Chemical signature of Eurois occulta L. outbreaks in the xylem cell wall of Salix glauca L. in Greenland.

Insect defoliations are a major natural disturbance in high-latitude ecosystems and are expected to increase in frequency and severity due to current climatic change. Defoliations cause severe reductions in biomass and carbon investments that affect the functioning and productivity of tundra ecosystems. Here we combined dendro-anatomical analysis with chemical imaging to investigate the direct and lagged effects of insect outbreaks on carbon investment. We analysed the content of lignin vs. holocellulose, i.e. unspecified carbohydrates in xylem samples of Salix glauca L. collected at Iffiartarfik, Nuuk fjord, Greenland, featuring two outbreak events of the moth Eurois occulta L. Cross sections of the growth rings corresponding to both outbreaks ±3 years were analysed using confocal Raman imaging to identify possible chemical signatures related to insect defoliation on fibres, vessels, and ray parenchyma cells and to get insight into species-specific defence responses. Outbreak years with narrower rings and thinner fibre cell walls are accompanied by a change in the content of cell-wall polymers but not their underlying chemistry. Indeed, during the outbreaks the ratio between lignin and carbohydrates significantly increased in fibre but not vessel cell walls due to an increase in lignin content coupled with a reduced content of carbohydrates. Parenchyma cell walls and cell corners did not show any significant changes in the cell-wall biopolymer content. The selective adjustment of the cell-wall composition of fibres but not vessels under stressful conditions could be related to the plants priority to maintain an efficient hydraulic system rather than mechanical support. However, the higher lignin content of fibre cell walls formed during the outbreak events could increase mechanical stiffness to the thin walls by optimizing the available resources. Chemical analysis of xylem traits with Raman imaging is a promising approach to highlight hidden effects of defoliation otherwise overlooked with classical dendroecological methods.

Winter precipitation - not summer temperature - is still the main driver for Alpine shrub growth.

High latitude and altitude environments are universally recognized as particularly sensitive to environmental changes and the current climate warming is inducing remarkable transformations on vegetation assemblage in these temperature-limited regions. However, next to the wealth of studies describing the effect of rising growing season temperature on trees, much less is known about the concurrent effects of precipitation and snowpack dynamics on the other key component of alpine vegetation represented by prostrate life forms. Selecting the most widespread shrub species in the North Hemisphere, we assembled a monospecific (Juniperus communis L.) network of 7 sites overarching the European Alps, measured the annual growth on >330 individuals and assessed the climate-growth associations for the last century (1910-2010) adopting a new model estimating the solid fraction of precipitation from unique highly-resolved daily climate records. Despite the high space-time variability of the yearly precipitation amount and distribution across the region, our analysis found a prominent, consistent and negative role of winter precipitation for shrub growth. Moreover, this crucial role of snow is maintained even in recent years, despite the persistent and significant warming trend. The presence of this underrated key factor for Alpine long-lived vegetation will require a thorough consideration. For the prostrate life form, not only temperature but also the solid fraction of winter precipitation should be considered to improve the projections of future growth trajectories.

Diverging shrub and tree growth from the Polar to the Mediterranean biomes across the European continent.

Climate warming is expected to enhance productivity and growth of woody plants, particularly in temperature-limited environments at the northernmost or uppermost limits of their distribution. However, this warming is spatially uneven and temporally variable, and the rise in temperatures differently affects biomes and growth forms. Here, applying a dendroecological approach with generalized additive mixed models, we analysed how the growth of shrubby junipers and coexisting trees (larch and pine species) responds to rising temperatures along a 5000-km latitudinal range including sites from the Polar, Alpine to the Mediterranean biomes. We hypothesize that, being more coupled to ground microclimate, junipers will be less influenced by atmospheric conditions and will less respond to the post-1950 climate warming than coexisting standing trees. Unexpectedly, shrub and tree growth forms revealed divergent growth trends in all the three biomes, with juniper performing better than trees at Mediterranean than at Polar and Alpine sites. The post-1980s decline of tree growth in Mediterranean sites might be induced by drought stress amplified by climate warming and did not affect junipers. We conclude that different but coexisting long-living growth forms can respond differently to the same climate factor and that, even in temperature-limited area, other drivers like the duration of snow cover might locally play a fundamental role on woody plants growth across Europe.