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Carbon allocation plays a key role in ecosystem dynamics and plant adaptation to changing environmental conditions. Hence, proper description of this process in vegetation models is crucial for the simulations of the impact of climate change on carbon cycling in forests. Here we review how carbon allocation modelling is currently implemented in 31 contrasting models to identify the main gaps compared with our theoretical and empirical understanding of carbon allocation. A hybrid approach based on combining several principles and/or types of carbon allocation modelling prevailed in the examined models, while physiologically more sophisticated approaches were used less often than empirical ones. The analysis revealed that, although the number of carbon allocation studies over the past 10 years has substantially increased, some background processes are still insufficiently understood and some issues in models are frequently poorly represented, oversimplified or even omitted. Hence, current challenges for carbon allocation modelling in forest ecosystems are (i) to overcome remaining limits in process understanding, particularly regarding the impact of disturbances on carbon allocation, accumulation and utilization of nonstructural carbohydrates, and carbon use by symbionts, and (ii) to implement existing knowledge of carbon allocation into defence, regeneration and improved resource uptake in order to better account for changing environmental conditions., (© The Author(s) 2019. Published by Oxford University Press.)

Climate teleconnections drive highly variable and synchronous seed production (masting) over large scales. Disentangling the effect of high-frequency (inter-annual variation) from low-frequency (decadal trends) components of climate oscillations will improve our understanding of masting as an ecosystem process. Using century-long observations on masting (the MASTREE database) and data on the Northern Atlantic Oscillation (NAO), we show that in the last 60 years both high-frequency summer and spring NAO, and low-frequency winter NAO components are highly correlated to continent-wide masting in European beech and Norway spruce. Relationships are weaker (non-stationary) in the early twentieth century. This finding improves our understanding on how climate variation affects large-scale synchronization of tree masting. Moreover, it supports the connection between proximate and ultimate causes of masting: indeed, large-scale features of atmospheric circulation coherently drive cues and resources for masting, as well as its evolutionary drivers, such as pollination efficiency, abundance of seed dispersers, and natural disturbance regimes.

European beech (Fagus sylvatica L.) can regenerate successfully from seeds after mixed-severity fires with mid-to-long fire return intervals (>60 years). However, if fire return interval is lower than the age of sexual maturity, post-fire seeding will be limited, leaving vegetative resprouting as the only viable option for recovery. This means that the forecasted increase in fire frequency driven by climate change may erode beech forest resilience to fire. Here, we surveyed tree regeneration in a European beech forest affected by two consecutive fires, in 2003 and 2017, and applied experimental clipping of tree saplings to address the following questions: (1) What is the fire resistance and post-fire recovery via resprouting of tree saplings? (2) Which factors drive post-fire resprouting of beech saplings? (3) Does post-fire clipping of tree saplings increase the probability of survival and resprouting vigor? We monitored 2195 beech saplings and 953 saplings of other tree species during three consecutive years, from 2018 to 2020. Almost all beech saplings were top-killed by fire, and two-thirds of them died completely. However, 3 years after the second fire, 30 per cent of beech saplings survived by resprouting from the base. Post-fire resprouting was less likely in small-diameter saplings and in those more injured by fire. Overall, the second fire did not cause a major decline of beech regeneration and consequently did not alter the dominant species composition of post-fire recovery. Given the low specific resistance to fire, post-fire resprouting of saplings is therefore a key component of beech resilience to short-interval fires. The effects of clipping on post-fire survival and resprouting vigor were very limited, suggesting the unsuitability of actively clearing burned beech regeneration as a post-fire management prescription. In conclusion, basal resprouting from beech saplings after fire-induced top-kill led to a higher-than-expected resilience of beech to short-interval fires (i.e. circa 15 years). [ABSTRACT FROM AUTHOR]

Drought will increasingly threaten forest ecosystems worldwide. Understanding how competition influences tree growth response to drought is essential for forest management aiming at climate change adaptation. However, published results from individual case studies are heterogeneous and sometimes contradictory. We reviewed 166 cases from the peer-reviewed literature to assess the influence of stand-level competition on tree growth response to drought. We monitored five indicators of tree growth response: mean sensitivity (inter-annual tree ring width variability); association between inter-annual growth variability and water availability; resistance; recovery; and resilience to drought. Vote counting did not indicate a consistent effect of competition on mean sensitivity. Conversely, higher competition for resources strengthened the association between water availability and inter-annual growth rates. Meta-analysis showed that higher competition reduced resistance (p < 0.001) and improved recovery (p < 0.05), but did not consistently affect resilience. Species, site and stand characteristics, and drought intensity were insignificant or poor predictors for the large variability among the investigated cases. Our review and meta-analysis show that competition does not affect the response of tree growth to drought in a unidirectional and universal way. Although density reduction (thinning) can alleviate growth declines during drought, the effects on growth after stress are uncertain. The large variability among investigated cases suggests that local-scale processes play a crucial role in determining such responses and should be explicitly evaluated and integrated into specific strategies for adaptation of forests to climate change. [ABSTRACT FROM AUTHOR]

There is evidence that variable and synchronous reproduction in seed plants (masting) correlates to modes of climate variability, e.g. El Niño Southern Oscillation and North Atlantic Oscillation. In this perspective, we explore the breadth of knowledge on how climate modes control reproduction in major masting species throughout Earth's biomes. We posit that intrinsic properties of climate modes (periodicity, persistence and trends) drive interannual and decadal variability of plant reproduction, as well as the spatial extent of its synchrony, aligning multiple proximate causes of masting through space and time. Moreover, climate modes force lagged but in-phase ecological processes that interact synergistically with multiple stages of plant reproductive cycles. This sets up adaptive benefits by increasing offspring fitness through either economies of scale or environmental prediction. Community-wide links between climate modes and masting across plant taxa suggest an evolutionary role of climate variability. We argue that climate modes may 'bridge' proximate and ultimate causes of masting selecting for variable and synchronous reproduction. The future of such interaction is uncertain: processes that improve reproductive fitness may remain coupled with climate modes even under changing climates, but chances are that abrupt global warming will affect Earth's climate modes so rapidly as to alter ecological and evolutionary links. This article is part of the theme issue 'The ecology and evolution of synchronized seed production in plants'. [ABSTRACT FROM AUTHOR]

Populations of many long-lived plants exhibit spatially synchronized seed production that varies extensively over time, so that seed production in some years is much higher than on average, while in others, it is much lower or absent. This phenomenon termed masting or mast seeding has important consequences for plant reproductive success, ecosystem dynamics and plant–human interactions. Inspired by recent advances in the field, this special issue presents a series of articles that advance the current understanding of the ecology and evolution of masting. To provide a broad overview, we reflect on the state-of-the-art of masting research in terms of underlying proximate mechanisms, ontogeny, adaptations, phylogeny and applications to conservation. While the mechanistic drivers and fitness consequences of masting have received most attention, the evolutionary history, ontogenetic trajectory and applications to plant–human interactions are poorly understood. With increased availability of long-term datasets across broader geographical and taxonomic scales, as well as advances in molecular approaches, we expect that many mysteries of masting will be solved soon. The increased understanding of this global phenomenon will provide the foundation for predictive modelling of seed crops, which will improve our ability to manage forests and agricultural fruit and nut crops in the Anthropocene. This article is part of the theme issue 'The ecology and evolution of synchronized seed production in plants'. [ABSTRACT FROM AUTHOR]

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Climate change may reduce forest growth and increase forest mortality, which is connected to high carbon costs through reductions in gross primary production and net ecosystem exchange. Yet, the spatiotemporal patterns of vulnerability to both short‐term extreme events and gradual environmental changes are quite uncertain across the species' limits of tolerance to dryness. Such information is fundamental for defining ecologically relevant upper limits of species tolerance to drought and, hence, to predict the risk of increased forest mortality and shifts in species composition. We investigate here to what extent the impact of short‐ and long‐term environmental changes determines vulnerability to climate change of three evergreen conifers (Scots pine, silver fir, Norway spruce) and two deciduous hardwoods (European beech, sessile oak) tree species at their southernmost limits of distribution in the Mediterranean Basin. Finally, we simulated future forest growth under RCP 2.6 and 8.5 emission scenarios using a multispecies generalized linear mixed model. Our analysis provides four key insights into the patterns of species' vulnerability to climate change. First, site climatic marginality was significantly linked to the growth trends: increasing growth was related to less climatically limited sites. Second, estimated species‐specific vulnerability did not match their a priori rank in drought tolerance: Scots pine and beech seem to be the most vulnerable species among those studied despite their contrasting physiologies. Third, adaptation to site conditions prevails over species‐specific determinism in forest response to climate change. And fourth, regional differences in forests vulnerability to climate change across the Mediterranean Basin are linked to the influence of summer atmospheric circulation patterns, which are not correctly represented in global climate models. Thus, projections of forest performance should reconsider the traditional classification of tree species in functional types and critically evaluate the fine‐scale limitations of the climate data generated by global climate models. Current and forecasted trends in tree growth from temperate and boreal tree species growing at the southernmost limit of their natural distribution in Europe. The tree growth projections by 2100 revealed a generalized decrease in growth under the climatic conditions derived from the RCP 8.5 emission scenario. [ABSTRACT FROM AUTHOR]

Human activities have the potential to enhance carbon sequestration by the world's forests and contribute to climate change mitigation. Voluntary carbon trading is currently the only option to pursue and reward carbon sequestration by forestry activities. Carbon credits for enhanced sequestration can be sold to partners wishing to offset their own emissions. Here we illustrate the steps taken to design guidelines for the generation of voluntary carbon credits by improved forest management in Piemonte, Italy. The guidelines have been developed in a joint effort by academia, regional administrations, forest owners and professional consultants. In particular, we show how to compute the baseline and the additionality of credit-generating forest management activities, and how to reconcile the generation of forest carbon credits with law requirements, technical limitations, and the provision of other ecosystem services. To illustrate the profitability of carbon credit generation, we simulated the application of carbon credit guidelines to two forest-rich mountain watersheds in the southern part of the Piemonte region. The two dominating tree species are beech (Fagus sylvatica L.) and chestnut (Castanea sativa Mill.). We computed current forest carbon stock and carbon credits generated in 20 years under business as usual and an alternative biomass retention scenario. The IFM resulted in an avoided harvest of 39,362 m3 for a net total of 64,014 MgCO2e after subtracting harvest emissions, or 38 Mg ha-1 throughout the permanence period of 20 years. These steps can be replicated in other mountain regions where there is interest in promoting this ecosystem service as an alternative or an addition to production-oriented forest management. [ABSTRACT FROM AUTHOR]

Forest fires in Italy are changing. More frequent heatwaves and drought increase the flammability of the vegetation; the abandonment of rural land produces 30.000 ha of newly afforested areas each year; and the wildland-urban interface is expanding with the sprawl of urbanized areas. However, forest fires are rarely understood and managed in their complexity. The public opinion is often misinformed on the causes and consequences of fires in the forest. Moreover, fire management relies almost exclusively on extinction and emergency response, resulting in high costs and limited efficacy versus extreme fire seasons. We advocate to increase the role and investments in wildfire prevention, which can be carried out by fuel-oriented silviculture, such as facilitating less flammable species or prescribed burning, in order to reduce the flammability of the vegetation and mitigate fire intensi - ty in high-leverage areas. A centralized structure is necessary to implement such a strategy and coordinate the competences and actions of all local administrations and actors involved. [ABSTRACT FROM AUTHOR]

Forests of the Apennines are characterised by high canopy cover and high tree species diversity (being at the interface between two major climatic zones of Europe), and provide important ecosystem functions to millions of people. They exemplify cutting-edge themes such as forest ecology in warmer climates, consequences of heavy land use, and resilience at the trailing edge of the distribution of many European forest species (Silver fir, Norway spruce, Beech, Black pine, Birch). We introduce the setting under the geological and climatological point of view and review the literature on the interactions between these long-term drivers and the specific, structural, and genetic diversity of these forest communities (e.g., effects of glacial refugia or tectonic/volcanic activity), followed by a brief outline of what little is known about natural disturbance regimes and their range of variability. Anthropogenic disturbances (fire, grazing) and land use changes (abandonment of cropland and pasture) have been by far the main drivers of forest dynamics at least for the last two millennia, determining for examples overageing of coppices, treeline advances, forest encroachment on former agricultural land. We suggest considerations about the interplay between these land use changes and disturbance drivers (e.g. fuel continuity), summarize comparisons between managed and unmanaged forests (e.g., increase in tree size, deadwood, biodiversity indicators), and elaborate on current proposals for climate-adapted management, highlighting specific and genetic diversity as an important source of resilience and adaptive potential. [ABSTRACT FROM AUTHOR]

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 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. [ABSTRACT FROM AUTHOR]

An increasing number of studies have reported on forest declines and vegetation shifts triggered by drought. In the Swiss Rhone valley (Valais), one of the driest inner-Alpine regions, the species composition in low elevation forests is changing: The sub-boreal Scots pine ( Pinus sylvestris L.) dominating the dry forests is showing high mortality rates. Concurrently the sub-Mediterranean pubescent oak ( Quercus pubescens Willd.) has locally increased in abundance. However, it remains unclear whether this local change in species composition is part of a larger-scale vegetation shift. To study variability in mortality and regeneration in these dry forests we analysed data from the Swiss national forest inventory ( NFI) on a regular grid between 1983 and 2003, and combined it with annual mortality data from a monitoring site. Pine mortality was found to be highest at low elevation (below 1000 m a.s.l.). Annual variation in pine mortality was correlated with a drought index computed for the summer months prior to observed tree death. A generalized linear mixed-effects model indicated for the NFI data increased pine mortality on dryer sites with high stand competition, particularly for small-diameter trees. Pine regeneration was low in comparison to its occurrence in the overstorey, whereas oak regeneration was comparably abundant. Although both species regenerated well at dry sites, pine regeneration was favoured at cooler sites at higher altitude and oak regeneration was more frequent at warmer sites, indicating a higher adaptation potential of oaks under future warming. Our results thus suggest that an extended shift in species composition is actually occurring in the pine forests in the Valais. The main driving factors are found to be climatic variability, particularly drought, and variability in stand structure and topography. Thus, pine forests at low elevations are developing into oak forests with unknown consequences for these ecosystems and their goods and services. [ABSTRACT FROM AUTHOR]