Additional informationReprints and permissions information is available online at www.nature.com/reprints. Publisher's note: Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. Competing financial interestsThe authors declare no competing financial interests. Europe PMC Funders GroupAuthor Manuscript Nat Clim Chang. Author manuscript; available in PMC 2017 December 01. Forest disturbances are sensitive to climate. However, our understanding of disturbance dynamics in response to climatic changes remains incomplete, particularly regarding large-scale patterns, interaction effects and dampening feedbacks. Here we provide a global synthesis of climate change effects on important abiotic (fire, drought, wind, snow and ice) and biotic (insects and pathogens) disturbance agents. Warmer and drier conditions particularly facilitate fire, drought and insect disturbances, while warmer and wetter conditions increase disturbances from wind and pathogens. Widespread interactions between agents are likely to amplify disturbances, while indirect climate effects such as vegetation changes can dampen long-term disturbance sensitivities to climate. Future changes in disturbance are likely to be most pronounced in coniferous forests and the boreal biome. We conclude that both ecosystems and society should be prepared for an increasingly disturbed future of forests.Natural disturbances, such as fires, insect outbreaks and windthrows, are an integral part of ecosystem dynamics in forests around the globe. They occur as relatively discrete events, and form characteristic regimes of typical disturbance frequencies, sizes and severities over extended spatial and temporal scales1,2. Disturbances disrupt the structure, composition and function of an ecosystem, community or population, and change resource availability or the physical environment3. In doing so, they create heterogeneity on the landscape4, foster diversity across a wide range of guilds and species5,6 and initiate ecosystem renewal or reorganization7,8.Disturbance regimes have changed profoundly in many forest ecosystems in recent years, with climate being a prominent driver of disturbance change9. An increase in disturbance occurrence and severity has been documented over large parts of the globe, for example, for fire10,11, insect outbreaks12,13 and drought14,15. Such alterations of disturbance regimes have the potential to strongly impact the ability of forests to provide ecosystem services to society6. Moreover, a climate-mediated increase in disturbances could exceed the ecological resilience of forests, resulting in lastingly altered ecosystems or shifts to non-forest ecosystems as tipping points are crossed16-18. Consequently, disturbance change is expected to be among the most profound impacts that climate change will have on forest ecosystems in the coming decades19.The ongoing changes in disturbance regimes in combination with their strong and lasting impacts on ecosystems have led to an in...

Climate warming is causing a shift in biological communities in favor of warm-affinity species (i.e., thermophilization). Species responses often lag behind climate warming, but the reasons for such lags remain largely unknown. Here, we analyzed multidecadal understory microclimate dynamics in European forests and show that thermophilization and the climatic lag in forest plant communities are primarily controlled by microclimate. Increasing tree canopy cover reduces warming rates inside forests, but loss of canopy cover leads to increased local heat that exacerbates the disequilibrium between community responses and climate change. Reciprocal effects between plants and microclimates are key to understanding the response of forest biodiversity and functioning to climate and land-use changes.

Much of our understanding of natural forest dynamics in the temperate region of Europe is based on observational studies in old-growth remnants that have emphasized small-scale gap dynamics and equilibrium stand structure and composition. Relatively little attention has been given to the role of infrequent disturbance events in forest dynamics. In this study, we analyzed dendroecological data from four stands and three windthrow patches in an old-growth landscape in the Dinaric Mountains of Bosnia and Herzegovina to examine disturbance history, tree life history traits, and compositional dynamics. Over all stands, most decades during the past 340 years experienced less than 10% canopy loss, yet each stand showed evidence of periodic intermediate-severity disturbances that removed > 40% of the canopy, some of which were synchronized over the study area landscape. Analysis of radial growth patterns indicated several life history differences among the dominant canopy trees; beech was markedly older than fir, while growth patterns of dead and dying trees suggested that fir was able to tolerate longer periods of suppressed growth in shade. Maple had the fastest radial growth and accessed the canopy primarily through rapid early growth in canopy gaps, whereas most beech and fir experienced a period of suppressed growth prior to canopy accession. Peaks in disturbance were roughly linked to increased recruitment, but mainly of shade-tolerant beech and fir; less tolerant species (i.e., maple, ash, and elm) recruited successfully on some of the windthown sites where advance regeneration of beech and fir was less abundant. The results challenge the traditional notions of stability in temperate old-growth forests of Europe and highlight the nonequilibrial nature of canopy composition due to unique histories of disturbance and tree life history differences. These findings provide valuable information for developing natural disturbance-based silvicultural systems, as well as insight into maintaining less shade-tolerant, but valuable broadleaved trees in temperate forests of Europe.

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Mountain forests are among the most important ecosystems in Europe as they support numerous ecological, hydrological, climatic, social, and economic functions. They are unique relatively natural ecosystems consisting of long-lived species in an otherwise densely populated human landscape. Despite this, centuries of intensive forest management in many of these forests have eclipsed evidence of natural processes, especially the role of disturbances in long-term forest dynamics. Recent trends of land abandonment and establishment of protected forests have coincided with a growing interest in managing forests in more natural states. At the same time, the importance of past disturbances highlighted in an emerging body of literature, and recent increasing disturbances due to climate change are challenging long-held views of dynamics in these ecosystems. Here, we synthesize aspects of this Special Issue on the ecology of mountain forest ecosystems in Europe in the context of broader discussions in the field, to present a new perspective on these ecosystems and their natural disturbance regimes. Most mountain forests in Europe, for which long-term data are available, show a strong and long-term effect of not only human land use but also of natural disturbances that vary by orders of magnitude in size and Europe PMC Funders Author ManuscriptsEurope PMC Funders Author Manuscripts frequency. Although these disturbances may kill many trees, the forests themselves have not been threatened. The relative importance of natural disturbances, land use, and climate change for ecosystem dynamics varies across space and time. Across the continent, changing climate and land use are altering forest cover, forest structure, tree demography, and natural disturbances, including fires, insect outbreaks, avalanches, and wind disturbances. Projected continued increases in forest area and biomass along with continued warming are likely to further promote forest disturbances. Episodic disturbances may foster ecosystem adaptation to the effects of ongoing and future climatic change. Increasing disturbances, along with trends of less intense land use, will promote further increases in coarse woody debris, with cascading positive effects on biodiversity, edaphic conditions, biogeochemical cycles, and increased heterogeneity across a range of spatial scales. Together, this may translate to disturbance-mediated resilience of forest landscapes and increased biodiversity, as long as climate and disturbance regimes remain within the tolerance of relevant species. Understanding ecological variability, even imperfectly, is integral to anticipating vulnerabilities and promoting ecological resilience, especially under growing uncertainty. Allowing some forests to be shaped by natural processes may be congruent with multiple goals of forest management, even in densely settled and developed countries.

Biodiversity time series reveal global losses and accelerated redistributions of species, yet no net loss in local species richness. To better understand how these patterns are linked, we quantify how individual species trajectories scale up to diversity changes using data from 68 vegetation resurvey studies of seminatural forests in Europe. Herb-layer species with small geographic ranges are being replaced by more widely distributed species and our results suggest this is less due to species abundances than to species nitrogen (N) niches. N-deposition accelerates extinctions of small-ranged, N-efficient plants and colonization by broadly distributed, N-demanding plants including non-natives. Despite no net change in species richness at the spatial scale of a study site, losses of small-ranged species reduce biome-scale (gamma) diversity. These results provide one mechanism to explain the directional replacement of smallranged species within sites and thus patterns of biodiversity change across spatial scales.

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