Effect of forest canopy composition on soil nutrients and dynamics of the understorey: mixed canopies serve neither vascular nor bryophyte strata

Question: The effect of overstorey composition on above‐ground dynamics of understorey vegetation is poorly understood. This study examines the understorey biomass, production and turnover rates of vascular and non‐vascular plants along a conifer–broadleaf gradient of resource availability and heterogeneity. Location: Canadian boreal forests of northwest Quebec and Ontario. Methods: We sampled mature stands containing various proportions of black spruce (Picea mariana (Mill.) BSP), trembling aspen (Populus tremuloides Michx.) and jack pine (Pinus banksiana Lamb.). Above‐ground biomass of the understorey vegetation was assessed through harvesting; annual growth rates were calculated as the differences between biomass in 2007 and 2008, as estimated by allometric relationships, and turnover rates were estimated as net primary production divided by the biomass in 2007. Results: Higher aspen presence, linked to greater nutrient availability in the forest floor, was generally associated with higher vascular biomass and production in the understorey. This effect was less pronounced in sites of high intrinsic fertility. In contrast, bryophyte biomass was positively associated with conifer abundance, particularly in wet sites of the Quebec study area. Non‐linear responses resulted in total understorey biomass being lower under mixed canopies than under pure aspen or pure conifer canopies. Turnover rates did not differ with overstorey composition. Conclusions: While resource availability is a main driver of understorey productivity, resources as drivers appear to differ with differences in understorey strata components, i.e. vascular versus non‐vascular plants. Resource heterogeneity induced by a mixed canopy had overall negative effects on understorey above‐ground productivity, as this productivity seemed to rely on species adapted to the specific conditions induced by a pure canopy.     

The Contrasting Effects of Aspen and Jack Pine on Soil Nutritional Properties Depend on Parent Material

Abstract The influence of forest stand composition on soil was investigated by comparing the forest floor (FH) and upper mineral soil (0–20 cm) nutritional properties of jack pine and aspen stands on two soil types of contrasting fertility, a coarse-textured and a fine-textured deposit, in a replicated design. The studied tree species are pioneers that are found after major disturbances in the southern boreal forest of western Quebec and that differ in their nutrient requirements but not in their growth rate. Soil organic matter as well as total and available N, P, K, Ca, Mg contents were determined and the relationships with nutrient accumulation in tree biomass were studied. On both soil types a greater total and available nutrient accumulation in the forest floor layer was observed in aspen than in jack pine whereas such differences between stand types could not be detected in the mineral soil. Differences in FH nutrient content between stand types were larger on coarse deposits than on fine-textured soils. These results support the hypothesis that tree species with greater nutrient requirements cause an enrichment of the surface soil at least in the short term. The modulation of tree species effect by soil type was contrary to the pattern observed in other studies since a greater expression of this effect was observed on poorer soils. Differences in soil nutrient content were related to levels of organic matter accumulation.     

Growth and mortality of trembling aspen (Populus tremuloides) in response to artificial defoliation

To simulate the effects of forest tent caterpillar (FTC) defoliation on trembling aspen growth and mortality, an artificial defoliation experiment was performed over three years in young aspen stands of northwestern Quebec. Defoliation plots of 15 × 15 m were established on three sites, together with associated control stands of pure trembling aspen. In 2007, root collar diameters were measured and positions of all trees were mapped prior defoliation. Severe FTC defoliation was simulated for three successive years (2007–2009) by manually removing all leaves from all but 7–10% of the trees present in the defoliation plots. Yearly surveys of growth and mortality were conducted until 2010 to evaluate defoliation effects on defoliated as well as surrounding undefoliated trees. In absence of other factors, growth and mortality of trembling aspen decreased and increased, respectively, after defoliation. Our study further revealed that small diameter trees died after one year of artificial defoliation, while larger-diameter trees died after repeated defoliations. Distributions of tree mortality tended to be aggregated at small scales (<5 m), corroborating gap patterns observed in mature stands following FTC outbreaks. This experiment revealed that trembling aspen mortality can be directly attributed solely to defoliation. Repeated defoliations during FTC outbreaks have the potential to profoundly modify stand productivity and structure by reducing tree growth and increasing tree mortality in the absence of predisposing factors.     

Climate‐diameter growth relationships of black spruce and jack pine trees in boreal Ontario,Canada

To predict the long‐term effects of climate change – global warming and changes in precipitation – on the diameter (radial) growth of jack pine (Pinus banksiana Lamb.) and black spruce (Picea mariana [Mill.] B.S.P.) trees in boreal Ontario, we modified an existing diameter growth model to include climate variables. Diameter chronologies of 927 jack pine and 1173 black spruce trees, growing in the area from 47°N to 50°N and 80°W to 92°W, were used to develop diameter growth models in a nonlinear mixed‐effects approach. Our results showed that the variables long‐term average of mean growing season temperature, precipitation during wettest quarter, and total precipitation during growing season were significant (alpha = 0.05) in explaining variation in diameter growth of the sample trees. Model results indicated that higher temperatures during the growing season would increase the diameter growth of jack pine trees, but decrease that of black spruce trees. More precipitation during the wettest quarter would favor the diameter growth of both species. On the other hand, a wetter growing season, which may decrease radiation inputs, increase nutrient leaching, and reduce the decomposition rate, would reduce the diameter growth of both species. Moreover, our results indicated that future (2041–2070) diameter growth rate may differ from current (1971–2000) growth rates for both species, with conditions being more favorable for jack pine than black spruce trees. Expected future changes in the growth rate of boreal trees need to be considered in forest management decisions. We recommend that knowledge of climate–growth relationships, as represented by models, be combined with learning from adaptive management to reduce the risks and uncertainties associated with forest management decisions.     

树木年龄和断面积对加拿大北方林树木死亡率的影响

以加拿大北部的杨树(Populus spp.)、斑克松(Pinus banksiana)、黑云杉(Picea mariana)为对象,采用长期定位试验,对134块固定样地的活立木及枯死木进行调查,并运用线性回归的方法研究树木年龄、断面积和林分类型对3种树木死亡率的影响.结果表明:随着树龄和断面积的增加,林木的死亡率呈上升趋势.杨树在斑克松林中的死亡率较高,而在黑云杉林中死亡率较低.在黑云杉林中,树龄是影响斑克松死亡率的主要因子;而在杨树林中,断面积是影响斑克松死亡率的重要因子;不同林分类型中树龄对黑云杉死亡率的影响显著.树种组成对树种的死亡率有显著影响;树木年龄、断面积和林分类型之间的交互效应对各树种的死亡率均有显著影响;不同林分类型中同一树种的死亡状况有明显差异.     

The effects of Laccaria proxima and fibrous pulp waste on the growth of nine container-grown conifer seedling species

 Japanese larch (Larix kaempferi), white spruce (Picea glauca), black spruce (Picea mariana), red spruce (Picea rubens), jack pine (Pinus banksiana), mugo pine (Pinus mugo), red pine (Pinus resinosa), Japanese black pine (Pinus thunbergii) and Douglas-fir (Pseudotsuga menziesii var. menziesii), were inoculated to test the effective host range of the ectomycorrhizal fungus Laccaria proxima and the possibility of utilizing pulp waste as a potting medium for containerized seedling production. Laccaria proxima tended to improve the container growth of Japanese black pine and white spruce, and significantly improved that of jack pine, mugo pine, black spruce, red spruce and Douglasfir. The growth of red pine and Japanese larch were only slightly improved with L. proxima. Pulp waste (33% by volume) had negative effects on tree seedling growth, except for Douglasfir (no significant effect). The interactions of Laccaria proxima and pulp waste varied; the hosts were significantly positive (P<0.01) in the case of jack pine and black spruce, but there was no significant effect for the rest. Negative effects were found with Japanese black pine. Use of pulp waste in seedling production of jack pine, black spruce, mugo pine, red spruce and Douglasfir inoculated with L. proxima and of Japanese black pine both with and without L. proxima is feasible, but further research is necessary to determine the optimal percentage of pulp waste that can be utilized in seedling production of tree species and the field performance of these seedlings. Accepted: 30 August 1995     

Modelling Variable Fire Severity in Boreal Forests: Effects of Fire Intensity and Stand Structure

It is becoming clear that fires in boreal forests are not uniformly stand-replacing. On the contrary, marked variation in fire severity, measured as tree mortality, has been found both within and among individual fires. It is important to understand the conditions under which this variation can arise. We integrated forest sample plot data, tree allometries and historical forest fire records within a diameter class-structured model of 1.0 ha patches of mono-specific black spruce and jack pine stands in northern Québec, Canada. The model accounts for crown fire initiation and vertical spread into the canopy. It uses empirical relations between fire intensity, scorch height, the percent of crown scorched and tree mortality to simulate fire severity, specifically the percent reduction in patch basal area due to fire-caused mortality. A random forest and a regression tree analysis of a large random sample of simulated fires were used to test for an effect of fireline intensity, stand structure, species composition and pyrogeographic regions on resultant severity. Severity increased with intensity and was lower for jack pine stands. The proportion of simulated fires that burned at high severity (e.g. >75% reduction in patch basal area) was 0.80 for black spruce and 0.11 for jack pine. We identified thresholds in intensity below which there was a marked sensitivity of simulated fire severity to stand structure, and to interactions between intensity and structure. We found no evidence for a residual effect of pyrogeographic region on simulated severity, after the effects of stand structure and species composition were accounted for. The model presented here was able to produce variation in fire severity under a range of fire intensity conditions. This suggests that variation in stand structure is one of the factors causing the observed variation in boreal fire severity.     

Spatial and temporal variations in boreal forest fire frequency in northern Alberta

Abstract. Spatial and temporal variations in fire frequency in the boreal forest of Wood Buffalo National Park (WBNP) were assessed using forest stand age, fire scar and historical data. I test the hypotheses that (1) fire frequency is higher in jack pine forests and aspen forests than in black spruce forests and white spruce forests, (2) these variations in fire frequency can be related to the mean waterbreak distance (MWD) around a site and (3) fire frequency has changed over the past 300 years. The fire cycles (the time required to burn an area equal in size to the entire study area) in jack pine forests (39 years) and in aspen forests (39 years) were significantly shorter than those in black spruce forests (78 years) and in white spruce forests (96 years). The length of the fire cycle varies inversely with the MWD around a site, and the MWD was significantly higher in jack pine and aspen forests than in black or white spruce forests. It is suggested that covariations between soil type and the MWD influence, respectively, variations in forest dominant and fire frequency. A change in fire frequency at 1860 was apparent in the fire history for all of WBNP, the black spruce dominated stands, and the near and medium MWD classes. The fire cycle estimates for these classes were all significantly shorter during the period 1750 to 1859 (fire cycles = 25–49 years) than they were in the period 1860 to 1989 (fire cycles = 59–89 years). The possible roles of changes in climate and aboriginal burning practices in causing the temporal change in fire frequency are discussed.     

Soil CO2 efflux in contrasting boreal deciduous and coniferous stands and its contribution to the ecosystem carbon balance

Similar nonsteady‐state automated chamber systems were used to measure and partition soil CO₂ efflux in contrasting deciduous (trembling aspen) and coniferous (black spruce and jack pine) stands located within 100 km of each other near the southern edge of the Boreal forest in Canada. The stands were exposed to similar climate forcing in 2003, including marked seasonal variations in soil water availability, which provided a unique opportunity to investigate the influence of climate and stand characteristics on soil CO₂ efflux and to quantify its contribution to the net ecosystem CO₂ exchange (NEE) as measured with the eddy‐covariance technique. Partitioning of soil CO₂ efflux between soil respiration (including forest‐floor vegetation) and forest‐floor photosynthesis showed that short‐ and long‐term temporal variations of soil CO₂ efflux were related to the influence of (1) soil temperature and water content on soil respiration and (2) below‐canopy light availability, plant water status and forest‐floor plant species composition on forest‐floor photosynthesis. Overall, the three stands were weak to moderate sinks for CO₂ in 2003 (NEE of ?103, ?80 and ?28 g C m^?2 yr^?1 for aspen, black spruce and jack pine, respectively). Forest‐floor respiration accounted for 86%, 73% and 75% of annual ecosystem respiration, in the three respective stands, while forest‐floor photosynthesis contributed to 11% and 14% of annual gross ecosystem photosynthesis in the black spruce and jack pine stands, respectively. The results emphasize the need to perform concomitant measurements of NEE and soil CO₂ efflux at longer time scales in different ecosystems in order to better understand the impacts of future interannual climate variability and vegetation dynamics associated with climate change on each component of the carbon balance.     

Neighborhood effects on soil properties,mycorrhizal attributes,tree growth,and nutrient status in afforested zones

A harmonious interspecies relationship is the key to the success of mixed afforestation. This study was conducted to assess the responses of afforestation species to their neighboring trees. We examined five types of stands—monocultures of Chinese pine (Pinus tabuliformis), black locust (Robinia pseudoacacia), sea‐buckthorn (Hippophae rhamnoides), and two mixtures (Chinese pine × black locust mixture and Chinese pine × sea‐buckthorn mixture)—in the Loess Plateau, northwestern China. The height and diameter at breast height of each tree species were measured, and rhizosphere soil, shoot, and root were sampled. In monocultures, black locust was taller than Chinese pine and sea‐buckthorn, while the height of Chinese pine and sea‐buckthorn was similar. In mixtures, Chinese pine grew better with sea‐buckthorn than alone as a result of modified soil properties and plant nutrition, but not with black locust. When Chinese pine was used as neighbors, it affected the level of arbuscular mycorrhizal (AM) colonization of black locust, soil properties and AM fungal spore density of black locust and sea‐buckthorn, but did not significantly affect their growth. Our results suggest that the reciprocal effects between tree species in mixture are not symmetric, and thus planning for efficient mixed afforestation requires knowledge of species‐specific growth rate, nutrient requirements, and species interactions.     

Forest floor properties across sharp compositional boundaries separating trembling aspen and jack pine stands in the southern boreal forest

Jack pine and trembling aspen are two early-seral boreal tree species with contrasting nutrient cycling strategies. Both species may form adjacent mono-specific stands separated by sharp compositional boundaries. We hypothesized that such boundaries result in wider functional ecotones. Spatial transitions in humus forms, forest floor chemistry and microbial communities were assessed across 32 m long transects set perpendicular to sharp compositional boundaries separating four jack pine and aspen stands. Split moving window analysis (SMWA) and moving window regression analysis (MWRA) were used to locate functional boundaries and ecotones. We found a gradual transition from moder (aspen) to mor (jack pine) humus spanning 16 m across the compositional boundary. An abrupt increase in forest floor water content at 3 m within jack pine stands was possibly due to aspen roots foraging for water beyond the boundary. The functional boundary and associated ecotone for forest floor pH, C:N ratio, Mg and ammonification were skewed toward jack pine stands, likely the result of aspen leaf dispersal. Low nitrification rates throughout jack pine stands and up to 11 m into aspen stands suggested that jack pine roots might extend far within aspen stands and produce metabolites that suppress nitrification. SMWA performed on the multivariate dataset of microbial fatty acids (FAs) revealed three distinct forest floor microbial communities that were skewed toward jack pine stands. Pine-type communities were associated to fungal FAs, pine-type and transition-type communities to non-fungal eukaryotic FAs, and aspen-type communities to bacterial FAs. Taken collectively, our data delimit a 24 m wide functional ecotone straddling sharp compositional boundaries separating trembling aspen and jack pine stands. We conclude that the functional diversity of boreal landscapes, where adjacent mono-specific stands are prevalent, is related to the patchiness of the landscape.     

The influence of recent climate change on tree height growth differs with species and spatial environment

Tree growth has been reported to increase in response to recent global climate change in controlled and semi-controlled experiments, but few studies have reported response of tree growth to increased temperature and atmospheric carbon dioxide (CO₂) concentration in natural environments. This study addresses how recent global climate change has affected height growth of trembling aspen (Populus tremuloides Michx) and black spruce (Picea mariana Mill B.S.) in their natural environments. We sampled 145 stands dominated by aspen and 82 dominated by spruce over the entire range of their distributions in British Columbia, Canada. These stands were established naturally after fire between the 19^th and 20^th centuries. Height growth was quantified as total heights of sampled dominant and co-dominant trees at breast-height age of 50 years. We assessed the relationships between 50-year height growth and environmental factors at both spatial and temporal scales. We also tested whether the tree growth associated with global climate change differed with spatial environment (latitude, longitude and elevation). As expected, height growth of both species was positively related to temperature variables at the regional scale and with soil moisture and nutrient availability at the local scale. While height growth of trembling aspen was not significantly related to any of the temporal variables we examined, that of black spruce increased significantly with stand establishment date, the anomaly of the average maximum summer temperature between May-August, and atmospheric CO₂ concentration, but not with the Palmer Drought Severity Index. Furthermore, the increase of spruce height growth associated with recent climate change was higher in the western than in eastern part of British Columbia. This study demonstrates that the response of height growth to recent climate change, i.e., increasing temperature and atmospheric CO₂ concentration, did not only differ with tree species, but also their growing spatial environment.     

Temporal variation of competition and facilitation in mixed species forests in Central Europe

Facilitation, reduced competition or increased competition can arise in mixed stands and become essential to the performance of these stands when compared to pure stands. Facilitation and over‐yielding are widely held to prevail on poor sites, whereas neutral interactions or competition, leading to under‐yielding of mixed versus pure stands, can occur on fertile sites. While previous studies have focused on the spatial variation of mixing effects, we examine the temporal variation of facilitation and competition and its effect on growth. The study is based on tree ring measurement on cores from increment borings from 559 trees of Norway spruce (Picea abies [L.] Karst.), European beech (Fagus sylvatica [L.]) and sessile oak (Quercus petraea (Matt.) Liebl.) in southern Germany, half of which were in pure stands and half in adjacent mixed stands. Mean basal area growth indices were calculated from tree ring measurements for pure and mixed stands for every species and site. The temporal variation, with positive correlations between species‐specific growth indices during periods of low growth and neutral or negative correlations during periods of high growth, is more distinct in mixed than in neighbouring pure stands. We provide evidence that years with low growth trigger over‐yielding of trees in mixed as opposed to pure stands, while years with high growth lead to under‐yielding. We discuss the relevance of the results in terms of advancing our understanding and modelling of mixed stands, extension of the stress gradient hypothesis, and the performance of mixed versus pure stands in the face of climate change.     

Ectomycorrhizal colonization on black spruce and jack pine seedlings outplanted in reforestation sites

Six-week-old, mycorrhiza-free, bareroot jack pine and black spruce seedlings were outplanted in ten reforestation sites, situated between 45–48° latitude N and 69–74° longitude W, within the province of Quebec, representing diverse operational forestry disturbances and ecological conditions. Two months after outplanting, root systems of black spruce seedlings had fewer mycorrhizae than those of jack pine seedlings. Ectomycorrhizal colonization on black spruce seedlings did not vary significantly with the reforestation site. Percent mycorrhizal colonization for these seedlings was positively correlated with seedling dry weight while with the jack pine seedlings, mycorrhizal colonization varied significantly with the outplanting site and there was no correlation between mycorrhizal formation and seedling dry weight. Multiple linear regressions showed pH to be a determinant soil factor for mycorrhizal colonization for the two species. Drainage was the other influential factor affecting colonization of black spruce while organic matter accumulation was more important for jack pine. Inoculation with selected ectomycorrhizal fungi could be more important for black spruce than for jack pine seedlings.     

Changes in diversity and storage function of ectomycorrhiza and soil organoprofile dynamics after introduction of beech into Scots pine forests

Diversity and storage function of mycorrhiza as well as soil organoprofile formation were investigated in a chronosequence of a pure Scots pine (Pinus sylvestris L.) stand, of Scots pine stands that were underplanted with beech (Fagus sylvatica L.) and in three pure beech stands of different age. Mycorrhiza diversity was higher in the pure beech stands compared to the pure pine stand. Beech and pine trees in the mixed stands had similar dominant mycorrhiza morphotypes. However, trees in two of the three pure beech stands were mycorrhized with other types. Mycorrhizal abundance and nutrient amounts of mycorrhizae associated with beech trees were higher in the mixed and in the pure beech stands compared to pine mycorrhizae indicating that nutrient uptake was higher in older beech than in older pine trees. Humus quality varied from pine to beech stands. Plant litter storage in the humus layer was highest in the youngest mixed stand and lowest in the oldest beech stand. Humus forms changed from moder grass-type in the pure Scots pine stand to mor-like moder and moder rich in fine humus with increasing age of beeches in the mixed stands. The older beech stands were characterised by oligomull and mull-like moder as the dominating humus forms. The ecologically favourable humus forms, i.e., nutrient rich humus forms in the older beech stands correlate well with the higher mycorrhizal diversity and abundance as well as the higher nutrient storage of their mycorrhizae in these stands. The results are also discussed with regard to the 'base-pump effect' of beech trees.     

Tree Species Traits Influence Soil Physical,Chemical, and Biological Properties in High Elevation Forests

Background

Previous studies have shown that plants often have species-specific effects on soil properties. In high elevation forests in the Southern Rocky Mountains, North America, areas that are dominated by a single tree species are often adjacent to areas dominated by another tree species. Here, we assessed soil properties beneath adjacent stands of trembling aspen, lodgepole pine, and Engelmann spruce, which are dominant tree species in this region and are distributed widely in North America. We hypothesized that soil properties would differ among stands dominated by different tree species and expected that aspen stands would have higher soil temperatures due to their open structure, which, combined with higher quality litter, would result in increased soil respiration rates, nitrogen availability, and microbial biomass, and differences in soil faunal community composition.

Methodology/Principal Findings

We assessed soil physical, chemical, and biological properties at four sites where stands of aspen, pine, and spruce occurred in close proximity to one-another in the San Juan Mountains, Colorado. Leaf litter quality differed among the tree species, with the highest nitrogen (N) concentration and lowest lignin∶N in aspen litter. Nitrogen concentration was similar in pine and spruce litter, but lignin∶N was highest in pine litter. Soil temperature and moisture were highest in aspen stands, which, in combination with higher litter quality, probably contributed to faster soil respiration rates from stands of aspen. Soil carbon and N content, ammonium concentration, and microbial biomass did not differ among tree species, but nitrate concentration was highest in aspen soil and lowest in spruce soil. In addition, soil fungal, bacterial, and nematode community composition and rotifer, collembolan, and mesostigmatid mite abundance differed among the tree species, while the total abundance of nematodes, tardigrades, oribatid mites, and prostigmatid mites did not.

Conclusions/Significance

Although some soil characteristics were unaffected by tree species identity, our results clearly demonstrate that these dominant tree species are associated with soils that differ in several physical, chemical, and biotic properties. Ongoing environmental changes in this region, e.g. changes in fire regime, frequency of insect outbreaks, changes in precipitation patterns and snowpack, and land-use change, may alter the relative abundance of these tree species over coming decades, which in turn will likely alter the soils.     

Competition and regeneration in quaking aspen–white fir (Populus tremuloides–Abies concolor) forests in the Northern Sierra Nevada,USA

Question: How does competition between quaking aspen (Populus tremuloides) and white fir (Abies concolor) affect growth and spatial pattern of each species? Location: The northern Sierra Nevada, California, USA. Methods: In paired plots in mixed aspen‐ (n=3) or white fir‐dominated (n=2) stands, we mapped trees and saplings and recorded DBH, height, species, and condition and took increment cores. We tallied seedlings by species. Tree ring widths were used as a measure of basal area change over the last decade, and canopy openness was identified using hemispherical photographs. Linear mixed models were used to relate neighborhood indices of competition, stand, and tree‐level variables to diameter increment. Spatial patterns of stems were identified using the Neighborhood Density Function. Results: White fir radial growth was higher in aspen‐ than white fir‐dominated plots. Individual‐level variables were more important for white fir than for aspen growth, while variables representing competitive neighborhood were important only for aspen. The forest canopy was more open in aspen‐ than white fir‐dominated stands, but ample aspen seedlings were observed in all stands. Canopy stems of aspen and white fir were randomly distributed, but saplings and small trees were clumped. Aspen saplings were repelled by canopy aspen stems. Conclusions: Variation in canopy openness explained more stand–stand variation in white fir than aspen growth, but high light levels were correlated with recruitment of aspen seedlings to the sapling class. Radial growth of aspen was predicted by indices of neighborhood competition but not radial growth of white fir, indicating that spacing and stem arrangement was more important for aspen than white fir growth. Fire suppression has removed a major disturbance mechanism that promoted aspen persistence and reduced competition from encroaching conifers, and current forests favor species that regenerate best by advance regeneration (white fir).     

Taxonomy,together with ontogeny and growing conditions,drives needleleaf species' sensitivity to climate in boreal North America

Currently, there is no consensus regarding the way that changes in climate will affect boreal forest growth, where warming is occurring faster than in other biomes. Some studies suggest negative effects due to drought‐induced stresses, while others provide evidence of increased growth rates due to a longer growing season. Studies focusing on the effects of environmental conditions on growth–climate relationships are usually limited to small sampling areas that do not encompass the full range of environmental conditions; therefore, they only provide a limited understanding of the processes at play. Here, we studied how environmental conditions and ontogeny modulated growth trends and growth–climate relationships of black spruce (Picea mariana) and jack pine (Pinus banksiana) using an extensive dataset from a forest inventory network. We quantified the long‐term growth trends at the stand scale, based on analysis of the absolutely dated ring‐width measurements of 2,266 trees. We assessed the relationship between annual growth rates and seasonal climate variables and evaluated the effects of various explanatory variables on long‐term growth trends and growth–climate relationships. Both growth trends and growth–climate relationships were species‐specific and spatially heterogeneous. While the growth of jack pine barely increased during the study period, we observed a growth decline for black spruce which was more pronounced for older stands. This decline was likely due to a negative balance between direct growth gains induced by improved photosynthesis during hotter‐than‐average growing conditions in early summers and the loss of growth occurring the following year due to the indirect effects of late‐summer heat waves on accumulation of carbon reserves. For stands at the high end of our elevational gradient, frost damage during milder‐than‐average springs could act as an additional growth stressor. Competition and soil conditions also modified climate sensitivity, which suggests that effects of climate change will be highly heterogeneous across the boreal biome.     

Characterization of tree macroremains production in a recently burned conifer forest in northern Québec,Canada

The production of plant macroremains was studied in a conifer forest twomonths after it burned in 1996 in northern Québec. The proportions ofvarious types of charred and uncharred pieces (needles, cones andwood) produced by black spruce (Piceamariana) and jack pine (Pinusbanksiana) were determined by sampling around individual trees.Both species produced equivalent masses of charred material, but pieces of woodcharcoal from jack pine are generally larger that those of black spruce. Theproportion of charred versus uncharred needles is the bestindicator of the species dominance in the forest. Although the fall of uncharredneedles is delayed from the time of a fire, they contribute to more than half ofall remains produced. Jack pine cones remain on the tree for a long time after afire, while charred cones of black spruce are dehiscent (cones come off thebranches easily). Trees are poor wood charcoal producers compared toundergrowth shrubs. As a result, the macroremains assemblage associated with afire event is made up of large amount of uncharred material from trees and alarge proportion of charred pieces produced by undergrowth vegetation. Modernassemblages of plant macroremains indicate that in order to reconstruct pastvegetation associated with fire disturbance, it is important to distinguishbetween the various types of remains, because wood charcoal is mainly producedby material that is already dead.     

Structural crown properties of Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica [L.]) in mixed versus pure stands revealed by terrestrial laser scanning

How tree morphology develops in mixed-species stands is essential for understanding and modelling mixed-stand dynamics. However, research so far focused on the morphological variation between tree species and neglected the variation within a species depending on intra- and interspecific competition. Our study, in contrast, addresses crown properties of nine mature Norway spruces (Picea abies [L.] Karst.) of a pure stand and compares them with ten spruces growing in mixture with European beech (Fagus sylvatica [L.]). The same was done with 11 pure stand beeches and 12 beeches growing in mixture with spruce. Through application of a terrestrial laser scanner and a new skeletonization approach, we deal with both species’-specific morphological traits such as branch angle, branch length, branch bending, crown volume and space occupation of branches within the crown, some of which were hardly accessible so far. Special attention is paid to distinct differences between trees growing in mixed and pure stands: for spruce, our study reveals significantly longer branches and greater crown volumes in the mixed stand when compared to the pure stand. In case of European beech, individuals growing in mixture show flatter branch angles, more distinct ramification, greater crown volumes and a lower share of a single branch’s space occupation in the total crown volume. The results show that the presented methods yield detailed information on the morphological traits analyzed in this study and that interspecific competition on its own may have a significant impact on crown structures. Implications for production ecology and stand dynamics of mixed-species forests are discussed.