Boreal Forest Species Research Articles

Predicted range shifts in North American boreal forest birds and the effect of climate change on genetic diversity in blackpoll warblers (Setophaga striata)

As North American species’ ranges shift northward in response to climate change, populations isolated in high-elevation habitat “islands” at the southern edge of distributions are predicted to decrease in size or be extirpated. Levels of genetic structure and gene flow and the number of private alleles held within these peripheral populations can be used as a measure of the potential loss of genetic diversity due to climate change. We use GIS-based climate niche models to project geographic distributions of 15 boreal forest bird species for the year 2080 under two carbon emissions scenarios to predict the extent to which ranges will shift, leading to the extirpation of isolated populations at the southern periphery of the boreal forest. Breeding distributions of nearly all boreal bird species are predicted to expand as they shift northward, but will dramatically decrease or be completely lost from mountain populations in New York, Vermont, and New Hampshire by 2080. To examine the effect of these shifts on gene pools of migratory bird species we genotyped 178 blackpoll warblers (Setophaga striata) at nine microsatellite loci, sampling four imperiled high-elevation populations and four northern populations. In S. striata 10.4 % of microsatellite alleles were confined to populations expected to be lost due to climate change. However, these accounted for a nonsignificant percent of the genetic structure, and loss of these alleles would not significantly erode species heterozygosity or allelic richness. Our results indicate that isolated southern populations of S. striata, and possibly other migratory species with high gene flow, do not represent genetically isolated, independently evolving units. Efforts to mitigate the effect of climate change on boreal forest birds should focus on species in which peripheral populations harbor significant genetic diversity.

Dead wood and polypore diversity in natural post-fire succession forests and managed stands – Lessons for biodiversity management in boreal forests

We tested the extent to which forestry mimics natural stand-replacing fire dynamics, and what factors best explain species richness and assemblages of polyporous fungi in natural northern boreal Picea abies-dominated forests and managed forests which were either clear-cut and planted with pine, or selectively logged. The amount of coarse woody debris (CWD) and diversity of polypores in old-growth forests and in forests at three natural post-fire successional stages (young, middle-aged, old) were compared with managed forests at comparable successional stages. Fire created a high amount of CWD, providing a supply of dead wood for saproxylic species for at least 50–70y. Dead wood volumes remained high throughout the natural forest succession, exceeding at all stages 18m3ha−1, the threshold value established in the study for the occurrence of red-listed species. The richness of polypore species was highest on young post-fire sites, and their species composition, including red-listed species, was comparable to that of old-growth forests (>300y) used as references. The amount of CWD in natural post-fire forests was over two to twenty times higher than that on clear-cut sites, and species diversity on clear-cut sites was considerably lower than on sites undergoing natural succession, with no occurrences of red-listed species. A relatively diverse polypore flora was found in selectively logged forests, likely due to rather a high volume and continuity of spruce CWD. The results indicate a strong but non-linear relationship between total volume of CWD and species richness of polypores, but successional stage and naturalness also explained part of the variation. The species assemblages were influenced most by the decay stage of CWD, tree species composition and naturalness of the site. The results demonstrate the importance of old-growth and young post-fire forests as landscape-level habitats and species pools, especially for red-listed species. We recommend that old-growth spruce forests be preserved whenever possible and that high CWD volumes be generated locally for red-listed species, for example, by prescribed burning. Regeneration forests for spruce, extended rotation periods and the use of logging techniques other than clear-cutting would help in maintaining species diversity in spruce-associated species in boreal forests.

The relative photosynthetic contribution of old and new fronds of the wintergreen fern Dryopteris carthusiana, Ontario, Canada1

Goldblum, D. and M. C. Kwit (Department of Geography, Northern Illinois University, DeKalb, IL 60115). The relative photosynthetic contribution of old and new fronds of the wintergreen fern Dryopteris carthusiana, Ontario, Canada. J. Torrey Bot. Soc. 139: 270–282. 2012.—A number of understory species in temperate and boreal forests are characterized by the wintergreen habit which entails retaining leaves for a portion of a second growing season. Several hypotheses have been offered to account for this trait, most commonly: old leaves serve as storage organs for nutrients required for spring growth or old leaves contribute to the overall carbon gain of the plant given their presence in the understory during the period of high radiation prior to canopy leafout. In this study, using field-derived light response curves, plant demographic data, leaf phenological data, and twice-hourly understory light levels, we model the relative contributions of old and new fronds to overall seasonal net carbon gain of a common fern, Dryopteris carthusiana, in a boreal and deciduous forest in Ontario, Canada. Approximately 43% of the total fern carbon gain in the deciduous forest occurs in the 40 days before canopy closure, compared to 46% in the boreal forest during the same period. In the deciduous forest, approximately 30% of the total fern net carbon assimilation occurs at the end of the growing season following overstory leaf fall, which was not the case in the boreal forest. Old fronds contribute 29% of overall carbon gain in the deciduous forest, but substantially more (63%) in the boreal forest. We estimate the total net carbon assimilation by D. carthusiana ferns in the deciduous forest at 90.7 kg C ha−1 for the growing season and 45.5 kg C ha−1 in the boreal forest. Our findings quantify the importance of the wintergreen habit for overall fern seasonal carbon gain, as well as the importance of early season high light conditions for understory plants in deciduous forests.

Size-mediated tree transpiration along soil drainage gradients in a boreal black spruce forest wildfire chronosequence

Boreal forests are crucial to climate change predictions because of their large land area and ability to sequester and store carbon, which is controlled by water availability. Heterogeneity of these forests is predicted to increase with climate change through more frequent wildfires, warmer, longer growing seasons and potential drainage of forested wetlands. This study aims at quantifying controls over tree transpiration with drainage condition, stand age and species in a central Canadian black spruce boreal forest. Heat dissipation sensors were installed in 2007 and data were collected through 2008 on 118 trees (69 Picea mariana (Mill.) Britton, Sterns & Poggenb. (black spruce), 25 Populus tremuloides Michx. (trembling aspen), 19 Pinus banksiana Lamb. (jack pine), 3 Larix laricina (Du Roi) K. Koch (tamarack) and 2 Salix spp. (willow)) at four stand ages (18, 43, 77 and 157 years old) each containing a well- and poorly-drained stand. Transpiration estimates from sap flux were expressed per unit xylem area, J(S), per unit ground area, E(C) and per unit leaf area, E(L), using sapwood (A(S)) and leaf (A(L)) area calculated from stand- and species-specific allometry. Soil drainage differences in transpiration were variable; only the 43- and 157-year-old poorly-drained stands had ∼ 50% higher total stand E(C) than well-drained locations. Total stand E(C) tended to decrease with stand age after an initial increase between the 18- and 43-year-old stands. Soil drainage differences in transpiration were controlled primarily by short-term physiological drivers such as vapor pressure deficit and soil moisture whereas stand age differences were controlled by successional species shifts and changes in tree size (i.e., A(S)). Future predictions of boreal climate change must include stand age, species and soil drainage heterogeneity to avoid biased estimates of forest water loss and latent energy exchanges.

Energy density and its variation in space limit species richness of boreal forest birds

AbstractAim An area’s ability to support species may be dependent not only on the total amount of available energy it contains but also on energy density (i.e. available energy per unit area). Acknowledging these two aspects of energy availability may increase mechanistic understanding of how increased energy availability results in increased species richness. We studied the relationship between energy density, its variation in space and boreal forest bird species richness and investigated two possible mechanisms: (1) metabolic constraints of organisms, and (2) increased resource availability for specialists.Location Protected areas in Finland’s boreal forest.Methods We tested whether bird species richness was best determined by total energy availability in an area or by energy density and its variation within the area, before and after including bird abundance in the models. We evaluated two main explanatory variables: tree growth reflecting the rate of energy production and tree volume as a measure of biomass. In addition, we modelled individual species’ responses to energy density and its variation, and evaluated the prediction of the metabolic constraints hypothesis that small species are limited by energy density whereas large species are limited by total energy availability in the area.Results Energy density and its variation were good predictors of species richness: together with abundance they explained 84% of variation in species richness (compared with 74% for abundance alone). Pure metabolic constraints were unlikely to explain this relationship. Instead, the mechanism probably involved increased habitat heterogeneity benefiting specialist species. Total energy availability was also an important factor determining species richness but its effect was indirect via abundance.Main conclusions Our results corroborate the importance of energy availability as a driver of species richness in forest bird communities, and they indicate that energy density and its variation in the landscape strongly influence species richness even after accounting for abundance.

Синтаксономия вторичных лесов средних стадий сукцессий центрально-возвышенной части Южного Урала

The secondary birch and aspen forests of middle stages of succession of the central elevated part of the Southern Urals are studied. 4 subassociations, 1 community, and 7 variants in the alliances of Aconito-Piceion and Piceion excelsae are allocated. It is shown that the floristic composition of aspen and birch secondary forests in the age of 60—80 years is almost identical to the natural forests. However, a slight increase the coenotical role of light-requiring species of grasslands and hemiboreal forests in the secondary communities of the class Brachypodio-Betuletea was noticed as well as some reduction of role the shade-tolerant species of nemoral complex and species of boreal forests of the class Vaccinio-Piceetea. Dominant tree layer under the canopy of secondary series is marked by an active growth of natural tree species.

Testing masting mechanisms of boreal forest species at different stand densities

Mast seeding is considered a reproductive trait resulting from several potential evolutionary forces. Although the mechanisms driving reproduction have been thoroughly investigated and discussed, their relative importance and possible coexistence remain an open question. Seed rain abundance and viability of balsam fir Abies balsamea, white spruce Picea glauca, and white birch Betula papyrifera were monitored during 1994–2007 along a chronosequence of developmental stages at different densities represented by four stands in the boreal forest of Quebec, Canada. The aim of the study was to verify whether seeding temporal dynamics and abundance were affected by stand density, and to test the causal relationships between the hypotheses of pollination efficiency and predator satiation. Seed rain abundance varied substantially among years and species, with the highest values being observed in 1994 and 1996, and attaining up to 28.2 × 103 seeds m−2 year−1. However, the annual dynamics of seed production was similar in the three sites with the higher densities. The greater proportions of germinating and dead seeds were observed in white birch, while balsam fir showed an average of 11.2% of damaged seeds, which were infected by larvae. The conifer species had the higher proportions of empty seeds. The causal models tested on balsam fir demonstrated that seed rain abundance influenced the amount of both viable and larval-infected seeds, although no causal relationship existed between these last two variables. The comparable and synchronous seed rain abundances observed among sites demonstrated that stand density can significantly affect the reproduction of trees, probably by limiting the access to resources, but not the annual dynamics of seeding. Pollination efficiency and predator satiation are important forces in the reproductive effort of trees and could be not mutually exclusive because both these mechanisms of natural selection synergistically coexist in driving the seeding dynamics of balsam fir.

Density estimation for small mammals from livetrapping grids: rodents in northern Canada

Management agencies and quantitative ecologists need robust estimates of population density. The best way of converting population estimates of livetrapped small mammals to population density is not clear. We estimated population density on livetrapping grids with 4 estimators applied to 3 species of boreal forest and 3 species of tundra rodents to test for relative differences in density estimators. We used 2 spatial estimators proposed by Efford (2009) and 2 traditional boundary-strip estimators designed for grid livetrapping. We, analyzed mark-recapture data from 104 trapping sessions from the boreal forest at Kluane, Yukon (n = 4,818 individuals), and 56 trapping sessions from tundra areas of Herschel Island and Komakuk Beach in northern Yukon (n = 1,327 individuals). For boreal forest rodents on average both boundary-strip methods produced density estimates larger than Efford's maximum-likelihood (ML) estimator by as much as 50% at all population densities up to 25 animals/ha. For tundra rodents both boundary-strip methods produced density estimates smaller than Efford's ML at low density (<1.5/ha) and larger than Efford's ML density by 36–63% at high density (25/ha). Efford's inverse prediction estimator produced larger density estimates than the ML estimator by 4% for the boreal forest and 32% for the tundra rodents. Relationships were high between all the estimators, such that trends in density could be inferred from all methods. Determining the bias in population density estimators in small mammals will require data from populations spatially closed and completely enumerated. For our small mammals Efford's ML estimator typically provided density estimates smaller than those produced by conventional boundary-strip estimators.

Soil-to-plant transfer of elements is not linear: Results for five elements relevant to radioactive waste in five boreal forest species

Element-specific concentration ratios (CRs) assuming that plant uptake of elements is linear are commonly used in radioecological modelling to describe the soil-to-plant transfer of elements. The goal of this study was to investigate the validity of the linearity assumption in boreal forest plants, for which only limited relevant data are available. The soil-to-plant transfer of three essential (Mo, Ni, Zn) and two non-essential (Pb, U) elements relevant to the safety of radioactive waste disposal was studied. Three understory species (blueberry, narrow buckler fern and May lily) and two tree species (Norway spruce and rowan) were included. Examining CRs as a function of soil concentration showed that CR was not constant but decreased with increasing soil concentrations for all elements and plant species. A non-linear equation fitted fairly well with the empirical data; the R 2-values for this equation were constantly higher than those for the linear fit. The difference between the two fits was most evident at low soil concentrations where the use of constant CRs underestimated transfer from soil to plants. Site-specific factors affected the transfer of Mo and Ni. The results suggested that systematic variation with soil concentrations explains a part of the large variation of empirically determined CRs, and the accuracy of modelling the soil-to-plant transfer might be improved by using non-linear methods. Non-linearity of soil-to-plant transfer has been previously reported for a few different species, elements and environments. The present study systematically tested the linearity assumption for five elements (both essential and non-essential) and in five boreal forest species representing different growth traits and phylogenies. The data supported non-linearity in all cases.

A fast and reliable method for the delineation of tree crown outlines for the computation of crown openness values and other crown parameters

Numerous crown parameters (e.g., leaf area index, diameter, height, volume) can be obtained via the analysis of tree crown photographs. In all cases, parameter values are functions of the position of the crown outline. However, no standardized method to delineate crowns exists. To explore the effect of different outlines on tree crown descriptors, in this case crown openness (CO), and facilitate the adoption of a standard method free of user bias, we developed the program Crown Delineator that automatically delineates any outline around tree crowns following predetermined sensibility settings. We used different outlines to analyze tree CO in contrasting settings: using saplings from four species in young boreal mixedwood forests and medium-sized hybrid poplar trees from a low-density plantation. In both cases, the estimated CO increases when calculated from a looser outline, which had a strong influence on understory available light simulations using a forest simulator. These results demonstrate that the method used to trace crown outlines is an important step in the determination of CO values. We provide a much-needed computer-assisted solution to help standardize this procedure, which can also be used in many other situations in which the delineation of tree crowns is needed (e.g., competition and crown shyness).

Genetic Differentiation of the Western Capercaillie Highlights the Importance of South-Eastern Europe for Understanding the Species Phylogeography

The Western Capercaillie (Tetrao urogallus L.) is a grouse species of open boreal or high altitude forests of Eurasia. It is endangered throughout most mountain range habitat areas in Europe. Two major genetically identifiable lineages of Western Capercaillie have been described to date: the southern lineage at the species' southernmost range of distribution in Europe, and the boreal lineage. We address the question of genetic differentiation of capercaillie populations from the Rhodope and Rila Mountains in Bulgaria, across the Dinaric Mountains to the Slovenian Alps. The two lineages' contact zone and resulting conservation strategies in this so-far understudied area of distribution have not been previously determined. The results of analysis of mitochondrial DNA control region sequences of 319 samples from the studied populations show that Alpine populations were composed exclusively of boreal lineage; Dinaric populations of both, but predominantly (96%) of boreal lineage; and Rhodope-Rila populations predominantly (>90%) of southern lineage individuals. The Bulgarian mountains were identified as the core area of the southern lineage, and the Dinaric Mountains as the western contact zone between both lineages in the Balkans. Bulgarian populations appeared genetically distinct from Alpine and Dinaric populations and exhibited characteristics of a long-term stationary population, suggesting that they should be considered as a glacial relict and probably a distinct subspecies. Although all of the studied populations suffered a decline in the past, the significantly lower level of genetic diversity when compared with the neighbouring Alpine and Bulgarian populations suggests that the isolated Dinaric capercaillie is particularly vulnerable to continuing population decline. The results are discussed in the context of conservation of the species in the Balkans, its principal threats and legal protection status. Potential conservation strategies should consider the existence of the two lineages and their vulnerable Dinaric contact zone and support the specificities of the populations.

Landscape context affects the success of habitat restoration: large‐scale colonization patterns of saproxylic and fire‐associated species in boreal forests

AbstractAim Restoration of habitats may be used as a conservation tool when ecosystems have lost their natural structure, dynamics or functioning over large areas. Controlled and planned use of fire could be an effective way to restore habitats of many threatened species in boreal forests where fire suppression has been effective. We asked whether the large‐scale landscape context affects the occurrence of rare and threatened species in forest habitats that have been burned to restore their fire‐related structures.Location Boreal forests in southern Finland.Methods We designed a large‐scale field experiment that included nine Pinus sylvestris forests (5–10 ha each) in southern Finland. Sites were located in two regions: (1) in eastern region with shorter management history and (2) in western region where intensive forestry has continued longer. We evaluated whether restoration of dead/burned wood is beneficial for rare and conservation‐dependent species and measured the recovery of pyrophilous and red‐listed insects (beetles and flatbugs) in burned forests, using standardized sampling effort. Altogether, 956 individuals of 29 red‐listed and pyrophilous species were sampled.Results Rare species colonized areas quickly, but there was a clear difference in species richness between the regions. The eastern forests harboured higher species richness after restoration. In these sites, the average species richness was 13.7 species per site, whereas in western forests it was 5.0 species per site. Similar pattern was also observed in subgroups: the corresponding numbers for pyrophilous species were 9.7 vs. 3.8, for red‐listed 8.7 vs. 2.3 and for red‐listed pyrophiles 4.7 vs. 1.2.Main conclusions Introducing fire back to boreal forests can aid in the recovery of rare species, but the landscape context considerably affects the success of restoring species. If restored habitats are located in landscapes that have lost their natural properties long ago, the success of restoration seems to be more challenging than in landscapes where habitats have been modified more recently.

Spatial genetic structure in Swiss stone pine suggests a region-specific population history of the species along the Carpathian range

Recently reported evidence supports the existence of ancient populations of cold-tolerant boreal forest species in the upper ranges of the Carpathians. Studies on allele length polymorphism designated several glacial refugia for Norway spruce (Picea abies) in the South Carpathians. Former molecular studies performed on live remnant populations of Swiss stone pine (Pinus cembra) and Scots pine (Pinus sylvestris) suggest that the Carpathians are important in terms of accumulation of genetic diversity

Assessment of assisted migration effects on spring bud flush in white spruce (Picea glauca[Moench] Voss) seedlings

In a changing climate, delaying the time of bud flush may be advantageous to boreal forest species to reduce the risk of spring frost damage. In this study, we examined the potential effect of assisted migration of tree seed on time to bud flush for white spruce. Flushing times of seedlings from 23 white spruce provenances from Ontario were observed under varying temperature conditions simulated in controlled environment chambers. Results indicated that time to bud flush varied considerably among provenances. Although higher temperatures significantly promoted bud flushing for all provenances, provenance-by-temperature interactions were negligible, indicating stable performance of white spruce provenances for this adaptive trait. Spatial patterns of variation among provenances in bud flushing were not consistent with patterns found in range-wide provenance tests. Assisted migration of tree seed across relatively short distances is unlikely to delay bud flushing time in white spruce. Some southern populations may flush earlier at more northerly sites, which would exacerbate spring frost risk. Tree improvement may be an effective approach to utilize amongand withinprovenance variation to enhance this fitness trait for better climatic adaptation.

Early effects of pre- and post-harvest herbicide application and partial cutting in regenerating aspen – jack pine mixtures in northeastern Ontario

Silvicultural options to obtain mixtures of broadleaved and conifer tree species in boreal forests remain limited, especially for mixtures of broadleaves and shade-intolerant conifers. In this study, we tested a series of treatment packages that included variation in harvesting pattern, site preparation, renewal treatments, and timing and patterns of tending to establish trembling aspen (Populus tremuloides Michx.) and jack pine (Pinus banksiana Lamb.) mixedwoods in intimate mixtures or spatial mosaics in northeastern Ontario. Preharvest spray resulted in fewer but better growth of aspen suckers compared with postharvest spray, which both reduced sucker density and decreased their vigour. Partial cutting reduced aspen regeneration in both the harvested and leave corridors but did not affect other broadleaved species, in this case predominantly red maple (Acer rubrum L.). After treatment, the rapid development of understory vegetation (shrubs, herbs, and grasses) and regeneration reduced total light transmittance at 0.5 m but had no effect at 4.0 m after 5 years. Future stands will be jack pine dominated following both pre- and post-harvest sprays, despite higher density, more uniform distribution, and better growth of broadleaved regeneration with preharvest spray, but will be balanced broadleaf and conifer mixtures in the partial cut areas.

Effects of Retention Felling on Epixylic Species in Boreal Spruce Forests in Southern Finland

We studied the effects of patch retention felling and soil scarification by harrowing on the coverage and species richness of epixylic species in boreal Norway spruce (Picea abies) forests in Southern Finland. The epixylics were investigated from both the retention tree groups (RTGs) and the surrounding felling areas before and after fellings and after scarification on consecutive years. The cover percentage of all included species groups was shown to decrease after the felling, especially in the felling areas (vascular plants, −0.4%; mosses, −27.8%; liverworts, −4.0%; and lichens,−2.1%). The decrease was considerable also in the RTGs. The amount of dead moss increased in both the areas indicating microclimatic changes. Species richness also declined rapidly after the first year, especially in the felling areas (vascular plants, −2.2%; mosses, −27.3%; liverworts, −30.3%; and lichens, −22.9%). Scarification also decreases covers and species richness of bryophytes. After the second year, the covers of the species groups generally started to regain, especially in the untreated RTGs. The size of RTG was in positive correlation with the total species number. Another main result indicates that it is possible to maintain much higher initial vegetation abundance and diversity in the RTGs than in the felling areas. Coarse woody debris formed by the frequent tree uprootings may also enhance the long-term survival of epixylics over forest regeneration period. RTGs should be at least 10 times larger than the size used in current Finnish forestry, so that they could function as species refugia.

Archaeal Communities in Boreal Forest Tree Rhizospheres Respond to Changing Soil Temperatures

Temperature has generally great effects on both the activity and composition of microbial communities in different soils. We tested the impact of soil temperature and three different boreal forest tree species on the archaeal populations in the bulk soil, rhizosphere, and mycorrhizosphere. Scots pine, silver birch, and Norway spruce seedlings were grown in forest humus microcosms at three different temperatures, 7-11.5°C (night-day temperature), 12-16°C, and 16-22°C, of which 12-16°C represents the typical mid-summer soil temperature in Finnish forests. RNA and DNA were extracted from indigenous ectomycorrhiza, non-mycorrhizal long roots, and boreal forest humus and tested for the presence of archaea by nested PCR of the archaeal 16S rRNA gene followed by denaturing gradient gel electrophoresis (DGGE) profiling and sequencing. Methanogenic Euryarchaeota belonging to Methanolobus sp. and Methanosaeta sp. were detected on the roots and mycorrhiza. The most commonly detected archaeal 16S rRNA gene sequences belonged to group I.1c Crenarchaeota, which are typically found in boreal and alpine forest soils. Interestingly, also one sequence belonging to group I.1b Crenarchaeota was detected from Scots pine mycorrhiza although sequences of this group are usually found in agricultural and forest soils in temperate areas. Tree- and temperature-related shifts in the archaeal population structure were observed. A clear decrease in crenarchaeotal DGGE band number was seen with increasing temperature, and correspondingly, the number of euryarchaeotal DGGE bands, mostly methanogens, increased. The greatest diversity of archaeal DGGE bands was detected in Scots pine roots and mycorrhizas. No archaea were detected from humus samples from microcosms without tree seedling, indicating that the archaea found in the mycorrhizosphere and root systems were dependent on the plant host. The detection of archaeal 16S rRNA gene sequences from both RNA and DNA extractions show that the archaeal populations were living and that they may have significant contribution to the methane cycle in boreal forest soil, especially when soil temperatures rise.

European aspen and hybrid aspen under changing environment – Leaf traits, growth and litter decomposition

Understanding the responses of species and ecosystems to human-induced global environmental change has become a high research priority. The main aim of this thesis was to investigate how certain environmental factors that relate to global change affect European aspen (Populus tremula), a keystone species in boreal forests, and hybrid aspen (P. tremula × P. tremuloides), cultivated in commercial plantations. The main points under consideration were the acclimatization potential of aspen through changes in leaf morphology, as well as effects on growth, leaf litter chemistry and decomposition. The thesis is based on two experiments, in which young aspen ( 8 years) were growing in environments exposed to multiple environmental stress factors (roadside and urban environments). The field studies included litter decomposition experiments. The results show that young aspen, especially the native European aspen, was sensitive to O3 in terms of visible leaf injuries. Elevated O3 resulted in reduced biomass allocation to roots and accelerated leaf senescence, suggesting negative effects on growth in the long term. Water and N availability modified the frost hardening of young aspen: High N supply, especially when combined with drought, postponed the development of frost hardiness, which in turn may predispose trees to early autumn frosts. This effect was more pronounced in European aspen. The field studies showed that mature aspen acclimatized to roadside and urban environments by producing more xeromorphic leaves. Leaf morphology was also observed to vary in response to interannual climatic variation, which further indicates the ability of aspen for phenotypic plasticity. Intraspecific variation was found in several of the traits measured, although intraspecific differences in response to the abiotic factors examined were generally small throughout the studies. However, some differences between clones were found in sensitivity to O3 and the roadside environment. Aspen leaf litter decomposition was retarded in the roadside environment, but only initially. By contrast, decomposition was found to be faster in the urban than the rural environment throughout the study. The higher quality of urban litter (higher in N, lower in lignin and phenolics), as well as higher temperature, N deposition and humus pH at the urban site were factors likely to promote decay. The phenotypic plasticity combined with intraspecific variation found in the studies imply that aspen has potential for withstanding environmental changes, although some global change factors, such as rising O3 levels, may adversely affect its performance. The results also suggest that the multiple environmental changes taking place in urban areas – which correspond closely with the main drivers of global change – can modify ecosystem functioning by promoting litter decomposition, mediated partly by alterations in leaf litter quality.

Trends in growing space efficiency of four boreal forest species based on yield table data

Yield tables are used to identify trends in growing space efficiency (GSE) and to relate GSE to self-tolerance and intraspecific competition. The method is useful when data specifically collected for this purpose are not available. Plonski’s normal yield tables for jack pine (Pinus banksiana Lamb.), paper birch (Betula papyrifera Marshall), trembling aspen (Populus tremuloides Michx.), and black spruce (Picea mariana (Mill.) B.S.P.) are used. An exponential volume–age function was partitioned into volume–area and area–age functions. The exponents of these two components form the B/D ratio, which is used to determine the mode of the stand at a given time, e.g., if B/D is &lt;3/2, then the stand is in area occupation mode, and if B/D is &gt;3/2, then the stand is in area exploitation mode. The dominant mode is the one most responsive to availability of growth resources, showing greater acceleration when resources are plentiful and more rapid deceleration when resources are scarce. Jack pine and paper birch are identified as area occupiers, whereas trembling aspen and black spruce are area exploiters and are therfore self-tolerant. Asymmetric competition was deemed to be present for paper birch throughout the life of the stand on site class I and for trembling aspen on all sites prior to senescence.

What determines the number of dominant species in forests?

In this work, the difference in number of dominant species in a community on global scale and successional trajectories was analyzed based on the published data. We explained the reasons of these differences using a resource availability hypothesis, proposed in this work, that the distribution of available resource determined the pattern of community dominance. The results showed that on global scale the number of dominant species of community varied across latitudinal forest zone, namely from single-species dominance in boreal and temperate forest to multi-species codominance, even no dominant species in tropical forest. This was consistent with the pattern of resource distribution on global scale. Similarly, in successional trajectories, the number of dominant species gradually radiated from single-species dominance to multi-species codominance, even no dominant species in tropical forest. The changing available resources in trajectories were responsible for this difference. By contrary, a community was often dominated by single species in temperate or boreal forest. This was determined by the low available resource, especially low available water and temperature. In boreal forest, low temperature greatly reduced availability of water and nutrient, which were responsible for the single-species dominance. In addition, the conclusion that high available resources sustained low dominance of community might be deduced, based on the fact that the dominance of community declined with the increasing of species diversity. To sum up, the richer the available resources were, the lower the dominance of community was, and vice versa. The hypothesis that the resource availability controlled the dominance of community could well elucidate the difference of community dominance on global and community scale.