•A database containing nearly 1000 European biomass equations was developed.•Biomass and leaf area allometry were influenced by stand structure.•Species traits were correlated with interspecific ...differences in responses to stand structure.
Biomass and leaf area equations are often required to assess or model forest productivity, carbon stocks and other ecosystem services. These factors are influenced by climate, age and stand structural attributes including stand density and tree species diversity or species composition. However, such covariates are rarely included in biomass and leaf area equations. We reviewed the literature and built a database of biomass and leaf area equations for 24 European tree species and 3 introduced species. The final dataset contained 973 equations. Most of the equations were site-specific and therefore restricted to the edaphic, climatic and stand structural conditions of the given site. To overcome this limitation, the database was used to develop regional species-specific equations that can be used in a wide range of stands and to quantify the effects of climate, age and stand structure on biomass or leaf area. The analysis showed considerable inter- and intra-specific variability in biomass relationships. The intra-specific variability was related to climate, age or stand characteristics, while the inter-specific variability was correlated with traits such as wood density, specific leaf area and shade tolerance. The analysis also showed that foliage mass is more variable than stem or total aboveground biomass, both within and between species, and these biomass components have contrasting responses to age and changes in stand structure. Despite the large number of published equations, many species are still not well represented. Therefore, generic equations were developed that include species-specific wood density instead of species identity. Further improvements may be possible if future studies quantify the stand structure of individual tree neighbourhoods instead of using the stand means for all trees sampled with the given stand.
1. When tree-species mixtures are more productive than monocultures, higher light absorption is often suggested as a cause. However, few studies have quantified this effect and even fewer have ...examined which light-related interactions are most important, such as the effects of species interactions on tree allometric relationships and crown architecture, differences in vertical or horizontal canopy structure, phenology of deciduous species or the mixing effects on tree size and stand density. 2. In this study, measurements of tree sizes and stand structures were combined with a detailed tree-level light model (Maestra) to examine the contribution of each light-related interaction on tree- and stand-level light absorption at 21 sites, each of which contained a triplet of plots including a mixture and monocultures of Fagus syivatica and Pinus sylvestris (63 plots). These sites were distributed across the current distribution of these species within Europe. 3. Averaged across all sites, the light absorption of mixtures was 14% higher than the mean of the monocultures. At the whole community level, this positive effect of mixing on light absorption increased as canopy volume or site productivity increased, but was unrelated to climate. At the species population or individual tree levels, the mixing effect on light absorption resulted from light-related interactions involving vertical canopy structure, stand density, the presence of a deciduous species (F. syivatica), as well as the effects of mixing on tree size and allometric relationships between diameter and height, crown diameter and crown length. 4. The mixing effects on light absorption were only correlated with the mixing effects on growth for P. sylvestris, suggesting that the mixing effects on this species were driven by the light-related interactions, whereas mixing effects on F. syivatica or whole community growth were probably driven by non-light-related interactions. 5. Synthesis. The overall positive effect of mixing on light absorption was the result of a range of light-related interactions. However, the relative importance of these interactions varied between sites and is likely to vary between other species combinations and as stands develop.
•Thinning intensity in uneven-aged interior Douglas-fir dominated stands influenced the spatial pattern of live trees.•Thinning influenced the spatial correlation of live tree diameters.•No spatial ...relationship existed between surviving and dead trees in unthinned plots.•The unthinned plots exhibited mainly small tree mortality.
Understanding the spatial patterns of trees and their interactions can reveal the ecological processes driving forest stand structure and stand development over time. We assessed temporal changes in tree spatial patterns in uneven-aged interior Douglas-fir (Pseudotsuga menziesii var. glauca (Beissn.) Franco) dominated stands in central British Columbia, Canada. Data were available on 24 plots in three blocks over 21 years, 18 of which had received pre-commercial thinning (PCT) treatments of varying intensity. We first applied the Clark and Evans aggregation index and the L function, a transformation of Ripley’s K function, to describe the spatial pattern of live trees in thinned and unthinned plots over time. Second, we analysed the spatial correlations between live tree diameters using the mark correlation function and the mark variogram. Third, the spatial pattern of dead trees in the unthinned plots after 21 years was analysed. Lastly, we tested the spatial relationship between dead and surviving trees in the unthinned plots. In all three blocks, the spatial patterns of live trees in the unthinned plots were clustered through time. The moderate thinning treatments had random or regular spatial patterns that remained unchanged through time, because of reduced mortality rates and low levels of ingrowth. The heavier thinning initially retained a clustered pattern at small inter-tree distances; however, this changed to a random pattern with increasing distance 15 years post-thinning and remained as such until the end of the study period. Tree diameters were not spatially autocorrelated in the thinned plots, although there was positive spatial correlation of tree diameters in the unthinned plots, probably due to competitive growth inhibition among neighbouring trees. Dead trees were primarily smaller in size and were significantly clustered at all spatial scales. The lack of spatial relationship between dead and surviving trees indicated that mortality was a random process. Our study contributes to a better understanding of how spatial patterns of trees change over time in these stands, which could help in the design of silvicultural regimes that mimic natural processes.
Mixed forests make up the majority of natural forests, and they are conducive to improving the resilience and resistance of forest ecosystems. Moreover, it is in the crown of the trees where the ...effect of inter- and intra-specific interaction between them is evident. However, our knowledge of changes in crown morphology caused by density, competition, and mixture of specific species is still limited. Here, we provide insight on stand structural complexity based on the study of four response crown variables (Maximum Crown Width Height, MCWH; Crown Base Height, CBH; Crown Volume, CV; and Crown Projection Area, CPA) derived from multiple terrestrial laser scans. Data were obtained from six permanent plots in Northern Spain comprising of two widespread species across Europe; Scots pine (Pinus sylvestris L.) and sessile oak (Quercus petraea (Matt.) Liebl.). A total of 193 pines and 256 oaks were extracted from the point cloud. Correlation test were conducted (ρ ≥ 0.9) and finally eleven independent variables for each target tree were calculated and categorized into size, density, competition and mixture, which was included as a continuous variable. Linear and non-linear multiple regressions were used to fit models to the four crown variables and the best models were selected according to the lowest AIC Index and biological sense. Our results provide evidence for species plasticity to diverse neighborhoods and show complementarity between pines and oaks in mixtures, where pines have higher MCWH and CBH than oaks but lower CV and CPA, contrary to oaks. The species complementarity in crown variables confirm that mixtures can be used to increase above ground structural diversity.
Identifying the factors controlling tree mortality is key to understanding the effects of ongoing global change on treeline movement and species composition. One potentially significant but little ...studied factor that impacts the formation of treelines is mortality caused by snow fungi. We studied the mortality patterns of Pinus cembra, a typical treeline species of the Central Alps of Switzerland, in two plantations located on opposite slopes of the Dischma valley (Northeast NE and Southwest SW) above the current treeline. In 1975, 33,000 P. cembra trees were planted at the NE site and 550 trees at the smaller SW site. All trees have been periodically monitored for survival since then. After 30 years, only ca. 5% of all P. cembra trees survived in both plantations. Two species of pathogenic snow fungus, Gremmeniella abietina and Phacidium. infestans, were major mortality agents for saplings. High rates of infection by Gremmeniella corresponded to late snowmelt and high ratios of rainfall to temperature. High rates of infection by Phacidium corresponded to earlier snowmelt and were spatially associated with P. cembra mature trees that had been present in the area before the time of planting. We provide experimental evidence that snow fungi are a primary cause of sapling tree mortality at treeline. Although additional evidence from different geographic regions and more natural treeline distributions is needed, our results suggest that the prevalence of snow fungi may prevent establishment of trees above the current treeline at present, and under future climate scenarios.
•Species coexistence was assessed in naturally regenerated mixed oak stands.•Each species is highly aggregated and the two species are strongly segregated.•The two species show similar growth rates ...and competitive abilities.•The oak mixture is very stable at the regeneration stage.•No silvicultural operation is required to maintain the mixture.
In most mixed-species forestry systems, regeneration is a critical step for individual tree survival and species coexistence. Species coexistence is driven by the fine-scale spatial patterns and competitive abilities of species. In naturally regenerated mixed oak (Quercus petraea and Quercus robur: QP and QR, respectively) stands, species coexistence was assessed by analysing the spatial distribution of both species and their growth responses to intra- and interspecific competition.
Four sites located in North Eastern France with an established (1.4–2.8-m-high) mixed QP and QR regeneration were selected for the study. In a first step, individual seedling location was recorded along line transects to analyse the spatial distribution of the two species. Univariate pair correlation functions were used to test for spatial aggregation for each species and bivariate pair correlation functions were used to test for spatial associations between the two species. In a second step, seedlings were measured in small plots to analyse their response to local competition. Mixed linear models were used to estimate the combined effects of density, mixture, seedling size and seedling status on seedling growth.
QP and QR were both highly aggregated in monospecific patches, and the two species were strongly segregated at a decametric scale. Transition zones between monospecific patches were reduced to a few meters. For both species, mixture had no effect on seedling growth. In all sites, the two species showed very similar diameter and height growth rates in response to density, mixture, seedling size and seedling status, and similar intra- and interspecific competitive abilities.
Highly patchy patterns of QP and QR regeneration strongly reduce interactions between the two species and, in transition zones between monospecific patches where the two species are mixed, no species is expected to outcompete its companion species. For each species, long-term persistence at the stand scale is determined by intraspecific competition rather than by interspecific competition. At this developmental stage, the QP-QR mixture appears to be very stable and no specific silvicultural treatment is required to maintain the mixture.
Alpine treeline ecosystems are generally expected to advance with increasing temperatures and after land-use abandonment. Multiple interacting factors modify this trend. Understanding the long-term ...processes underlying treeline advance is essential to predict future changes in structure and function of mountain ecosystems. In a valley in the Central Swiss Alps, we re-assessed a 40-year-old survey of all treeline trees (>0.5 m height) and disentangled climate, topographical, biotic, and disturbance (land use and avalanche risk) factors that have led to treeline advance with a combination of ground-based mapping, decision tree, and dendroecological analyses. Between the first ground survey in 1972/73 and the resurvey in 2012, treeline advanced on average by 10 meters per decade with a maximum local advance of 42 meters per decade. Larch consistently advanced more on south-facing slopes, while pine advance was greater on north-facing slopes. Newly established spruce mostly represented infilling below the previous treeline. The forefront of treeline advance above 2330 m a.s.l. occurred mainly on favorable microsites without competing dwarf shrub vegetation. At slightly lower elevations, treeline advanced mainly on sites that were used for agriculture at the beginning of the 20th century. This study indicates that although treeline advances under the effect of climate warming, a combination of additional ecological factors controls this advance at regional and local scales.
KEY MESSAGE : Density was more important in shaping crown structure than neighbor species identity. Both species showed high crown plasticity at alternative levels, which may explain species ...coexistence in mixed broadleaved forests with functionally similar species. Understanding crown response to local competition is essential to predicting stand development in mixed stands. We analyzed data from an 8-year-old field experimental plantation mixing two species according to a crossed gradient of density and species proportion to quantify the effect of a broad range of local neighborhood conditions on the development of young trees at multiple crown levels. We used Fagus sylvatica and Acer pseudoplatanus, as two model deciduous species. They are considered functionally equivalent at the young stages, but with contrasting architectural patterns. For both species: (1) changes in density explained more of the variation on crown development than species proportion (2) much of the effect of competition was accounted for by variables at the stem level, while branch and leaf development within crowns were not directly altered by competition. Both species were able to modify their crowns at the stem level to compete with intra- and inter-specific neighbors: Acer and Fagus were taller with a highest proportion of Fagus as neighbors; Fagus displayed a lower crown base when the proportion of Fagus decreased, while Acer had a lower crown base when the proportion of Fagus around it increased. Both species showed common shapes in allometric relationships but contrasting responses at alternative crown levels. Acer exhibited broader intra-specific variation in its height–diameter relationship and in its crown length, while Fagus displayed higher individual variation of branch development and leaf area than Acer. This study demonstrates that differences in crown development strategy of each species in response to changes in local neighborhood conditions are an important factor in maintaining species coexistence in broadleaved forests and designing mixtures that persist over time.
•We studied the mortality in unmanaged 100–150years old Norway spruce forests.•Climate extremes acted together with competition to cause sharp growth declines.•Most dead trees had one to several ...growth declines lasting from few years to decades.•Trees which died had significantly lower initial radial growth.•More frequent future droughts may contribute to higher rates of mortality in subalpine forests.
Climatic variability often is thought to be most important for ecosystem development at ecotones, while competition is thought to be most important farther from ecotones, where neighboring plants compete for scarce resources. However, climatic variability may also modulate consequences of competition, especially under recent and future climate change. Norway spruce (Picea abies) forests are among the most important ecosystems in the mountain regions of Europe and provide various ecosystem services. Many of these forests are currently in a self-thinning, stem-exclusion phase. Understanding processes governing forest dynamics during this phase is necessary for understanding future forest structure and processes as well as effects of climatic variability on ongoing forest development. We studied growth and mortality patterns in unmanaged 100–150years-old Norway spruce forests that originated after stand-replacing disturbances in the Parangalitsa Reserve in Bulgaria. We collected data on forest structure and tree ring samples from 648 live and dead trees (DBH >4cm) to analyze onset, pattern and duration of mortality, as well as contributing factors.
We found that climate extremes acted together with competition to cause sharp growth declines lasting from a few years to several decades and, in some cases, eventually led to death. The majority of dead trees had one to several consecutive growth declines, most of which initiated in response to extreme summer droughts during periods of high within-stand competition (after trees were 40–50years old). Our tree-ring analysis revealed that some suppressed trees that died were more drought-sensitive than living trees. Other climate extremes such as unusually cold winters or summers also contributed to sharp growth reductions in some cases. Trees that died had significantly lower initial radial growth, which suggests that in the absence of external disturbances, the outcome of mortality in the stem-exclusion stage may be pre-determined from factors that determine initial growth rates. Spatial distribution data showed that there was no significant aggregation of dead and live trees and that in almost all cases, neither live nor dead trees were clustered.
Our findings contribute to understanding mortality processes in self-thinning subalpine Norway spruce forests in Europe and show that under climate change scenarios that include more frequent future droughts, even forests in which competition is thought to be the main driver of dynamics, may experience higher rates of mortality.
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