinant of seed dispersal dynamics, individual behaviour should be taken into account for a better understanding of ecosystem functions under current and future environmental conditions. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

It is generally accepted that the spatial distribution of neutral genetic diversity within a species' native range mostly depends on effective population size, demographic history, and geographic position. However, it is unclear how genetic diversity at adaptive loci correlates with geographic peripherality or with habitat suitability within the ecological niche. Using exome‐wide genomic data and distribution maps of the Alpine range, we first tested whether geographic peripherality correlates with four measures of population genetic diversity at > 17,000 SNP loci in 24 Alpine populations (480 individuals) of Swiss stone pine (Pinus cembra) from Switzerland. To distinguish between neutral and adaptive SNP sets, we used four approaches (two gene diversity estimates, FST outlier test, and environmental association analysis) that search for signatures of selection. Second, we established ecological niche models for P. cembra in the study range and investigated how habitat suitability correlates with genetic diversity at neutral and adaptive loci. All estimates of neutral genetic diversity decreased with geographic peripherality, but were uncorrelated with habitat suitability. However, heterozygosity (He) at adaptive loci based on Tajima's D declined significantly with increasingly suitable conditions. No other diversity estimates at adaptive loci were correlated with habitat suitability. Our findings suggest that populations at the edge of a species' geographic distribution harbour limited neutral genetic diversity due to demographic properties. Moreover, we argue that populations from suitable habitats went through strong selection processes, are thus well adapted to local conditions, and therefore exhibit reduced genetic diversity at adaptive loci compared to populations at niche margins. [ABSTRACT FROM AUTHOR]

Abstract: Fungi are both agents of disease and mutualistic partners of plants. Previous studies have tested the effects of abiotic or biotic factors on plant‐associated fungal communities in isolation. However, to better understand patterns of plant–fungal associations, the combined effects of abiotic and biotic drivers across environmental gradients may be important. We investigated the effects of temperature, pH, soil moisture, vegetation cover and distance to host plant on the occurrence and abundance of fungi associated with Swiss stone pine (Pinus cembra). We did this by DNA metabarcoding 288 soil samples taken across and beyond the elevation range of P. cembra (i.e. 1,850–2,250 m a.s.l.) in two valleys in the Swiss Alps. We modelled the effects of abiotic and biotic factors on DNA read abundance of pathogenic and mutualistic fungal operational taxonomic units (OTUs) associated with P. cembra. We also tested whether abiotic and biotic factors differentially affected fungi of varying host specificity (i.e. host generalists, host specialists). We found that the occurrences of both host generalist and specialist fungi exceeded the current elevational range of their host plant. Abiotic factors had only minor effects on the abundances of all fungal OTUs. However, we found positive effects of the host plant on the abundance of a host specialist pathogenic fungus, providing support for a Janzen–Connell effect of high pathogen accumulation close to conspecific host plants. We also found a positive response to the host plant in a specialist ectomycorrhizal fungus, suggesting an “inverse” Janzen–Connell effect. Synthesis. Our findings imply that negative distance dependence shapes not only the distribution of host‐specific fungal pathogens, but also host‐specific fungal mutualists. We conclude that the occurrence of both pathogenic and mutualistic fungi beyond the current elevational range of host plants may determine their potential range shifts under projected climate warming. [ABSTRACT FROM AUTHOR]

Many animals hoard seeds for later consumption and establish seed caches that are often located at sites with specific environmental characteristics. One explanation for the selection of non-random caching locations is the avoidance of pilferage by other animals. Another possible hypothesis is that animals choose locations that hamper the perishability of stored food, allowing the consumption of unspoiled food items over long time periods., We examined seed perishability and pilferage avoidance as potential drivers for caching behaviour of spotted nutcrackers ( Nucifraga caryocatactes) in the Swiss Alps where the birds are specialized on caching seeds of Swiss stone pine ( Pinus cembra)., We used seedling establishment as an inverse measure of seed perishability, as established seedlings cannot longer be consumed by nutcrackers. We recorded the environmental conditions (i.e. canopy openness and soil moisture) of seed caching, seedling establishment and pilferage sites., Our results show that sites of seed caching and seedling establishment had opposed microenvironmental conditions. Canopy openness and soil moisture were negatively related to seed caching but positively related to seedling establishment, i.e. nutcrackers cached seeds preferentially at sites where seed perishability was low. We found no effects of environmental factors on cache pilferage, i.e. neither canopy openness nor soil moisture had significant effects on pilferage rates. We thus could not relate caching behaviour to pilferage avoidance., Our study highlights the importance of seed perishability as a mechanism for seed-caching behaviour, which should be considered in future studies. Our findings could have important implications for the regeneration of plants whose seeds are dispersed by seed-caching animals, as the potential of seedlings to establish may strongly decrease if animals cache seeds at sites that favour seed perishability rather than seedling establishment. [ABSTRACT FROM AUTHOR]

Tree-ring series from a single site and a single species were used as proxies to reconstruct past summer temperatures over 550 years in the Eastern Carpathian Mountains. The chronology was built using standard procedures in order to provide comparable information about this under-sampled region while preserving the low-frequency signals. The studied site offered abundant samples of both living trees and dead wood, which were carefully selected in order to minimize heterogeneity sources owing to a reduced availability of suitable material as e.g. in the Alps. The ring-width chronology spanned over 550 years with satisfying replication and showed an even segment length. The chronology correlated to temperature over quite a narrow window, temperatures of June and July only being significant. The reconstruction showed that the last 180 years was the warmest period with only three short episodes of anomalies. We present evidence that the summer temperatures in the Eastern Carpathian Mountains showed divergences as compared to the Alps and a clear regionality, the coolest period over the last 600 years occurring during 1720-1850. [ABSTRACT FROM AUTHOR]

In low temperature-adapted plants, including treeline trees, light-saturated photosynthesis is considerably less sensitive to temperature than growth. As a consequence, all plants tested so far show increased nonstructural carbohydrate (NSC) tissue concentrations when exposed to low temperatures. Reduced carbon supply is thus an unlikely cause for low temperature range limits of plants. For altitudinal treeline trees there is, however, a possibility that high NSC genotypes have been selected., Here, we explored this possibility using afforestations with single-provenance conifers along elevational gradients in the Southern Chilean Andes and the Swiss Alps., Tree growth was measured at each of four approximately equidistant elevations at and below the treeline. Additionally, at the same elevations, needle, branch and stem sapwood tissues were collected to determine NSC concentrations., Overall, growth decreased and NSC concentrations increased with elevation. Along with previous empirical and experimental studies, the findings of this study provide no indication of NSC reduction at the treeline; NSC increased in most species (each represented by one common population) towards their upper climatic limit. The disparity between carbon acquisition and structural carbon investment at low temperature (accumulation of NSC) thus does occur even among genotypes not adapted to treeline environments. [ABSTRACT FROM AUTHOR]

Understanding the interplay between environmental factors contributing to treeline formation and how these factors influence different life stages remains .a major research challenge. We used an afforestation experiment including 92 000 trees to investigate the spatial and temporal dynamics of tree mortality and growth at treeline in the Swiss Alps. Seedlings of three high-elevation conifer species (Larix decidua, Pinus mugo ssp. uncinata, and Pinus cembra) were systematically planted along an altitudinal gradient at and above the current treeline (2075 to 2230 m above sea level [a.s.l.]) in 1975 and closely monitored during the following 30 years. We used decision-tree models and generalized additive models to identify patterns in mortality and growth along gradients in elevation, snow duration, wind speed, and solar radiation, and to quantify interactions between the different variables. For all three species, snowmelt date was always the most important environmental factor influencing mortality, and elevation was always the most important factor for growth over the entire period studied. Individuals of all species survived at the highest point of the afforestation for more than 30 years, although mortality was greater above 2160 m a.s.l., 50-100 m above the current treeline. Optimal conditions for height growth differed from those for survival in all three species: early snowmelt (ca. day of year 125-140 [where day 1 is 1 January]) yielded lowest mortality rates, but relatively later snowmelt (ca. day 145-150) yielded highest growth rates. Although snowmelt and elevation were important throughout all life stages of the trees, the importance of radiation decreased over time and that of wind speed increased. Our findings provide experimental evidence that tree survival and height growth require different environmental conditions and that even small changes in the duration of snow cover, in addition to changes in temperature, can strongly impact tree survival and growth patterns at treeline. Further, our results show that the relative importance of different environmental variables for tree seedlings changes during the juvenile phase as they grow taller. [ABSTRACT FROM AUTHOR]

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) andbarrier 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 ( FST = 0.19) than among Alpine populations ( FST = 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. [ABSTRACT FROM AUTHOR]

1. Changes in the frequency and intensity of disturbances are expected to occur during the coming decades as a consequence of climatic changes. Mountain forests are sensitive to climate variability, disturbances and changes in human activities; this is particularly true for subalpine forests located close to the limits of tree-growth. Here we test the role of surface fires on the structure and the dynamics of a subalpine forest compared to a control stand not affected by fire events for at least two centuries. 2. The fire events are deduced from fire scars, the age-structure from tree-ring counting, regeneration from sapling and seedling counts, necromass from the volume of woody debris, and the understorey structure from shrub and herb cover, height and biomass. Land-use history is assessed from livestock and human inventories. 3. Four surface fires occurred during the last 200 years in the burned stand. Tree density is greater in the burned stand, and, even 50 years after the last fire, burning could still be having a positive effect on regeneration density. Land-use history, mainly grazing by cattle and sheep, explains the modern forest structure through its promotion of Larix decidua. The understorey composition is the same within the two stands, suggesting a rapid recovery of dwarf shrubs ( Rhododendron ferrugineum and Vaccinium myrtillus) after the latest fire, which coincided with the final change in land-use. The exception is V. vitis-idaea cover, which is higher in the control stand. The removal of livestock in the 1940s indirectly triggered a decline in L. decidua regeneration, which was formerly promoted by grazing and trampling. Conversely, Pinus cembra seedlings have begun to establish during the last 70 years. 4. Synthesis. Expansion of forests dominated by P. cembra is expected during the 21st century, unless current global warming leads to a significant and lethal increase in fire frequency, that is, a reduction of fire-return intervals, which would reduce the number of sexually mature trees. The re-introduction of grazing could maintain the mixed subalpine forests, whereas surface fires would alter the woody debris load, promoting greater regeneration of L. decidua and P. cembra. [ABSTRACT FROM AUTHOR]

It has been predicted that global climate change will lead to increasing drought in the Alps during the 21st century, as well as an increased fire risk, fires being currently rare in these mountains. Herein we describe fire frequency reconstruction using high-resolution analyses of macroscopic sedimentary charcoal, pollen, and plant macrofossils. Sediments were sampled from a subalpine pond within the dry western French Alps. Results show that the early-Holocene expansion of Pinus cembra (7200 calibrated years BP) occurred in Acer/Alnus incana/Betula woodlands, which were affected by fires with moderate mean fire-free intervals (MFFI = 173 ± 61 yr [mean ± SE]). Superposed Epoch Analyses show that the abundance of P. cembra macroremains decreased significantly after burning, although they never disappeared entirely. Statistics suggest that fires spread through cembra pine communities; they were not stand-replacing fires but mainly surface fires, probably killing nonreproductive pines. An increase in fire frequency occurred 6740 years ago, when four fires appear to have occurred within 140 years. These fires may have been associated with a regional drought and could have affected the composition of the subalpine forest by depleting the local P. cembra population in the short term. The predicted increase in drought in the future could, therefore, affect the cembra pine ecosystem in the Alps if fire frequency is reduced to intervals of less than 80 years. [ABSTRACT FROM AUTHOR]

Questions: What is the structure of the anthropogenic upper forest-grassland ecotone and are there differences in the spatial relationships between the tree species involved? Location: Valfurva Valley, Italian central Alps. Methods: We conducted a spatial distribution and structure analysis in three 1-ha permanent plots along an altitudinal gradient, from the treeline to the sub-alpine forest. We reconstructed the age structure from cores from each individual with diameter > 4 cm at 50 cm height. Results: All tree species and age classes examined had a clumped structure. The cluster tendency was more evident at the treeline where the environmental conditions are more severe. In the sub-alpine forest there was a repulsion between Pinus cembra and Pious mugo but at the treeline P. cembra was frequently found downslope from P. mugo. Conclusions: Although human influence has been the main driving force in shaping the present forest structure, in the last few decades natural dynamics have become the predominant force acting on forest structure and processes, showing a higher magnitude as altitude increases. Our results emphasize the existence of facilitating and interfering mechanisms between different species. P. cembra seems to be favoured compared to the other tree species. [ABSTRACT FROM AUTHOR]

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

In this study, charcoal-based data for Pinus cembra L . (arolla pine) were gathered from soil and travertine sequences in order to reconstruct its historical biogeography at the landscape level in the inner western Alps during the Holocene. The study sites are located between 1700 and 2990 m a.s.l., in the southern (Queyras Massif and Ubaye Valley) and the northern (Maurienne Valley) parts of the inner French Alps. Charcoal fragments were extracted from sediments by water sieving, using meshes of 5, 2, 0.8 and 0.4 mm. The charcoal mass of P. cembra was determined in each charcoal assemblage. Accelerator mass spectrometry and conventional 14C measurements were used to date the fragments. Supported by 40 14C datings, the fragments show that, over 2000 m a.s.l., P. cembra accounts for around 40% (mean value) of identified fragments. Data reveal that arolla pine once extended between 260 and 375 m above the present-day local tree lines. It was established in the southern and the northern French Alps from at least c. 9000 and 6000 cal yr bp, respectively. While present-day populations of P. cembra are very fragmented in the inner French Alps, charcoal records indicate large past occurrences of this tree since the early Holocene. Human disturbance since the Neolithic seems to be the main reason for the regression of the arolla pine woodlands. On the south-facing slopes of the study sites, currently deforested, this species extended up to 2800 m a.s.l. In the northern areas, charcoal records of the P. cembra expansion are consistent with the regional pollen archives, but in the southern massifs charcoal records indicate its presence c. 2600 years earlier than other palaeobotanical studies suggest. This discrepancy highlights the necessity to crosscheck data using several different proxies in order to assess the validity of conclusions regarding tree development in space and time. [ABSTRACT FROM AUTHOR]