Forest diversity-productivity relationships have been intensively investigated in recent decades. However, few studies have considered the interplay between species and structural diversity in ...driving productivity. We analyzed these factors using data from 52 permanent plots in southwestern Germany with more than 53,000 repeated tree measurements. We used basal area increment as a proxy for productivity and hypothesized that: (1) structural diversity would increase tree and stand productivity, (2) diversity-productivity relationships would be weaker for species diversity than for structural diversity, and (3) species diversity would also indirectly impact stand productivity via changes in size structure. We measured diversity using distance-independent indices indices. We fitted separate linear mixed-effects models for fir, spruce and beech at the tree level, whereas at the stand level we pooled all available data. We tested our third hypothesis using structural equation modeling. Structural and species diversity acted as direct and independent drivers of stand productivity, with structural diversity being a slightly better predictor. Structural diversity, but not species diversity, had a significant, albeit asymmetric, effect on tree productivity. The functioning of structurally diverse, mixed forests is influenced by both structural and species diversity. These sources of trait diversity contribute to increased vertical stratification and crown plasticity, which in turn diminish competitive interferences and lead to more densely packed canopies per unit area. Our research highlights the positive effects of species diversity and structural diversity on forest productivity and ecosystem dynamics.
Improving our understanding of the potential of forest adaptation is an urgent task in the light of predicted climate change. Long‐term alternatives for susceptible yet economically important tree ...species such as Norway spruce (Picea abies) are required, if the frequency and intensity of summer droughts will continue to increase. Although Silver fir (Abies alba) and Douglas fir (Pseudotsuga menziesii) have both been described as drought‐tolerant species, our understanding of their growth responses to drought extremes is still limited. Here, we use a dendroecological approach to assess the resistance, resilience, and recovery of these important central Europe to conifer species the exceptional droughts in 1976 and 2003. A total of 270 trees per species were sampled in 18 managed mixed‐species stands along an altitudinal gradient (400–1200 m a.s.l.) at the western slopes of the southern and central Black Forest in southwest Germany. While radial growth in all species responded similarly to the 1976 drought, Norway spruce was least resistant and resilient to the 2003 summer drought. Silver fir showed the overall highest resistance to drought, similarly to Douglas fir, which exhibited the widest growth rings. Silver fir trees from lower elevations were more drought prone than trees at higher elevations. Douglas fir and Norway spruce, however, revealed lower drought resilience at higher altitudes. Although the 1976 and 2003 drought extremes were quite different, Douglas fir maintained consistently the highest radial growth. Although our study did not examine population‐level responses, it clearly indicates that Silver fir and Douglas fir are generally more resistant and resilient to previous drought extremes and are therefore suitable alternatives to Norway spruce; Silver fir more so at higher altitudes. Cultivating these species instead of Norway spruce will contribute to maintaining a high level of productivity across many Central European mountain forests under future climate change.
The two extreme droughts in 1976 and 2003 affected negatively the radial growth response of Norway spruce, Silver and Douglas fir in the Black forest at all elevations. The 1976 drought had a less pronounced effect than the 2003 summer drought; however, firs were noticeably more resistant and resilient to extreme drought than spruce. Spruce was the most affected species when comparing performances of drought indices, and Silver fir the least affected. Douglas fir showed consistently the highest growth rates.
•First meta-analysis on effects of thinning to mitigate drought stress in trees.•Thinning improves the physiological and growth response of trees to drought.•Growth benefits before, during and after ...drought increase with thinning intensity.•Drought-mitigation through thinning differs between conifers and broadleaves.•Benefits for post-drought growth decrease with time since the last thinning.
Increasing frequency of extremely dry and hot summers in some regions emphasise the need for silvicultural approaches to increase the drought tolerance of existing forests in the short term, before long-term adaptation through species changes may be possible. The aim of this meta-analysis was to assess the potential of thinning for improving tree performance during and after drought. We used results from 23 experiments that employed different thinning intensities including an unthinned control and focused on the response variables: radial growth, carbon- and oxygen-isotopes in tree-rings and pre-dawn leaf-water potential. We found that thinning effects on the growth response to drought differed between broadleaves and conifers, although these findings are based on few studies only in broadleaved forests. Thinning helped to mitigate growth reductions during drought in broadleaves, most likely via increases of soil water availability. In contrast, in conifers, comparable drought-related growth reductions and increases of water-use efficiency were observed in all treatments but thinning improved the post-drought recovery and resilience of radial growth. Results of meta-regression analysis indicate that benefits of both moderate and heavy thinning for growth performance following drought (recovery and resilience) decrease with time since the last intervention. Further, growth resistance during drought became smaller with stand age while the rate of growth recovery following drought increased over time irrespective of treatment. Heavy but not moderate thinning helped to avoid an age-related decline in medium-term growth resilience to drought. For both closed and very open stands, growth performance during drought improved with increasing site aridity but for the same stands growth recovery and resilience following drought was reduced with increasing site aridity. This synthesis of experiments from a wide geographical range has demonstrated that thinning, in particular heavy thinning, is a suitable approach to improve the growth response of remaining trees to drought in both conifers and broadleaves but the underlying processes differ and need to be considered.
Silviculture for old-growth attributes Bauhus, Jürgen; Puettmann, Klaus; Messier, Christian
Forest ecology and management,
07/2009, Volume:
258, Issue:
4
Journal Article, Conference Proceeding
Peer reviewed
Open access
Silviculture to maintain old-growth forest attributes appears to be an oxymoron since the late developmental phases of forest dynamics, described by the term old-growth, represent forests that have ...not experienced human intervention or timber removal for a long time. In the past, silvicultural systems applied to old-growth aimed to convert it into simplified, more productive regrowth forests substantially different in structure and composition. Now it is recognised that the maintenance of biodiversity associated with structural and functional complexity of late forest development successional stages cannot rely solely on old-growth forests in reserves. Therefore, in managed forests, silvicultural systems able to develop or maintain old-growth forest attributes are being sought. The degree to which old-growth attributes are maintained or developed is called “old-growthness”. In this paper, we discuss silvicultural approaches that promote or maintain structural attributes of old-growth forests at the forest stand level in (a) current old-growth forests managed for timber production to retain structural elements, (b) current old-growth forests requiring regular, minor disturbances to maintain their structure, and (c) regrowth and secondary forests to restore old-growth structural attributes. While the functions of different elements of forest structure, such as coarse woody debris, large veteran trees, etc., have been described in principle, our knowledge about the quantity and distribution, in time and space, of these elements required to meet certain management objectives is rather limited for most ecosystems. The risks and operational constraints associated with managing for structural attributes create further complexity, which cannot be addressed adequately through the use of traditional silvicultural approaches. Silvicultural systems used in the retention and restoration of old-growthness can, and need, to employ a variety of approaches for managing spatial and temporal structural complexity. We present examples of silvicultural options that have been applied in creative experiments and forestry practice over the last two decades. However, these largely comprise only short-term responses, which are often accompanied by increased risks and disturbance. Much research and monitoring is required still to develop and optimise new silvicultural systems for old-growthness for a wide variety of forest ecosystem types.
1. Promoting mixed-species forests is an important strategy for adaptation and risk reduction in the face of global change. Concurrently, a main challenge in ecology is to quantify the effects of ...species diversity on ecosystem functioning. In forests, the effects of individual tree species on ecosystem functions depend largely on their dimensions, which are commonly predicted using allometric equations. However, little is known about how diversity influences allometry or how to incorporate this effect into allometric equations. Ignoring the effects of interspecific interactions on allometric relationships may result in severely biased predictions. 2. This study examined the effects of tree-species diversity, competition and tree social status on crown-projection area (cpa), height (h) and live-crown length (lcl) of trees using a European-wide data set containing 17 target species and 12 939 trees. The cpa, h and lcl were predicted as functions of stem diameter at 1·3 m, tree-species diversity, tree height relative to the stand mean height (rh) and a competition index (CI) that accounted for stand density and interspecific differences in competitive ability based on species-specific wood density or shade tolerance. 3. Averaged across species, diameter had the greatest effect on cpa and lcl, followed by the competition index, while rh had the greatest effect on lcl. Tree-species diversity had the smallest effect on cpa, h and lcl. Interspecific variability in cpa, h or lcl responses to diversity, CI, or rh was sometimes related to wood density or shade tolerance. 4. Synthesis. This study shows the strong influence of stand structure and species composition on allometric relationships. These influences can be quantified using measures of competition, tree-species diversity and relative tree height so that general equations can be developed for a given species to be applied to a wide range of species compositions and stand structures. This new approach will greatly improve predictions of biomass and carbon stocks in structurally and compositionally diverse forests.
1. Partitioning of tree mortality into different modes of death allows the tracing and mechanistic modelling of individual key processes of forest dynamics each varying depending on site, species and ...individual risk factors. This, in turn, may improve long-term predictions of the development of old-growth forests. 2. Six different individual tree mortality modes (uprooted and snapped (both with or without rot as a predisposing factor), standing dead and crushed by other trees) were analysed, and statistical models were derived for three tree species (European beech Fagus sylvatica, hornbeam Carpinus betulus and common ash Fraxinus excelsior) based on a repeated inventory of more than 13 000 trees in a 28 ha near-natural deciduous forest in Central Germany. 3. The frequently described U-shaped curve of size-dependent mortality was observed in beech and hornbeam (but not ash) and could be explained by the joint operation of processes related to the six distinct mortality modes. The results for beech, the most abundant species, suggest that each mortality mode is prevalent in different life-history stages: small trees died mostly standing or being crushed, medium-sized trees had the highest chance of survival, and very large trees experienced increased rates of mortality, mainly by uprooting or snapping. Reduced growth as a predictor also played a role but only for standing dead, all other mortality modes showed no relationship to tree growth. 4. Synthesis. Tree mortality can be partitioned into distinct processes, and species tend to differ in their susceptibility to one or more of them. This forms a fundamental basis for the understanding of forest dynamics in natural forests, and any mechanistic modelling of mortality in vegetation models could be improved by correctly addressing and formulating the various mortality processes.
Climate change is expected to pose major direct and indirect threats to groundwater-dependent forest ecosystems. Forests that concurrently experience increased rates of water extraction may face ...unprecedented exposure to droughts. Here, we examined differences in stem growth and xylem hydraulic architecture of 216 oak trees from sites with contrasting groundwater availability, including sites where groundwater extraction has led to reduced water availability for trees over several decades. We expected reduced growth and xylem hydraulic capacity for trees at groundwater extraction sites both under normal and unfavourable growing conditions. Compared to sites without extraction, trees at sites with groundwater extraction showed reduced growth and hydraulic conductivity both during periods of moderate and extremely low soil water availability. Trees of low vigour, which were more frequent at sites with groundwater extraction, were not able to recover growth and hydraulic capacity following drought, pointing to prolonged drought effects. Long-term water deficit resulting in reduced CO
assimilation and hydraulic capacity after drought are very likely responsible for observed reductions in tree vitality at extraction sites. Our results demonstrate that groundwater access maintains tree function and resilience to drought and is therefore important for tree health in the context of climate change.
Habitat trees, which provide roosting, foraging and nesting for multiple taxa, are retained in managed forests to support biodiversity conservation. To what extent their spatial distribution ...influences provisioning of habitats has rarely been addressed. In this study, we investigated whether abundance and richness of tree-related microhabitats (TreMs) differ between habitat trees in clumped and dispersed distributions and whether the abundance of fifteen groups of TreMs is related to tree distribution patterns. To identify habitat trees, we quantified TreMs in temperate mountain forests of Germany. We determined clumping (the Clark–Evans index), size of the convex hull, diameter at breast height, as well as altitude, slope and aspect of sites for their possible influence on TreMs. We additionally determined the difference in TreM abundance and richness among four options of selecting five habitat trees per ha from 15 candidates: (a) the most clumped trees, (b) five randomly selected and dispersed trees, (c) the single tree with highest abundance or richness of TreMs and its four closest neighbors and (d) a “reference selection” of five trees with known highest abundance or richness of TreMs irrespective of their distribution. The degree of clumping and the size of the convex hull influenced neither the abundance nor richness of TreMs. The reference selection, option (d), contained more than twice the number of TreMs compared to the most clumped, (a), or random distributions, (b), of five habitat trees, while option (c) assumed an intermediate position. If the goal of habitat tree retention is to maximize stand-level abundance and richness of TreMs, then it is clearly more important to select habitat trees irrespective of their spatial pattern.
Phosphorus is one of the major limiting factors of primary productivity in terrestrial ecosystems and, thus, the P demand of plants might be among the most important drivers of soil and ecosystem ...development. The P cycling in forest ecosystems seems an ideal example to illustrate the concept of ecosystem nutrition. Ecosystem nutrition combines and extents the traditional concepts of nutrient cycling and ecosystem ecology. The major extension is to consider also the loading and unloading of nutrient cycles and the impact of nutrient acquiring and recycling processes on overall ecosystem properties. Ecosystem nutrition aims to integrate nutrient related aspects at different scales and in different ecosystem compartments including all processes, interactions and feedbacks associated with the nutrition of an ecosystem. We review numerous previous studies dealing with P nutrition from this ecosystem nutrition perspective. The available information contributes to the description of basic ecosystem characteristics such as emergence, hierarchy, and robustness. In result, we were able to refine Odum's hypothesis on P nutrition strategies along ecosystem succession to substrate related ecosystem nutrition and development. We hypothesize that at sites rich in mineral‐bound P, plant and microbial communities tend to introduce P from primary minerals into the biogeochemical P cycle (acquiring systems), and hence the tightness of the P cycle is of minor relevance for ecosystem functioning. In contrast, tight P recycling is a crucial emergent property of forest ecosystems established at sites poor in mineral bound P (recycling systems). We conclude that the integration of knowledge on nutrient cycling, soil science, and ecosystem ecology into holistic ecosystem nutrition will provide an entirely new view on soil–plant–microbe interactions.
The phenomenon of overyielding in speciesdiverse plant communities is mainly attributed to complementary resource use. Vertical niche differentiation belowground might be one potential mechanism for ...such complementarity. However, most studies that have analysed the diversity/productivity relationship and belowground niche differentiation have done so for fully occupied sites, not very young tree communities that are in the process of occupying belowground space. Here we used a 5—6 year old forest diversity experiment to analyse how fine-root (<2 mm) production in ingrowth cores (0—30 cm) was influenced by tree species identity, as well as the species diversity and richness of tree neighbourhoods. Fineroot production during the first growing season after the installation of ingrowth cores increased slightly with tree species diversity, and four-species combinations produced on average 94.8% more fine-root biomass than monocultures. During the second growing season, fine-root mortality increased with tree species diversity, indicating an increased fine-root turnover in species-rich communities. The initial overyielding was attributable to the response to mixing by the dominant species, Pseudotsuga menziesii and Picea abies, which produced more fine roots in mixtures than could be expected from monocultures. In species-rich neighbourhoods, P. abies allocated more fine roots to the upper soil layer (0—15 cm), whereas P. menziesii produced more fine roots in the deeper layer (15—30 cm) than in species-poor neighbourhoods. Our results indicate that, although there may be no lasting overyielding in the fineroot production of species-diverse tree communities, increasing species diversity can lead to substantial changes in the production, vertical distribution, and turnover of fine roots of individual species.
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