•Terrestrial laser scanning (TLS) used to examine Fagus sylvatica crown characteristics.•Crowns in mixed stands were wider and lower-reaching than those in pure stands.•Magnitude of mixing effect ...increased in the two sites with higher productivity.•Innovative TLS metrics revealed site effects not captured by traditional metrics.
Competition with neighboring trees of different species can affect crown size and shape. However, whether intra-specific differences in crown characteristics in mixed stands compared to pure stands are dependent on site conditions remains poorly understood. We used terrestrial laser scanning (TLS) to examine the differences in Fagus sylvatica crown characteristics at four sites, each of which contained pure stands of F. sylvatica and their mixture with Pinus sylvestris. These sites covered the area where the mixture occurs in Europe from south to north, representing a gradient of F. sylvatica productivity, defined as the mean increment of annual volume growth in pure F. sylvatica stands. Despite the large range in productivity, F. sylvatica trees in mixtures had larger crowns regardless of site conditions, with a higher proportion of their crown volume in the lower canopy compared to trees in pure stands. Larger crown volumes were related to higher live crown ratios and greater crown expansion, depending on the site. The magnitude of the mixing effect was variable among the crown characteristics evaluated, but overall our findings provide evidence that for a given species combination and density, the effect of mixture increased in the two most productive sites. TLS-derived novel crown metrics revealed that the mixing effect was affected by productivity, which was not captured by traditionally measured crown variables.
How seedling mortality and browsing affects species composition of regenerating forests has been mostly studied on a small scale. Yet, large-scale analyses based on extensive data are essential for ...robust prediction of species composition in young forests. In this study, we used a dataset from a national inventory of young forests (1-4 metres in height) to investigate the species composition of young forests across Sweden. We found that most of the regenerated forest area (almost 90%) was planted with Norway spruce (southern Sweden) and Scots pine (northern Sweden). Regeneration of Norway spruce was generally relatively successful but as a consequence of seedling mortality and browsing, almost 40% of the area regenerated with Scots pine will probably not develop into pine-dominated stands. Thus, low survival of Scots pine seedlings and trees can profoundly change the trajectory of species composition of the young forest from what was originally intended, and a large proportion of the young stands may develop into mixtures of conifers and broadleaves. While such mixtures may benefit certain biodiversity and ecosystem services, a loss of Scots pine dominated stands may also have adverse impacts on the economic returns as well as pine-dependent biodiversity and recreational values.
Airborne laser scanning (ALS) data is increasingly being used for accurate estimations of forest inventory attributes, such as tree height and volume. However, determining tree species information, ...an important attribute required by inventories for a range of forest management applications, is challenging to extract from ALS data. This is due to complex species assemblages and vertical structures in certain forest environments, and a lack of spectral information for most ALS sensors. Our objective was to estimate tree species compositions in a Canadian boreal forest environment using ALS data and point-based deep learning techniques. To accomplish this, we utilised existing polygonal forest resource information to develop a large comprehensive dataset of tree species compositions across a 630,000 ha forest management area. We then used an adapted PointAugment generative adversarial network (GAN) coupled with the dynamic graph convolutional neural network to estimate tree species proportions. This innovative deep learning approach resulted in an overall R2adj of 0.61 for all tree species proportions. A weighted F1 score of 63% was observed when classifying the leading species of a plot, 66% for leading genus, and 85% when distinguishing between coniferous and deciduous dominated plots. Furthermore, the PointAugment GAN successfully generated augmented point clouds of forest plots which can alleviate issues associated with limited training samples for use in deep learning. This approach demonstrates the capability of point-based deep learning techniques to accurately estimate tree species compositions from ALS point clouds within an expansive forested region, characterized by a complex management history and a diversity of tree species. The source code along with the pretrained models are available at https://github.com/Brent-Murray/point-dl/tree/main/Pytorch/models/PointAugment.
Understanding the interplay between environmental factors contributing to treeline formation and how these factors influence different life stages remains a major research challenge. We used an ...afforestation experiment including 92 000 trees to investigate the spatial and temporal dynamics of tree mortality and growth at treeline in the Swiss Alps. Seedlings of three high-elevation conifer species (
Larix decidua
,
Pinus mugo
ssp.
uncinata
, and
Pinus cembra
) were systematically planted along an altitudinal gradient at and above the current treeline (2075 to 2230 m above sea level a.s.l.) in 1975 and closely monitored during the following 30 years. We used decision-tree models and generalized additive models to identify patterns in mortality and growth along gradients in elevation, snow duration, wind speed, and solar radiation, and to quantify interactions between the different variables. For all three species, snowmelt date was always the most important environmental factor influencing mortality, and elevation was always the most important factor for growth over the entire period studied. Individuals of all species survived at the highest point of the afforestation for more than 30 years, although mortality was greater above 2160 m a.s.l., 50-100 m above the current treeline. Optimal conditions for height growth differed from those for survival in all three species: early snowmelt (ca. day of year 125-140 where day 1 is 1 January) yielded lowest mortality rates, but relatively later snowmelt (ca. day 145-150) yielded highest growth rates. Although snowmelt and elevation were important throughout all life stages of the trees, the importance of radiation decreased over time and that of wind speed increased. Our findings provide experimental evidence that tree survival and height growth require different environmental conditions and that even small changes in the duration of snow cover, in addition to changes in temperature, can strongly impact tree survival and growth patterns at treeline. Further, our results show that the relative importance of different environmental variables for tree seedlings changes during the juvenile phase as they grow taller.
•We curate long-term datasets from a treeline research site in the European Alps.•We synthesise 50 years of monitoring and experimental research on treeline ecology.•Our findings confirm the crucial ...role of tree growth and establishment limitations.•This research improves understanding of global change impacts on treeline ecotones.•These high-quality, easily accessible datasets foster transdisciplinary syntheses.
Understanding the dynamics of treeline ecotones under global change requires long-term ecological and environmental data. The Stillberg ecological treeline research site in the Swiss Alps was established in 1975 by planting 92,000 seedlings of Larix decidua, Pinus cembra and Pinus mugo ssp. uncinata, and has been continuously monitored since then. Here, we present curated long-term data acquired over almost 50 years at the Stillberg site, and we synthesise the major research findings. The long-term datasets comprise 6.5 million ecological and environmental records from the 40-year afforestation experiment, as well as from a 9-year free-air CO2 enrichment experiment crossed with a 6-year soil warming experiment, a 12-year nutrient addition experiment and an 8-year multifactorial tree seedling recruitment experiment. Our datasets further include 38 million records of 25 meteorological parameters measured at an hourly resolution from 1975 to 1996, and at a 10 min resolution since 1997. We provide all datasets and the corresponding metadata as open research data. Almost five decades of research in this treeline ecotone showed high mortality after tree establishment that was closely related to microclimatic variability. The two Pinus species survived at a much lower rate than L. decidua, due to indirect pathogen interactions. Furthermore, CO2 enrichment only increased growth of L. decidua, while warming increased growth of P. mugo ssp. uncinata and two Vaccinium shrub species. Enhanced nutrient availability stimulated growth in tree and understorey shrub species. In addition, soil warming and CO2 enrichment stimulated microbial activity and decreased soil carbon stocks. These findings improve our understanding of ecological processes in the treeline ecotone under global change and confirm the importance of tree growth and establishment limitations. The enhanced availability and quality of these long-term data are expected to foster whole-system approaches and transdisciplinary research syntheses, supporting the development of effective global change adaptation strategies.
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•Volume increased with increases in pine proportion and at the highest density.•Lodgepole pine had larger diameter at breast height and larger crowns in mixtures.•Neither mixture nor density affected ...gall rust damage in lodgepole pine.•Either mixture or higher spruce density reduced attack by weevil on spruce.•Mixing may increase spruce lumber quality and value by reducing damage.
Monocultures tend to yield higher total stand volumes and are simple to manage. Yet, mixed species stands may result in similar stand volumes while providing benefits such as mitigating damage from insects and disease. To understand the effects of stand density and species mixture and their interactions on stand yield, tree size and morphology, and damage in monocultures and mixtures, we analyzed a 25-year-old experiment in interior British Columbia, Canada. The lodgepole pine (Pl)-interior hybrid spruce (Sx) experiment included three densities—1000, 1500, and 2000 stems per hectare (SPH)—and five species mixtures—1:0, 3:1, 1:1, 1:3, and 0:1 Pl:Sx. Results 25 years after stand establishment showed that stand volume was significantly larger with an increasing proportion of Pl across all stand densities. Pl had 10% larger diameters in the 1000 SPH than in the 2000 SPH and when mixed with Sx (1:1). Pl had larger crowns in mixtures regardless of density. Mixture proportion did not affect gall rust incidence or stem form in Pl, but reduced attack in Sx by spruce weevil. Our findings suggest that mixing Pl-Sx and high planting density decrease weevil attacks in Sx, which reduce loss in timber quality. Yet, Pl quality may decrease when mixed with Sx, due to larger Pl crowns. These results may be used to improve the implementation of management strategies that decrease trade-offs between yields, desired market tree sizes, and timber loss from pest and pathogens, while making the stands more resilient to further climate change impact.
We assessed the impacts of three approaches to thinning from below with varying spatial patterns on several stand and individual tree variables for interior Douglas-fir ( Pseudotsuga menziesii var. ...glauca (Beissn.) Franco), interior spruce ( Picea glauca (Moench) Voss × Picea engelmannii Engelm.), and lodgepole pine ( Pinus contorta Dougl. Ex Loud. var. latifolia Englem.) in central British Columbia, Canada. The three thinning treatments were two experimental “clumped” treatments (3 m Clumped and 5 m Clumped) and the Standard (more uniform spacing) thinning treatment that was employed operationally at that time. We used long-term data from 24 plots measured five times over 21 years. Thinning increased stand basal area increment, with the plots that received the 5 m Clumped treatment having significantly higher periodic annual relative basal area increment than the unthinned Control plots. The responses for the two clumped treatments were not any lower than the Standard. The 3 m Clumped treatment was best if one is concerned about fast recovery of the growing space; however, the 5 m Clumped spacing treatment may be preferable if higher individual tree vigour is needed for resistance and resilience to fire, insects, and disease.
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•Size-asymmetric and inter-specific competition drove tree growth in mixtures.•Tree growth and size of Scots pine was higher in mixtures compared to monocultures.•Drought drove tree ...growth for both species, but temperature only affects pine.•Weather conditions modified inter-specific tree competition for both species.•Growth reduction of Norway spruce in mixtures may be offset by faster Scots pine growth.
Mixed forests are suggested as a strategic adaptation of forest management to climate change. Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) are tree species of high economic and ecological value for European forestry. Both species coexist naturally in a large part of their distributions but there is a lack of knowledge on the ecological functioning of mixtures of these species and how to manage such stands. This paper analyses these species’ intra- and inter-specific competition, including size-symmetric vs. size-asymmetric competition, and explore the effect of weather conditions on tree growth and competition. We studied basal area growth at tree level for Scots pine and Norway spruce in mixed versus pure stands in 22 triplets of fully-stocked plots along a broad range of ecological conditions across Europe. Stand inventory and increment cores provided insights into how species mixing modifies tree growth compared with neighbouring pure stands. Five different competition indices, weather variables and their interactions were included and checked in basal area growth models using a linear mixed model approach. Interspecific size-asymmetric competition strongly influenced growth for both tree species, and was modulated by weather conditions. However, species height stratification in mixed stands resulted in a greater tree basal area growth of Scots pine (10.5 cm2 year−1) than in pure stands (9.3 cm2 year−1), as this species occupies the upper canopy layer. Scots pine growth depended on temperature and drought, whereas Norway spruce growth was influenced only by drought. Interspecific site-asymmetric competition increased in cold winters for Scots pine, and decreased after a drought year for Norway spruce. Although mixtures of these species may reduce tree size for Norway spruce, our results suggest that this could be offset by faster growth in Scots pine. How inter-specific competition and weather conditions alter tree growth may have strong implications for the management of Scots pine-Norway spruce mixtures along the rotation period into the ongoing climate change scenario.
1. There is increasing evidence that species diversity enhances the temporal stability (TS) of community productivity in different ecosystems, although its effect at the population and tree levels ...seems to be negative or neutral. Asynchrony in species responses to environmental conditions was found to be one of the main drivers of this stabilizing process. However, the effect of species mixing on the stability of productivity, and the relative importance of the associated mechanisms, remain poorly understood in forest communities. 2. We investigated the way mixing species influenced the TS of productivity in Pinus sylvestris L. and Fagus sylvatica L. forests, and attempted to determine the main drivers among overyielding, asynchrony between species annual growth responses to environmental conditions, and temporal shifts in species interactions. We used a network of 93 experimental plots distributed across Europe to compare the TS of basal area growth over a 15-year period (1999-2013) in mixed and monospecific forest stands at different organizational levels, namely the community, population and individual tree levels. 3. Mixed stands showed a higher TS of basal area growth than monospecific stands at the community level, but not at the population or individual tree levels. The TS at the community level was related to asynchrony between species growth in mixtures, but not to overyielding nor to asynchrony between species growth in monospecific stands. Temporal shifts in species interactions were also related to asynchrony and to the mixing effect on the TS. 4. Synthesis. Our findings confirm that species mixing can stabilize productivity at the community level, whereas there is a neutral or negative effect on stability at the population and individual tree levels. The contrasting findings regarding the relationships between the temporal stability and asynchrony in species growth in mixed and monospecific stands suggest that the main driver in the stabilizing process may be the temporal niche complementarity between species rather than differences in species' intrinsic responses to environmental conditions.
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