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1.

Key message

Comparisons between compression and opposite wood formation in prostrating Pinus mugo indicate that the secondary meristem can produce more tracheids with thicker walls by also increasing the number of contemporaneously differentiating cells, rather than only increasing the duration or the rate of cell formation.

Context

Although cambium tissues within a stem experience the same climatic conditions, the resulting wood structure and properties can strongly differ. Assessing how meristem differently regulates wood formation to achieve different anatomical properties can help understanding the mechanisms of response and their plasticity.

Aims

We monitored the formation of compression (CW) and opposite (OW) wood within the same stems to understand whether achieved differences in wood structure are caused by modifications in the process of cell formation.

Methods

We collected weekly microcores of compression and opposite wood from the curved stem of ten treeline prostrating mountain pines (Pinus mugo Turra ssp. mugo) at the Majella massif in Central Italy.

Results

Results indicate that cambium formed approximately 1.5 times more cells in CW than OW, despite that CW cell differentiation only extended 2 weeks longer and the residence time of CW cells in the wall-thickening phase was only 20% longer. Differences in their formation were thus mainly related to both the rates and the width of the enlarging and wall-thickening zones (i.e., the number of cells simultaneously under differentiation) and less to duration of cell formation.

Conclusion

We conclude that to achieve such a different wood structures, the efficiency of the secondary meristem, in addition of altered rate of cell division and differentiation, can also modify the width of the developing zones. Thus, deciphering what rules this width is important to link environmental conditions with productivity.
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2.
3.

? Key message

Long-term strict protection of woodland communities may lead to their compositional simplification and homogenisation.

? Context

In the past, it has often been postulated that structures and processes typical for natural forests should be mimicked by silvicultural activities in the case of managed tree stands.

? Aims

To determine which features and traits of natural woodland communities (alongside typical old-growth attributes) should be imitated in managed forests, as well as which should not (and for what reasons).

? Methods

Tree data from five permanent study plots (of a total area of 15.44 ha) established in 1936 in the core area of the Bia?owie?a National Park (NE Poland) are used to calculate several quantitative indices describing the temporal dynamics (in terms of stand structure and composition) of eight major woodland community types.

? Results

Most structural attributes revealed rather high stability over time. In contrast to these, during the observation period, noticeable changes in the composition of particular Bia?owie?a woodland communities have been taking place, related to declining occurrence and reduced roles characteristic for a large number of tree species.

? Conclusion

In many ways, natural forests can serve as an important model for managed forest stands. However, in certain circumstances, silvicultural treatments counteracting natural developmental trends may appear to be indispensable, especially when more diverse and stable tree species composition (at a given spatial and temporal scale) is indicated or desirable.
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4.

Key message

Multiple lines of evidence suggest acoustic wave velocity (AWV) would provide a rapid and efficient method to indirectly select for superior pulp yield in Eucalyptus globulus breeding programs.

Context

Eucalyptus globulus is one of the most widely planted hardwood species in temperate regions of the world and is primarily grown for pulpwood.

Aims

To determine if acoustic wave velocity (AWV) can be used to indirectly select for kraft pulp yield in E. globulus.

Methods

Genetic group effects, additive and non-additive variance components, and genetic correlations were estimated for AWV and pulpwood traits, including Kraft pulp yield. In a separate trial, the relative position of quantitative trait loci (QTL) for these traits was compared.

Results

Estimated narrow-sense heritabilities for AWV and pulp yield were both 0.26, and these traits were strongly genetically correlated (0.84). Furthermore, co-located QTL for these traits were identified. Further evidence that AWV could be used to indirectly select for pulp yield was provided by the ranking of genetic groups—Otways and King Island had the highest AWV and pulp yield and Strzelecki and Tasmania the lowest. There was no evidence of dominance variation in wood property traits.

Conclusion

Together, these findings suggest that AWV could be used as a selection criterion for kraft pulp yield in E. globulus breeding programs.
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5.

Key message The application of the ITOC model allows the estimation of available biomass potentials from forests on the basis of National Forest Inventory data. The adaptation of the model to country-specific situations gives the possibility to further enhance the model calculations.

Context

With the rising demand for energy from renewable sources, up-to-date information about the available amount of biomass on a sustainable basis coming from forests became of interest to a wide group of stakeholders. The complexity of answering the question about amounts of biomass potentials from forests thereby increases from the regional to the European level.

Aims

The described ITOC model aims at providing a tool to develop a comparable data basis for the actual biomass potentials for consumption.

Methods

The ITOC model uses a harmonized net annual increment from the National Forest Inventories as a default value for the potential harvestable volume of timber. The model then calculates the total theoretical potential of biomass resources from forests. By accounting for harvesting restrictions and losses, the theoretical potential of biomass resources from forests is reduced and the actual biomass potentials for consumption estimated.

Results

The results from ITOC model calculations account for the difference between the amounts of wood measured in the forests and the actual biomass potentials which might be available for consumption under the model assumptions.

Conclusion

The gap between forest resource assessments and biomass potentials which are available for consumption can be addressed by using the ITOC model calculation results.
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6.

Context and aim

This study aimed to examine the effect of the tension wood G‐layer on the viscoelastic properties of wood.

Methods

Tension wood and opposite wood samples were obtained from six French Guianese tropical rainforest species (Sextonia rubra, Ocotea guyanensis, Inga alba, Tachigali melinoni, Iyranthera sagotiana and Virola michelii); the tension wood of the former three of these species had a G‐layer, whilst the tension wood from the latter three had no G‐layer. Tensile dynamic mechanical analysis (DMA) was performed on green never dried wood samples in the longitudinal direction with samples submerged in a water bath at a temperature (30°C) and frequency (1 Hz) representative of the conditions experienced by wood within a living tree. Then, DMA was repeated with samples conditioned to an air-dried state. Finally, samples were oven-dried to measure longitudinal shrinkage.

Results

Tension wood did not always have a higher longitudinal storage (elastic) modulus than opposite wood from the same tree regardless of the presence or absence of a G‐layer. For the species containing a G‐layer, tension wood had a higher damping coefficient and experienced a greater longitudinal shrinkage upon drying than opposite wood from the same species. No difference was found in damping coefficients between tension wood and opposite wood for the species that had no G‐layer.

Conclusion

It is proposed that the different molecular composition of the G-layer matrix has an influence on the viscoelasticity of wood, even if a biomechanical gain is not yet clear. This study shows that rheological properties and longitudinal shrinkage can be used to detect the presence of a G‐layer in tension wood.
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7.

? Key message

New types of distribution functions are needed to model the dynamics of stands where important age classes are represented by few trees. In this study the gamma shape mixture model and two simulation methods were used for generating tree diameter data.

? Context

To analyse forest dynamics, it is necessary to know distribution of the characteristics (mainly tree diameters) of trees forming particular developmental phases. In many forest inventories, the measurement of large diameter at breast height (DBH) samples is practically impossible. In this case, DBH distributions can be generated using theoretical models.

? Aims

The aim of this study was to assess the precision of the approximation of empirical DBH data using the gamma shape mixture (GSM) model and kernel density estimation. The strengths and weaknesses of the two simulation methods were presented and discussed.

? Methods

The GSM model was adopted to approximate empirical DBH data collected in 20 near-natural stands. Two simulation methods were used: (a) the procedure based on a multimodal distribution and gamma random numbers (MDGR procedure) and (b) MCMC techniques with Metropolis–Hastings sampling (MH method).

? Results

The GSM model precisely fitted the investigated DBH distributions. The MDGR procedure was slightly more precise than the MH method, especially in the case of the samples of 250 DBHs. The level of homogeneity within the drawn DBH sets was similar for all samples.

? Conclusion

The GSM model is very flexible. The DBH random variates, generated with the use of analysed procedures, represented all tree generations being significant from a biological point of view.
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8.

? Context

Physiological ecologists have been fascinated by height- or position-linked differences of leaf morphology within tall trees >25 m, but the exact cause is still debated, i.e., is it due to light or height-induced water stress?

? Aims

The aim of this study was to demonstrate that relatively small trees (<15 m) have leaf morphologies that vary with height and that such variation depends on site-moisture variability.

? Methods

Leaves were collected from Robinia pseudoacacia trees at two sites in China with contrasting moisture variability to gather baseline data on leaf morphology parameters.

? Results

Most measured parameters changed regularly with height. Water potential linearly decreased with height. Leaf area and stomata area decreased with height, while leaf mass per area, carbon isotope composition (δ 13C), and stomata density increased with height. Mesophyll and epidermal cell width decreased with height, while leaf thickness and palisade cell length increased with height. All the morphology parameters between two sites were also significantly different.

? Conclusions

Based on the field results, it is concluded that minor variations in water potential at the time of leaf growth influence leaf morphology at both site-level and height-level. Controlled environment experiments will be conducted to confirm this conclusion.
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9.

Key message

Recent growth changes (1980–2007) in Western European forests strongly vary across tree species, and range from +42% in mountain contexts to ?17% in Mediterranean contexts. These changes reveal recent climate warming footprint and are structured by species' temperature (?) and precipitation (+) growing conditions.

Context

Unprecedented climate warming impacts forests extensively, questioning the respective roles of climatic habitats and tree species in forest growth responses. National forest inventories ensure a repeated and spatially systematic monitoring of forests and form a unique contributing data source.

Aims

A primary aim of this paper was to estimate recent growth changes in eight major European tree species, in natural contexts ranging from mountain to Mediterranean. A second aim was to explore their association with species’ climatic habitat and contemporary climate change.

Methods

Using >315,000 tree increments measured in >25,000 NFI plots, temporal changes in stand basal area increment (BAI) were modelled. Indicators of climate normals and of recent climatic change were correlated to species BAI changes.

Results

BAI changes spanned from ?17 to +42% over 1980–2007 across species. BAI strongly increased for mountain species, showed moderate/no increase for generalist and temperate lowland species and declined for Mediterranean species. BAI changes were greater in colder/wetter contexts than in warmer/drier ones where declines were observed. This suggested a role for climate warming, further found more intense in colder contexts and strongly correlated with species BAI changes.

Conclusion

The predominant role of climate warming and species climatic habitat in recent growth changes is highlighted in Western Europe. Concern is raised for Mediterranean species, showing growth decreases in a warmer climate with stable precipitation.
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10.

Key message

The disturbance of a research plot by a windstorm allowed us to study the role of the seedling bank in the regeneration processes. The released advance regeneration dominated among the saplings; taller individuals retained their position until the end of the study. Pioneer species occurred sporadically. Seven years after the disturbance, the windthrow was covered by a dense thicket of young trees.

Context

The dominant role played by advance regeneration in natural regeneration processes after intense wind disturbances is still a matter of dispute.

Aims

We took advantage of a windstorm in one of our research plots to study the role of the seedling bank released by the disturbance in the regeneration processes.

Methods

We collected data in 70 plots, recording the survivorship of seedlings, annual height growth, and signs of browsing. The height ranking was analyzed with Kendall’s concordance coefficient, and the height growth rates were compared using Dunn’s test.

Results

The density of seedlings increased from 6.7/m2 in 2008 to 8.1/m2 in 2010 and then decreased to 1.2/m2 in 2015. The density of saplings increased continuously from 0.14 to 1.9/m2. The highest size differentiation occurred in sycamore maple; the individuals which were taller before the windstorm retained their position until the year 2015. The only species that was recruited mainly from germinants was European hornbeam.

Conclusion

The advance regeneration released by the windstorm played a major role in the regeneration process, while pioneer species occurred only sporadically. Seven years after the disturbance, the windthrow was already covered by a dense thicket of young trees.
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11.

Key message

Cattle grazing and overstory cover restrict understory growth and interact in shaping the understory community structure in Mediterranean conifer plantations.

Context

Understanding how silvicultural manipulations drive understory structure and function in Mediterranean pine plantations is essential for their multifunctional management.

Aims

This paper aims to study the interactive effects of cattle grazing and overstory thinning on understory structure and function.

Methods

Ten plots (0.25 ha) were selected in East Mediterranean mature Pinus brutia plantation (rainfall = 600 mm year?1) representing thinned (≈100 trees ha?1, leaf area index (LAI) ≈ 1.6) and non-thinned (≈230 trees ha?1, LAI ≈ 3.5) areas. Two subplots (100 m2) within each plot were fenced in 2000 and 2006 while a third one remained grazed. Understory growth and species composition were measured in 2010.

Results

Thinning and grazing exclusion both positively influenced woody growth with their combined effect during 10 years leading to 20-fold increase in vegetation volume. An increase (15-fold) in herbaceous biomass was recorded 4 years after grazing exclusion but disappeared 10 years after exclusion due to increased woody cover. Species richness was not influenced by grazing but was positively affected by thinning. Understory composition was affected by grazing × thinning interaction with herbaceous ephemerals and short woody species being more frequent in grazed, thinned areas while larger woody species were more associated with ungrazed, non-thinned areas.

Conclusion

Grazing impacts on forest understories depend on overstory cover. We propose variable grazing-thinning combinations to meet multiple management objectives.
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12.

Key message

Growth and wood chemical properties are important pulpwood traits. Their narrow-sense heritability ranged from 0.03 to 0.49 in Eucalyptus urophylla × E. tereticornis hybrids, indicating low to moderate levels of genetic control. Genetic correlations were mostly favorable for simultaneous improvement on growth and wood traits. Additive and non-additive genetic effects should be considered in making a hybrid breeding strategy.

Context

Eucalypt hybrids are widely planted for pulpwood production purposes. Genetic variations and correlations for growth and wood chemical traits remain to be explored in Eucalyptus interspecific hybrids.

Aims

Our objectives were to clarify the heritability of growth and wood chemical traits and determine the genetic correlations between traits and between trials in E. urophylla × E. tereticornis hybrids.

Methods

Two trials of 59 E. urophylla × E. tereticornis hybrids derived from an incomplete factorial mating design were investigated at age 10 for growth (height and diameter) and wood chemical properties (basic density, cellulose content, hemi-cellulose content, lignin content, and syringyl-to-guaiacyl ratio). Mixed linear models were used to estimate genetic parameters.

Results

Narrow-sense heritability estimates were 0.13?0.22 in growth and 0.03?0.49 in wood traits, indicating low to moderate levels of additive genetic control. Genetic correlations were mostly positively significant for growth with basic density and cellulose content but negatively significant with hemi-cellulose and lignin contents, being favourablefavorable for pulpwood breeding purpose. Type-B correlations between sites were significant for all the traits except diameter and lignin content.

Conclusion

Hybrid superiority warrants the breeding efforts. An appropriate breeding strategy should be able to capture both additive and non-additive genetic effects.
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13.

Key message

Mixing sessile oak and Scots pine in central France to reduce intraspecific competition for water resources did not improve the ability of these two species to withstand severe drought during the summer.

Context

In order to reduce the impact of increasingly extreme droughts on forests, managers must adapt their practices to future climate conditions. Maintaining a greater diversity of tree species in temperate forest ecosystems is one of the recommended options.

Aims

We addressed how interactions between sessile oak and Scots pine in mixed forests in central France affect their functional response to drought.

Methods

We characterized the carbon isotope composition (δ13C) in the tree growth rings formed during wet (2001, 2007) or dry (2003, 2004) summers for each of the two species growing both in pure and in mixed stands in order to compare the effect of stand composition on variations in carbon isotope discrimination (Δ13C) among contrasted years.

Results

The severe drought in 2003 induced a strong decrease in Δ13C for all trees and in all stands as compared to 2001. This decrease was greater in pine than in oak. There was no significant difference between pure and mixed stands in the response of either species to drought.

Conclusion

Mixing sessile oak and Scots pine in stands in central France does not improve the ability of either species to withstand severe drought during the summer.
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14.

Key message

The changes in the relative biomass allocation to roots in juvenile stands of fast-growing ( Leucaena leucocephala Lam., Moringa oleifera Lam., and Jatropha curcas L.) and slow-growing ( Anacardium occidentale L. and Parkia biglobosa Jacq.) afforestation species are driven mainly by ontogeny rather than resource availability. However, silvicultural management aiming at increasing availability of water and particularly nutrients enhances biomass production in all species.

Context

Understanding the patterns of biomass allocation among tree species in response to ontogeny and to variation in resource availability is key to the successful restoration of degraded land using forest plantations.

Aims

This study assessed the effects of resource availability and ontogeny on biomass accumulation and partitioning in five semi-arid afforestation species.

Methods

The aboveground and belowground biomass production of fast-growing Leucaena leucocephala Lam., Moringa oleifera Lam., and Jatropha curcas L. and slow-growing Anacardium occidentale L. and Parkia biglobosa Jacq. was monitored following the application of manure (1 kg plant?1) and/or supplemental irrigation (0.5 L per sapling daily) during the first two rainy seasons and the intervening dry season on degraded cropland in Northern Benin.

Results

Biomass accumulation in the fast-growing species was positively impacted by fertilization and irrigation during both rainy seasons. The slow-growing species responded positively to the silvicultural treatments during the dry and second rainy season. The application of fertilizer alone increased the biomass of P. biglobosa by up to 335% during the dry season. Fifteen months after planting, manure-treated L. leucocephala accumulated the most biomass (2.9 kg tree?1). The root fraction decreased with increasing tree size in all species. The comparison of root versus shoot allocation in trees of equal size indicated that the treatment-induced shifts in biomass partitioning were controlled by ontogeny, which explained 86–95% of the variation in root-shoot biomass relationships.

Conclusion

While ontogeny was the main driver of biomass partitioning, increased resource availability induced a larger production of biomass, overall leading to greater aboveground production in all species.
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15.

Key message

In Europe, P. nigra wood presents a density pattern of longitudinal variation with an increase from east to west. However, no latitudinal tendencies were detected. Compared to other Portuguese resinous species, P. nigra revealed higher density, identical radial growth and intra-ring heterogeneity, which presents advantages for industry purposes. The environmental factors (Sites effect) manifest more strongly in the latewood components while the Trees/Sites effect is more strongly expressed in the earlywood components.

Context

Although P. nigra Arnold is one of the most important conifers in Europe, little is known about the wood’s characteristics in the southwest European region.

Aims

Our aims are to outline a first approach to study the growth and wood quality in P. nigra in Portugal comparing to other European natural stands and other resinous species.

Methods

Inter- and intra-wood density variation of P. nigra from six Portuguese sites was studied using microdensitometry. Analysis of variance (ANOVA) was performed in three subsets: 50 common rings, core (juvenile wood) and peripheral analysis (mature wood).

Results

The average ring density was 0.588 g cm?3, with maximum values in the north and low altitudes. Regarding growth traits, no latitudinal and altitudinal tendencies were detected. Compared to the main timber species in Portugal (P. pinaster Aiton), P. nigra showed similar radial growth, higher density but lower intra-ring density homogeneity. The Sites effect mainly influenced latewood density components, while the Trees/Sites effect primarily influenced earlywood components. The Rings effect was found to be relatively low, with a density decrease in the tree’s first years followed by an increase in the periphery. Growth traits showed a reduction from pith to bark.

Conclusion

Considering the quality (density) and growth features of the Black pine, this species could be useful for the reforestation of mountainous Southern Europe areas that are not favourable for other species.
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16.

Key message

In Appalachian hardwood forests, density, stem size, and productivity affected growth during drought for red oak, but not white oak species. Minor effects of density suggest that a single low thinning does little to promote drought resilience for oaks in the region.

Context

Management is increasingly focused on promoting resilience to disturbance. Because stand density can modulate climate-growth relationships, thinning may be an adaptation strategy that promotes resistance/resilience to drought.

Aims

We examined how density, manipulated via thinning, stem size, and site productivity, influences the drought response of northern red, black, chestnut, and white oak.

Methods

We modeled the role of density, stem size, and site productivity on resistance, recovery, and resilience during two drought events.

Results

Chestnut and white oak displayed greater resistance, recovery, and/or resilience than did northern red and black oak. For black oak, density and stem size negatively affected resistance during the first and second drought, respectively. Density, stem size, and site productivity had no effect on chestnut and white oak.

Conclusion

The lack of sensitivity of chestnut and white oak to the ranges of density, stem size, and site productivity observed in this study and generally better resistance, recovery, and resilience suggests that management focused on the maintenance of these species, as opposed to a single silvicultural low thinning, may be a possible strategy for sustaining the growth and productivity of oak species in Appalachian hardwood stands. Drought response as affected by alternative thinning interventions should be evaluated.
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17.

Key message

Pinus sylvestris seedlings quickly expand their roots to deeper soil layers while Pseudotsuga menziesii concentrates its root system in the topsoil, thereby running the risk of desiccation during long dry spells, as indicated by lower survival after simulated summer drought.

Context

Pseudotsuga menziesii (Douglas-fir) is regarded as a promising species to maintain the productivity of Central European lowland forests given the projected increase of long dry spells.

Aims

Will the species be able to regenerate from seed and spread outside plantations in a drier temperate Europe?

Methods

We measured the relative growth rate, biomass allocation, root architecture, and phenotypic plasticity of Pseudotsuga menziesii seedlings sown in a common garden and grown under current precipitation and prolonged drought, respectively. The species’ competitive ability with respect to Pinus sylvestris L., the most drought-tolerant native conifer in Central Europe, was assessed during three growing seasons.

Results

Pinus sylvestris seedlings had higher relative growth rates than did Pseudotsuga menziesii seedlings, first in terms of aboveground biomass and later in terms of shoot height. This resulted in heavier and taller seedlings after three growing seasons under both moist and dry conditions. Shorter vertical roots corresponded with lower survival of Pseudotsuga menziesii seedlings under dry conditions.

Conclusion

Fast root proliferation allows Pinus sylvestris seedlings to reach deeper water pools that are less rapidly depleted during transient drought. By contrast, the shallow root system might put Pseudotsuga menziesii seedlings at the risk of desiccation during prolonged dry spells.
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18.

Key message

We demonstrate that, beyond leaf phenology, the phenological cycles of wood and fine roots present clear responses to environmental drivers in temperate and boreal trees. These drivers should be included in terrestrial ecosystem models.

Context

In temperate and boreal trees, a dormancy period prevents organ development during adverse climatic conditions. Whereas the phenology of leaves and flowers has received considerable attention, to date, little is known regarding the phenology of other tree organs such as wood, fine roots, fruits, and reserve compounds.

Aims

Here, we review both the role of environmental drivers in determining the phenology of tree organs and the models used to predict the phenology of tree organs in temperate and boreal forest trees.

Results

Temperature is a key driver of the resumption of tree activity in spring, although its specific effects vary among organs. There is no such clear dominant environmental cue involved in the cessation of tree activity in autumn and in the onset of dormancy, but temperature, photoperiod, and water stress appear as prominent factors. The phenology of a given organ is, to a certain extent, influenced by processes in distant organs.

Conclusion

Inferring past trends and predicting future trends of tree phenology in a changing climate requires specific phenological models developed for each organ to consider the phenological cycle as an ensemble in which the environmental cues that trigger each phase are also indirectly involved in the subsequent phases. Incorporating such models into terrestrial ecosystem models (TEMs) would likely improve the accuracy of their predictions. The extent to which the coordination of the phenologies of tree organs will be affected in a changing climate deserves further research.
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19.

? Key message

The optimal management of larch (Larix olgensis) plantations in Northeast China consisted of 2 or 3 thinnings and a rotation length of 55–61 years when economic profitability, wood production, and carbon sequestration were simultaneously maximized. Wood production ranged from 5.4 to 11.7 m3 ha?1 a?1, depending on site quality.

? Context

L. olgensis is an important tree species in the northeast forest region of China, playing a significant role in the establishment of fast-growing and high-yielding plantation forests in China. However, the management of these plantations has not been optimized in previous studies.

? Aims

The objective of the study was to find the optimal combinations of thinning times, thinning types, and rotation length for L. olgensis stands when both timber production and carbon stock are considered.

? Methods

First, a growth and yield model was developed to simulate the dynamics of larch plantations. Then, the models were linked with the Hooke and Jeeves optimization algorithm to optimize forest management for two commonly used planting densities and three site qualities.

? Results

Two thinnings were found to be suitable for larch plantations when the stand density at 10 years was 2125 trees/ha (corresponding to a planting density of 2500 trees/ha) whereas three thinnings were recommended when the density at 10 years was 2800 trees/ha (planting density of 3300 trees/ha). When the stand density was 2800 trees/ha, the optimal rotation length was 61, 58, and 55 years for site indices (SI) 12, 16, and 20 m (dominant height at 30 years), respectively. The mean annual wood production was 5.4 m3 ha?1 for SI 12, 8.2 m3 ha?1 for SI 16, and 11.7 m3 ha?1 for SI 20. The results were nearly the same for the lower initial stand density. The better the site quality of the stand, the earlier the thinnings were conducted.

? Conclusion

In multifunctional forestry, optimal rotation lengths of larch plantations were 10–20 years longer than advised in the current silvicultural recommendations for Northeast China.
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20.

? Key message

Natural regeneration of P. abies (L.) H. Karst. may reach high densities in lower mountain elevations. The highest densities were found in sites with moderate light availability, with low pH, and not near the riverbank. However, age-height classes differed in the predicted magnitude of response, but were consistent in response directions. Mosses and understory species typical of coniferous forests were positively correlated with regeneration density.

? Context

Norway spruce Picea abies (L.) H. Karst. in Central Europe is at risk under climate change scenarios, particularly in mountain regions. Little is known about the impact of environmental factors on the natural regeneration of P. abies in low-elevation mountain forests.

? Aims

We aimed to assess impacts of distance from the riverbank, soil pH, and light availability on natural P. abies regeneration. We hypothesized that (1) natural P. abies regeneration would depend on light availability and soil pH and (2) there are understory plant species which may indicate the microsites suitable for natural regeneration of P. abies.

? Methods

The study was conducted in the Sto?owe Mountains National Park (SW Poland, 600–800 m a.s.l.). We established 160 study plots (25 m2) for natural regeneration, light availability, soil pH, and understory vegetation assessment.

? Results

The highest densities of natural regeneration of P. abies were observed in sites with moderate light availability (0.1–0.2 of open sky) and low pH (3.5–4.5), and located relatively far from the riverbank. Cover of 22 understory plant species were correlated with natural P. abies regeneration densities, mostly positively.

? Conclusion

Different stages of natural regeneration of P. abies revealed different regeneration niches. Most understory plant species (bryophytes and herbs typical of coniferous forests) do not compete with natural regeneration of P. abies.
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