Landscape-variability of the carbon balance across managed boreal forests

Boreal forests are important global carbon (C) sinks and, therefore, considered as a key element in climate change mitigation policies. However, their actual C sink strength is uncertain and under debate, particularly for the actively managed forests in the boreal regions of Fennoscandia. In this study, we use an extensive set of biometric- and chamber-based C flux data collected in 50 forest stands (ranging from 5 to 211 years) over 3 years (2016–2018) with the aim to explore the variations of the annual net ecosystem production (NEP; i.e., the ecosystem C balance) across a 68 km² managed boreal forest landscape in northern Sweden. Our results demonstrate that net primary production rather than heterotrophic respiration regulated the spatio-temporal variations of NEP across the heterogeneous mosaic of the managed boreal forest landscape. We further find divergent successional patterns of NEP in our managed forests relative to naturally regenerating boreal forests, including (i) a fast recovery of the C sink function within the first decade after harvest due to the rapid establishment of a productive understory layer and (ii) a sustained C sink in old stands (131–211 years). We estimate that the rotation period for optimum C sequestration extends to 138 years, which over multiple rotations results in a long-term C sequestration rate of 86.5 t C ha⁻¹ per rotation. Our study highlights the potential of forest management to maximize C sequestration of boreal forest landscapes and associate climate change mitigation effects by developing strategies that optimize tree biomass production rather than heterotrophic soil C emissions.     

Projected climate change effects on Alberta's boreal forests imply future challenges for oil sands reclamation

Climate change will drive significant changes in vegetation cover and also impact efforts to restore ecosystems that have been disturbed by human activities. Bitumen mining in the Alberta oil sands region of western Canada requires reclamation to “equivalent land capability,” implying establishment of vegetation similar to undisturbed boreal ecosystems. However, there is consensus that this region will be exposed to relatively severe climate warming, causing increased occurrence of drought and wildfire, which threaten the persistence of both natural and reclaimed ecosystems. We used a landscape model, LANDIS‐II, to simulate plant responses to climate change and disturbances, forecasting changes to boreal forests within the oil sands region. Under the most severe climate forcing scenarios (representative concentration pathway [RCP] 8.5) the model projected substantial decreases in forest biomass, with the future forest being dominated by drought‐ and fire‐tolerant species characteristic of parkland or prairie ecosystems. In contrast, less extreme climate forcing scenarios (RCPs 2.6 and 4.5) had relatively minor effects on forest composition and biomass with boreal conifers continuing to dominate the landscape. If the climate continues to change along a trajectory similar to those simulated by climate models for the RCP 8.5 forcing scenario, current reclamation goals to reestablish spruce‐dominated boreal forest will likely be difficult to achieve. Results from scenario modeling studies such as ours, and continued monitoring of change in the boreal forest, will help inform reclamation practices, which could include establishment of species better adapted to warmer and drier conditions.     

Different facets of tree sapling diversity influence browsing intensity by deer dependent on spatial scale

Browsing of tree saplings by deer hampers forest regeneration in mixed forests across Europe and North America. It is well known that tree species are differentially affected by deer browsing, but little is known about how different facets of diversity, such as species richness, identity, and composition, affect browsing intensity at different spatial scales. Using forest inventory data from the Hainich National Park, a mixed deciduous forest in central Germany, we applied a hierarchical approach to model the browsing probability of patches (regional scale) as well as the species‐specific proportion of saplings browsed within patches (patch scale). We found that, at the regional scale, the probability that a patch was browsed increased with certain species composition, namely with low abundance of European beech (Fagus sylvatica L.) and high abundance of European ash (Fraxinus excelsior L.), whereas at the patch scale, the proportion of saplings browsed per species was mainly determined by the species’ identity, providing a “preference ranking” of the 11 tree species under study. Interestingly, at the regional scale, species‐rich patches were more likely to be browsed; however, at the patch scale, species‐rich patches showed a lower proportion of saplings per species browsed. Presumably, diverse patches attract deer, but satisfy nutritional needs faster, such that fewer saplings need to be browsed. Some forest stand parameters, such as more open canopies, increased the browsing intensity at either scale. By showing the effects that various facets of diversity, as well as environmental parameters, exerted on browsing intensity at the regional as well as patch scale, our study advances the understanding of mammalian herbivore–plant interactions across scales. Our results also indicate which regeneration patches and species are (least) prone to browsing and show the importance of different facets of diversity for the prediction and management of browsing intensity and regeneration dynamics.     

The effect of biodiversity on tree productivity: from temperate to boreal forests

Aim An important issue regarding biodiversity concerns its influence on ecosystem functioning. Experimental work has led to the proposal of mechanisms such as niche complementarity. However, few attempts have been made to confirm these in natural systems, especially in forests. Furthermore, one of the most interesting unresolved questions is whether the effects of complementarity on ecosystem functioning (EF) decrease in favour of competitive exclusions over an increasing productivity gradient. Using records from permanent forest plots, we asked the following questions. (1) Is tree productivity positively related to diversity? (2) Does the effect of diversity increase in less productive forests? (3) What metric of diversity (e.g. functional or phylogenetic diversity) better relates to tree productivity? Location Temperate, mixed and boreal forests of eastern Canada. Methods Over 12,000 permanent forest plots, from temperate to boreal forests, were used to test our hypotheses in two steps. (1) Stepwise regressions were used to identify the best explanatory variables for tree productivity. (2) The selected climatic and environmental variables, as well as density and biodiversity indices, were included in a structural equation model where links (paths) between covarying variables are made explicit, making structural equation modelling the best tool to explore such complicated causal networks. Results This is the first large‐scale demonstration of a strong, positive and significant effect of biodiversity on tree productivity with control for climatic and environmental conditions. Important differences were noted between the two forest biomes investigated. Main conclusions We show for the first time that complementarity may be less important in temperate forests growing in a more stable and productive environment where competitive exclusion is the most probable outcome of species interactions, whereas in the more stressful environment of boreal forests, beneficial interactions between species may be more important. The present work is also a framework for the analysis of large datasets in biodiversity–ecosystem functioning (B‐EF) research.     

Simulation of the effects of deer browsing on forest dynamics

I introduced forest dynamics to a deterministic herbivore-vegetation model to examine the effects of vertically stratified forest structure on the dynamics of the herbivore–vegetation system, the resilience and stable states of vegetation, and the interactions between deer populations and vegetation. I constructed a model based on data from field studies performed in Hokkaido, northern Japan. Three phases of state were identified for a given deer density: (1) understory vegetation is maintained with a equilibrium proportion of canopy gaps in the absence of deer; (2) if the equilibrium proportion of canopy gaps is greater than that in the equilibrium state in the absence of deer, the understory vegetation can be maintained; and (3) the understory vegetation cannot be maintained. At the boundary between phases 2 and 3, the herbivore population level had discontinuous effects on vegetation. When the deer density was held below the threshold, forest vegetation had resilience to recover to the equilibrium stable state at the given deer density, although the equilibrium canopy gap ratio and vegetation biomass differed with deer density. However, the forest vegetation–herbivore system could not be maintained in a stable state without artificial deer population management if food limitation was the only mechanism to keep the deer population at low levels. The deer population must be kept below the boundary between phases 1 and 2 to maintain the forest regeneration processes. The level cannot be determined by observing the deer population; careful observation of forest regeneration processes is required.     

Effects of post-fire conditions on soil respiration in boreal forests with special reference to Northeast China forests

Forest fires frequently occur in boreal forests, and their effects on forest ecosystems are often significant in terms of carbon flux related to climate changes. Soil respiration is the second largest carbon flux in boreal forests and the change in soil respiration is not negligible. Environmental factors controlling the soil respiration, for example, soil temperature, are altered by such fires. The abnormal increase in soil temperature has an important negative effect on soil microbes by reducing their activities or even by killing them directly with strong heat. On the other hand, although vegetation is directly disturbed by fires, the indirect changes in soil respiration are followed by changes in root activities and soil microbes. However, there is very limited information on soil respiration in the forests of Northeast China. This review, by combining what is known about fire influence on soil respiration in boreal forests from previous studies of post-fire effects on soil conditions, soil microbes, and forest regeneration, presents possible scenarios of the impact of anticipated post-fire changes in forest soil respiration in Northeast China.     

Effects of post-fire conditions on soil respiration in boreal forests with special reference to Northeast China forests

Forest fires frequently occur in boreal forests,and their effects on forest ecosystems are often significant in terms of carbon flux related to climate changes.Soil respiration is the second largest carbon flux in boreal forests and the change in soil respiration is not negligible.Environmental factors controlling the soil respiration,for example,soil temperature,are altered by such fires.The abnormal increase in soil temperature has an important negative effect on soil microbes by reducing their activities or even by killing them directly with strong heat.On the other hand,although vegetation is directly disturbed by fires,the indirect changes in soil respiration are followed by changes in root activities and soil microbes.However,there is very limited information on soil respiration in the forests of Northeast China.This review,by combining what is known about fire influence on soil respiration in boreal forests from previous studies of post-fire effects on soil conditions,soil microbes,and forest regeneration,presents possible scenarios of the impact of anticipated post-fire changes in forest soil respiration in Northeast China.     

A demographic study of deer browsing impacts on Trillium grandiflorum

When white-tailed deer populations reach high densities, they have negative and often dramatic effects on forest herb populations. However, it is not clear how deer affect the demographic processes of plant populations. We examined how the structure and dynamics of Trillium grandiflorum (Michx.) Salisb. populations are affected by deer browsing in the Upper Great Lakes region by sampling populations from nine study sites in a forested landscape in 1998 and 1999. We constructed a stage-based matrix population model for the regional population. Our model indicated that the long-term growth rate of the population to be –3.56% per year ( = 0.965). Mortality rates were highest for seeds (97.5%) followed by seedlings (29.1%), and lower for all remaining stage classes (4.9 to 8.5%). The observed stage distribution significantly differed from the stable stage distribution, and the damping ratio ( = 1.103) indicated the population would not reach its stable stage distribution anytime soon. In the absence of deer browsing, the long-term growth rate would improve to between –3.46% and –1.61% per year. A moderate drought during the study could account for the negative population growth rate, but deer browsing accelerates the rate of decline. Population growth is most sensitive to the proportion of plants remaining in the nonflowering stage, and deer browsing reduces this proportion. Browsing damage was relatively low in this study (5.4% of stems in 1998, 11.5% in 1999) compared to another study of browsing impacts on T. grandiflorum, indicating deer could have far more severe demographic consequences in populations subject to higher levels of browsing.     

Complementarity effects are strengthened by competition intensity and global environmental change in the central boreal forests of Canada

Increases in niche complementarity have been hypothesised to reduce the intensity of interspecific competition within natural forests. In regions currently experiencing potentially enhanced growth under global environmental change, niche complementarity may become even more beneficial. However, few studies have provided direct evidence of this mechanism. Here, we use data from 180 permanent sample plots in Manitoba, Canada, with a full spatial mapping of all stems, to show that complementarity effects on average increased with neighbourhood competition intensity and temporally rising CO₂, warming and water availability. Importantly, complementarity effects increased with both shade tolerance and phylogenetic dissimilarity between the focal tree and its neighbours. Our results provide further evidence that increasing stand functional and phylogenetic diversity can improve individual tree productivity, especially for individuals experiencing intense competition and may offer an avenue to maintain productivity under global environmental change.     

The evolution of nettle resistance to heavy deer browsing

We examined whether heavy browsing by sika deer, Cervus nippon Temminck, changed morphological characteristics of a Japanese nettle, Urtica thunbergiana Sieb. et Zucc., in Nara Park, where a large population of sika deer has been maintained for more than 1,200 years. Wild nettles of Nara Park exhibited smaller leaf area, 11–223 times more stinging hairs per leaf, and 58–630-times higher stinging hair densities than those of other areas where there was no evidence of sika deer browsing. There were no significant differences in stinging hair length between the areas. Nettles from Nara Park that were cultivated from seeds in a greenhouse retained a larger number and higher density of stinging hairs. In the field, nettles of Nara Park were less frequently browsed by sika deer and showed higher survivorship than nettles that were transplanted from an unbrowsed area into Nara Park. These results indicate that: (1) the U. thunbergiana population of Nara Park has an extremely high stinging hair density compared with those of unbrowsed areas; (2) this characteristic has a genetic basis, and (3) stinging hairs serve as a defensive structure against sika deer, contributing to an increase in survivorship. Thus, we conclude that a U. thunbergiana population in Nara Park, with extremely high stinging hair densities, has evolved through natural selection due to heavy browsing by sika deer.     

Spatial stratification of host-seeking Diptera in boreal forests of northern Europe

The stratification of haematophagous Diptera was assessed in two boreal forests in northern Sweden by placing traps baited with carbon dioxide at 1.5 m, 5.0 m and 10.0 m above the ground. More than 40 000 specimens were captured, including 617 biting midges (Ceratopogonidae), 4029 mosquitoes (Culicidae) and 36 092 black flies (Simuliidae). Catches at the various trap heights reflected the general vertical distribution of the preferred hosts, with mammalophilic flies predominating (68.6%) in catches at 1.5 m and ornithophilic flies (42.4%) in catches at 10.0 m; however, most flies that use host birds at ground level were caught in the lowest traps (e.g. 85.1% of Simulium annulus were collected at 1.5 m). Within-species variation in vertical patterns between forests suggests plasticity in responses to environmental factors such as vegetative structure.     

Sensitivity of managed boreal forests in Finland to climate change, with implications for adaptive management

This study investigated the sensitivity of managed boreal forests to climate change, with consequent needs to adapt the management to climate change. Model simulations representing the Finnish territory between 60 and 70 degrees N showed that climate change may substantially change the dynamics of managed boreal forests in northern Europe. This is especially probable at the northern and southern edges of this forest zone. In the north, forest growth may increase, but the special features of northern forests may be diminished. In the south, climate change may create a suboptimal environment for Norway spruce. Dominance of Scots pine may increase on less fertile sites currently occupied by Norway spruce. Birches may compete with Scots pine even in these sites and the dominance of birches may increase. These changes may reduce the total forest growth locally but, over the whole of Finland, total forest growth may increase by 44%, with an increase of 82% in the potential cutting drain. The choice of appropriate species and reduced rotation length may sustain the productivity of forest land under climate change.     

Productivity of forests in the Eurosiberian boreal region and their potential to act as a carbon sink –- a synthesis

Based on review and original data, this synthesis investigates carbon pools and fluxes of Siberian and European forests (600 and 300 million ha, respectively). We examine the productivity of ecosystems, expressed as positive rate when the amount of carbon in the ecosystem increases, while (following micrometeorological convention) downward fluxes from the atmosphere to the vegetation (NEE = Net Ecosystem Exchange) are expressed as negative numbers. Productivity parameters are Net Primary Productivity (NPP=whole plant growth), Net Ecosystem Productivity (NEP = CO₂ assimilation minus ecosystem respiration), and Net Biome Productivity (NBP = NEP minus carbon losses through disturbances bypassing respiration, e.g. by fire and logging). Based on chronosequence studies and national forestry statistics we estimate a low average NPP for boreal forests in Siberia: 123 gC m^–2 y^–1. This contrasts with a similar calculation for Europe which suggests a much higher average NPP of 460 gC m^–2 y^–1 for the forests there. Despite a smaller area, European forests have a higher total NPP than Siberia (1.2–1.6 vs. 0.6–0.9 × 10¹⁵ gC region^–1 y^–1). This arises as a consequence of differences in growing season length, climate and nutrition. For a chronosequence of Pinus sylvestris stands studied in central Siberia during summer, NEE was most negative in a 67-y old stand regenerating after fire (– 192 mmol m^–2 d^–1) which is close to NEE in a cultivated forest of Germany (– 210 mmol m^–2 d^–1). Considerable net ecosystem CO₂-uptake was also measured in Siberia in 200- and 215-y old stands (NEE:174 and – 63 mmol m^–2 d^–1) while NEP of 7- and 13-y old logging areas were close to the ecosystem compensation point. Two Siberian bogs and a bog in European Russia were also significant carbon sinks (– 102 to – 104 mmol m^–2 d^–1). Integrated over a growing season (June to September) we measured a total growing season NEE of – 14 mol m^–2 summer^–1 (– 168 gC m^–2 summer^–1) in a 200-y Siberian pine stand and – 5 mol m^–2 summer^–1 (– 60 gC m^–2 summer^–1) in Siberian and European Russian bogs. By contrast, over the same period, a spruce forest in European Russia was a carbon source to the atmosphere of (NEE: + 7 mol m^–2 summer^–1 = + 84 gC m^–2 summer^–1). Two years after a windthrow in European Russia, with all trees being uplifted and few successional species, lost 16 mol C m^–2 to the atmosphere over a 3-month in summer, compared to the cumulative NEE over a growing season in a German forest of – 15.5 mol m^–2 summer^–1 (– 186 gC m^–2 summer^–1; European flux network annual averaged – 205 gC m^–2 y^–1). Differences in CO₂-exchange rates coincided with differences in the Bowen ratio, with logging areas partitioning most incoming radiation into sensible heat whereas bogs partitioned most into evaporation (latent heat). Effects of these different surface energy exchanges on local climate (convective storms and fires) and comparisons with the Canadian BOREAS experiment are discussed. Following a classification of disturbances and their effects on ecosystem carbon balances, fire and logging are discussed as the main processes causing carbon losses that bypass heterotrophic respiration in Siberia. Following two approaches, NBP was estimated to be only about 13–16 mmol m^–2 y^–1 for Siberia. It may reach 67 mmol m^–2 y^–1 in North America, and about 140–400 mmol m^–2 y^–1 in Scandinavia. We conclude that fire speeds up the carbon cycle, but that it results also in long-term carbon sequestration by charcoal formation. For at least 14 years after logging, regrowth forests remain net sources of CO₂ to the atmosphere. This has important implications regarding the effects of Siberian forest management on atmospheric concentrations. For many years after logging has taken place, regrowth forests remain weaker sinks for atmospheric CO₂ than are nearby old-growth forests.     

Community structure of insect herbivores in a hybrid system: examining the effects of browsing damage and plant genetic variation

Abstract.  1. Using three genetic classes of willows, Salix eriocephala , Salix sericea , and their interspecific F1 hybrid, the influence of browsing damage and the importance of genetic class on insect community structure were evaluated.
2. Three-year-old plants grown from seeds generated from controlled crosses were placed in a common garden after a damage treatment was imposed on them (plants were either left undamaged during the previous winter or they had 50% of the previous year's growth removed). Clipping damage caused large increases in mean shoot length for plants.
3. The abundance of eight species of insect herbivores was determined for every plant to evaluate community structure for three genetic classes across the two damage levels. Based on manova , damage treatment had a modest effect on the relative abundance of herbivores (i.e. their proportional representation). In contrast, dramatic differences were detected among genetic classes for relative abundance; in cases where damage treatment influenced relative abundance of herbivores, the importance of genetic class was at least 20-fold greater than that of damage treatment. No interaction between genetic class and browsing treatment was detected for community structure.
4. The weak response of the herbivore community to clipping damage, contrasted to the large response to genetic class, was very surprising because mean shoot length was greatly altered by damage treatment. These findings, coupled together with previous research, suggest that plant genetic differences can act as the primary basis for herbivore community structure, while the effects of browsing may not be as common.     

Spatial hints of forest ecotone indicating forest succession, a case of larch forests in Baihuashan Reserve, north China

In cold or alpine areas of northern China, birch forests and larch forests are the two primary forest types. These forests are also characteristic of a south branch of boreal forests in Asia. Some ecologists argue that larch forests can replace birch forests, but this still remains a question due to fragmentary or short observations. The ecotone between a larch forest patch and a birch forest patch is the arena in which the two species interplay and compete with each other, and studies of these areas are meaningful to understanding forest succession. In the alpine area of the Baihuashan Reserve, northern China, we sampled a larch-birch forest ecotone with eight plots in four transects and then analyzed population structures of larches and birches. The results show that the edges of the larch forest patch are composed of many larch saplings or young trees, but the edges of the birch forest patch are mainly composed of old birches. Across the ecotone, the larches, on average, are taller than the birches. These facts suggest that larch saplings can permeate into birch forest patches, probably by seed dispersal, germination, success-ful competition and growth, but birch saplings cannot permeate into larch forest patches. Therefore, on the ecotone, larch forest patches can steadily expand by unceasing permeation into birch forest patches, whereas birch forest patches progressively recede due to ultimate death of the old and poor recruitment of the young. Larch forest patches replace birch forest patches in a stepwise manner, causing succession from birch forests to larch forests. This study not only confirms that larch forests can naturally replace birch forests, but also introduces a simple and reliable method, employing spatial hints, to study forest succession. Additionally, the findings are of benefit to cultivation or development of larch forests in cold or alpine areas of the North Temperate Zone, which can be a huge carbon sink.