Effects of soil temperature on biomass production and allocation in seedlings of four boreal tree species

One-year old seedlings of trembling aspen (Populus tremuloides Michx.), black spruce (Picea mariana (Mill.) B.S.P.), white spruce (Picea glauca (Moench) Voss), and jack pine (Pinus banksiana Lamb.) were subject to seven soil temperatures (5, 10, 15, 20, 25, 30 and 35 °C) for 4 months. All aspen seedlings, about 40% of jack pine, 20% of white spruce and black spruce survived the 35 °C treatment. The seedlings were harvested at the end of the fourth month to determine biomass and biomass allocation. It was found that soil temperature, species and interactions between soil temperature and species significantly affected root biomass, foliage biomass, stem biomass and total mass of the seedling. The relationship between biomass and soil temperature was modeled using third-order polynomials. The model showed that the optimum soil temperature for total biomass was 22.4, 19.4, 16.0 and 13.7 °C, respectively, for jack pine, aspen, black spruce and white spruce. The optimum soil temperature was higher for leaf than for root in jack pine, aspen and black spruce, but the trend was the opposite for white spruce. Among the species, aspen was the most sensitive to soil temperature: the maximum total biomass for aspen was about 7 times of the minimum value while the corresponding values were only 2.2, 2.4 and 2.3 times, respectively, for black spruce, jack pine and white spruce. Soil temperature did not significantly affect the shoot/root (S/R) ratio, root mass ratio (RMR), leaf mass ratio (LMR), or stem mass ratio (SMR) (P>0.05) with the exception of black spruce which had much higher S/R ratios at low (5 °C) and high (30 °C) soil temperatures. There were significant differences between species in all the above ratios (P<0.05). Aspen and white spruce had the smallest S/R ratio but highest RMR while black spruce had the highest S/R but lowest RMR. Jack pine had the highest LMR but lowest SMR while aspen had the smallest LMR but highest SMR. Both LMR and SMR were significantly higher for black spruce than for white spruce.     

Corrigendum to: Growth of white spruce underplanted beneath spaced and unspaced aspen stands in northeastern B.C.—10 year results

Establishing white spruce (Picea glauca (Moench) Voss) by planting it under established aspen (Populus tremuloides Michx.), stands has substantial potential as a technique for regenerating boreal mixedwood stands. The presence of an aspen overstory serves to ameliorate frost and winter injury problems and suppresses understory vegetation that may compete with white spruce. In this study we examine the growth of white spruce during the first 10 years after being planted underneath a 39-year-old stand of trembling aspen following thinning and fertilization. Results indicate successful establishment and reasonable growth rates of white spruce planted under thinned and unthinned aspen stands, even with aspen basal area of 51 m² ha⁻¹. Thinning of overstory aspen to 1000 or 2000 stems ha⁻¹ did not increase light reaching seedlings, but did result in improvements in light above the shrub layer and in diameter and height growth of the underplanted seedlings. However, these increases in growth of underplanted spruce may not justify the expense of thinnings. Fertilization of these stands prior to planting had no effect on spruce growth. Growth of spruce underplanted at this site near Fort Nelson was similar to that at two other stands near Dawson Creek, B.C.     

Growth of white spruce underplanted beneath spaced and unspaced aspen stands in northeastern B.C.—10 year results

Establishing white spruce (Picea glauca (Moench) Voss) by planting it under established aspen (Populus tremuloides Michx.), stands has substantial potential as a technique for regenerating boreal mixedwood stands. The presence of an aspen overstory serves to ameliorate frost and winter injury problems and suppresses understory vegetation that may compete with white spruce. In this study we examine the growth of white spruce during the first 10 years after being planted underneath a 39 year-old stand of trembling aspen following thinning and fertilization. Results indicate successful establishment and reasonable growth rates of white spruce planted under thinned and unthinned aspen stands, even with aspen basal area of 51 m² ha⁻¹. Thinning of overstory aspen to 1000 or 2000 stems ha⁻¹ did not increase light reaching seedlings, but did result in improvements in light above the shrub layer and in diameter and height growth of the underplanted seedlings. However, these increases in growth of underplanted spruce may not justify the expense of thinnings. Fertilization of these stands prior to planting had no effect on spruce growth. Growth of spruce underplanted at this site near Fort Nelson was similar to that at two other stands near Dawson Creek, B.C.     

A comparison of riparian plant community response to herbivory by beavers (Castor canadensis) and ungulates in Canada's boreal mixed-wood forest

Herbivores, such as beavers (Castor canadensis) and ungulates, can dramatically alter the species composition and structure of riparian vegetation communities. We examined how four different levels of beaver presence and ungulate density influenced the structure and composition of riparian vegetation communities in the mixed-wood boreal region of east-central Alberta, Canada. Beavers, as central-place foragers, focus on similar species utilized by the six ungulate species in our study area. However, we found that vegetation responded differently relative to the percent cover and heights of emergent, shrub, herbaceous dicots, and tree species depending on the varying combinations of beaver presence and ungulate densities. There was an obvious negative effect of ungulate herbivory on shrub heights and cover; however, we were unable to find any significant effects on the overall composition and structure of vegetation communities that were exclusively related to beaver activities. In areas with no beavers and lower ungulate densities there was a positive growth response in preferred species (e.g., Salix spp., Prunus spp.), which clearly demonstrated the degree of combined influence that these herbivores have on riparian communities. In these areas, Salix spp. was four times shorter than in areas where beavers were not present and ungulate densities were lower. In addition percent cover was lowest for emergent vegetation and herbaceous dicots in areas with higher ungulate densities. Many studies focus on single-species effects on forage resources, yet our research shows that the study of combined effects of multiple herbivores can provide a new insight into these complex systems.     

Partitioning of soil respiration into its autotrophic and heterotrophic components by means of tree-girdling in old boreal spruce forest

Forests accumulate much less carbon than the amount fixed through photosynthesis because of an almost equally large opposing flux of CO₂ from the ecosystem. Most of the return flux to the atmosphere is through soil respiration, which has two major sources, one heterotrophic (organisms decomposing organic matter) and one autotrophic (roots, mycorrhizal fungi and other root-associated microbes dependent on recent photosynthate). We used tree-girdling to stop the flow of photosynthate to the belowground system, hence, blocking autotrophic soil activity in a 120-yr-old boreal Picea abies forest. We found that at the end of the summer, two months after girdling, the treatment had reduced soil respiration by up to 53%. This figure adds to a growing body of evidence indicating (t-test, d.f. = 7, p < 0.05) that autotrophic respiration may contribute more to total soil respiration in boreal (mean 53 ± 2%) as compared to temperate forests (mean 44 ± 3%). Our data also suggests that there is a seasonal hysteresis in the response of total soil respiration to changes in temperature. We propose that this reflects seasonality in the tree below-ground carbon allocation.     

Effects of seed water content and storage temperature on the germination parameters of white spruce, black spruce and lodgepole pine seed

The effect of seed water content (WC) (2–3, 5–6 and 22–25%, on a fresh weight basis), storage temperature (+4, −20, −80 and −196°C) and storage duration (6, 12, 24, 48 and 60 months) on the germination of white spruce (Picea glauca (Moench) Voss), black spruce (Picea mariana (Mill.) B.S.P.) and lodgepole pine (Pinus contorta Dougl. ex Loud. var. latifolia Engelm.) seed was investigated. Germination of white spruce control (untreated) seeds and seeds adjusted to 2–3% and 5–6% WC declined after 48 months of storage at −80 and −196°C, with a further decline at 60 months at −20, −80, −196°C. Germination remained high when control white spruce seeds and seeds with 2–3, 5–6% WC were stored at +4°C, over all storage durations. Generally, black spruce and lodgepole pine exhibited high germination at all storage temperatures at 2–3% and 5–6% WC as well as the control (untreated) seed, for up to 60 months in storage. Germination declined for all three species when seed was conditioned to 22–25% WC. This loss in germination was partially recovered in white spruce seed stored at +4, −20 and −80°C after storage durations of 24, 12 and 48 months, respectively, and in black spruce seeds stored at −20 and −196°C after storage durations of 24 months. Mean germination time (MGT) was relatively constant for all species, under all conditions, except for seed conditioned to 22–25% WC, where MGT increased for white spruce seed stored 48 months at −80 and −196°C, and for black spruce seed stored 24 months at +4 and −80°C and 60 months at −196°C. These results show that the optimal storage temperatures are 4°C for white spruce, and 4, −20, −80, and −196°C for black spruce and lodgepole pine, and 2–6% water content is optimal for all 3 species at these temperatures.     

Transversal tracheid dimension in thinned black spruce and Jack pine stands in the boreal forest

Abstract

Commercial thinning has recently been applied in the boreal forest of Quebec (Canada) to increase the volume growth of the residual trees. We aimed to discover if the growth response influences the transversal tracheid dimension of thinned black spruce (Picea mariana (Mill.) B.S.P.) and Jack pine (Pinus banksiana Lamb.) stands at 0.2 m stem height. All 20 studied stands have shown a significant growth response after the treatment (p<0.0001), especially trees with the lowest radial growth before thinning in comparison with the stand mean. Growing conditions met by suppressed trees before thinning might favour them in the competition for light, water and biomass production after the treatment. Trees with a positive growth response did not significantly modify their measured transversal tracheid dimension except for trees which increased twofold their ring-width after thinning. In this case, lumen area and radial cell diameter extended significantly. However individuals with a growth decrease after thinning registered significantly lower values in their ring width, earlywood and latewood cell numbers (p<0.0001) in both species.     

Long-term response of planted conifers, natural regeneration, and vegetation to harvesting, scalping, and weeding on a boreal mixedwood site

This study reports 14th-year response of a boreal mixedwood stand to different harvest intensities (uncut, 50% partial cut with and without removal of residuals after 3 years, and clearcut), spot site preparation treatments (none and scalped), and chemical weeding frequencies (none, single, and multiple) in northeastern Ontario. The response variables include the survival and growth of planted white spruce (Picea glauca [Moench] Voss) and jack pine (Pinus banksiana Lamb.), height and density of natural regeneration and shrubs, and cover of shrubs and non-woody vegetation. Harvesting and weeding generally improved survival and growth of planted trees, although white spruce survival did not significantly differ among the three weeding frequencies. Harvesting tended to increase heights of hardwood (mostly trembling aspen (Populus tremuloides Michx.)) and conifer (largely balsam fir (Abies balsamea (L.) Mill.).) natural regeneration, cover and density of shrubs, and cover of herbs, lichens, and ferns. Chemical weeding reduced height, density and cover of shrubs, height and density of hardwood regeneration, and fern cover, but increased moss and lichen cover. Spot scalping did not significantly affect planted seedling, natural regeneration, or the vegetation.Maximum survival and growth of planted white spruce and jack pine were achieved using a combination of clearcutting and multiple weeding. However, partial cutting followed by a single weeding produced acceptable survival and reasonable growth of planted trees, particularly for white spruce. Partial canopy removal alone substantially reduced the amount of hardwood regeneration, relative to clearcutting, but did not adequately suppress understory shrubs. Significant improvement in seedling growth following multiple weedings was evident primarily in the complete canopy removal treatments: 50% partial cut with removal of residuals after 3 years and clearcut. While the effects of harvesting and weeding on planted crop trees found in the 5th-year assessments generally persisted at year 14, survival decreased, likely due to light competition from developing hardwood and shrubs.     

Effects of overstory retention and site preparation on growth of planted white spruce seedlings in deciduous and coniferous dominated boreal plains mixedwoods

Survival and growth of planted white spruce was assessed under partial harvest treatments and different site preparation techniques in mixedwood forests of two compositions prior to logging: deciduous dominated (d-dom) – primarily comprised of mature trembling aspen (Populus tremuloides Michx.) and coniferous dominated (c-dom) – primarily comprised of mature white spruce (Picea glauca (Moench) Voss). Levels of overstory retention were 0% (clearcut), 50% and 75% of original basal area, and site preparation techniques were inverted mounding, high speed mixing, scalping and control (no treatment). The survival and growth of white spruce were assessed seven years after planting. The experiment was established as a part of the Ecosystem Management Emulating Natural Disturbance (EMEND) experiment located in northern Alberta, Canada. In the c-dom, the 50% and 75% retention of overstory resulted in reduced growth and survival of white spruce seedlings compared to clearcuts. In contrast, in the d-dom, the seedlings performed best in sites that had 50% of the overstory retained. For the c-dom, the mounding and mixing treatments yielded the best growth of spruce seedlings, while scalping yielded the worst. In the d-dom, spruce growth was highest in sites with the mixing treatment. In the d-dom, growth and survival of the planted spruce was greater than in the c-dom. The natural regeneration of deciduous trees was suppressed by the retention of canopy regardless of original composition.     

Western Carpathian mountain spruce forest after a windthrow: Natural regeneration in cleared and uncleared areas

A catastrophic windstorm called “bora” occurred in The Tatra National Park in Slovakia in 2004 and destroyed an extensive area of mountain spruce forests. Part of the windthrow area was left uncleared for natural development, but most of the area was cleared. We analyzed natural regeneration in cleared and uncleared windthrows, and in reference closed canopy forest in 2007 and 2008 with the aim of assessing the regeneration capability of the windthrown forests and proposing the best method for their restoration.     

Influence of fire intensity on structure and composition of jack pine stands in the boreal forest of Quebec: Live trees,understory vegetation and dead wood dynamics

North American jack pine (Pinus banksiana Lamb.) stands are generally characterized by an even-aged structure resulting from high intensity fires (HIF). However, non-lethal fires of moderate intensity (MIF), which leave behind surviving trees, have also been reported. The objectives of this study were two-fold: (1) assess the concurrent dynamics of live trees, understory vegetation and different types of coarse woody debris (CWD) during succession after HIF; and (2) document how MIF affects stand structure component dynamics compared to HIF. Stands affected by both HIF and MIF were selected. Tree characteristics and age structure, understory biomass, and CWD volume were assessed. Our results suggest that the structural succession of jack pine stands following HIF comprises three stages: young stands (<48 years), premature and mature stands (58–100 years) and old stands (>118 years). Canopy openness and jack pine density significantly decreased with time since HIF, while black spruce density and CWD volume significantly increased. The highest structural diversity was measured in the premature and mature stands. Compared to HIF, MIF increased mean jack pine basal area, decreased average stand density, delayed the replacement of jack pine by black spruce replacement in the canopy, decreased CWD volume, and significantly increased bryophytes mass. MIF increased the diversity of live trees and generally decreased CWD structural diversity. The study confirms the diversity of natural disturbance magnitude and successional processes thereby initiated. Thereafter, it appeared to be relevant for adjustment of disturbance emulating forest-management systems.     

Legacies from natural forest dynamics: Different effects of forest management on wood-inhabiting fungi in pine and spruce forests

The species composition of wood-inhabiting fungi (polypores and corticoids) was investigated on 1138 spruce logs and 992 pine logs in 90 managed and 34 natural or near-natural spruce and pine forests in SE Norway.Altogether, the study included 290 species of wood-inhabiting fungi. Comparisons of logs with similar properties (standardized tree species, decay class, dimension class) in natural and managed forests showed a significant reduction in species number per log in managed spruce forests, but not in managed pine forests. The species number per log in managed spruce forests was 10-55% lower than on logs from natural spruce forests. The reduction was strongest on logs of large dimensions. A comparison of 200-400 spruce logs from natural and managed forests showed a 25% reduction in species richness corresponding to a conservative loss of ca. 40 species on a regional scale.A closer inspection revealed that species confined to medium and very decayed spruce logs were disfavored in managed forests, whereas species on early decay classes and decay generalists were unaffected. Similarly, species preferring large spruce logs were disfavored in managed forests. Forest management had strongest impact on low-frequent species in the spruce forests (more than 50% reduction), whereas common species were modestly affected. Corticoid fungi were more adversely affected than polypore fungi.These results indicate that wood-decaying fungi in pine forests are more adapted to forest disturbances than spruce-associated species. Management measures securing a continuous supply of dead wood are more important in spruce forests than in pine forests.     

Estimating the net carbon balance of boreal open woodland afforestation: A case-study in Québec’s closed-crown boreal forest

Black spruce (Picea mariana (Mill.) B.S.P.) is the dominant tree species in the Canadian province of Québec’s boreal ecosystem, particularly in the black spruce-feathermoss (BSFM) domain (between the 49th and the 52nd parallels). While black spruce is generally well adapted to regenerate after wildfires, regeneration failure can sometimes occur, resulting in the irreversible conversion of closed-crown BSFM to open black spruce-lichen woodlands (OW). With OWs representing approximately 7% (1.6 M ha) of Québec’s BSFM domain, the afforestation of OWs carries significant theoretical potential for carbon (C) sequestration, which has not yet been evaluated. The main objectives of the study were then: (i) to estimate the theoretical C balance of OW afforestation within the closed-crown BSFM domain in Québec’s boreal forest; (ii) to calculate, using the life cycle analysis (LCA) method, all the GHG emissions related to black spruce OW afforestation in the closed-crown BSFM domain of Québec. The CO2FIX v. 3.1 model was used to calculate the biological C balance between the baseline (natural OW of site index 9 at age 50) and afforestation (black spruce plantation of site index 6 at age 25) scenarios, using the best estimates available for all five recommended C compartments (aboveground biomass, belowground biomass, litter, deadwood, and soil). The simulation revealed a biological C balance of 77.0 t C ha⁻¹, 70 years following afforestation, for an average net sequestration rate of 1.1 t C ha⁻¹ year⁻¹. Biological C balance only turns positive after 27 years. When integrating the uncertainties related to both the plantation growth yield and the wildfire disturbance, the average sequestration rate varies between 0.2 and 1.9 t C ha⁻¹ year⁻¹. GHG emissions are 1.3 t CO₂ equiv. ha⁻¹ for all afforestation-related operations, which is less than 0.5% of the biological C balance after 70 years. Thus, GHG emissions do not significantly affect the net C balance of the afforestation project simulated. Several recommendations are made, mostly centered on the factors influencing the growth rate of carbon stocks and the impact of natural disturbances, to minimize the range of uncertainties associated to the sequestration potential and maximize the mitigation benefits of an OW afforestation project.     

Long-term dynamics of large woody debris in a managed boreal forest stream

Little is known about how past forest management in Sweden influenced the quantity and quality of large woody debris (LWD) in streams. The present study provides information of the long-term dynamics of LWD in a reach of a boreal stream intersecting a managed forest. Dendrochronological methods were used to reconstruct mortality years of the pieces of LWD and the general history of fire and cuttings of the surrounding riparian forest. Today, spruce dominates among the living trees, whereas the LWD is dominated by birch in the forest and by pine in the stream. Fire frequency prior to active fire suppression was similar to values reported from boreal forests. Pine trees were more abundant in the riparian forest before selective logging operations and active fire suppression began in the 1800s. Many of the pieces of LWD found in the stream today died more than 200 years ago and derived from a cohort of pines that generated in the early 1600s. Pine LWD in stream channels is highly resistant to decomposition and can reside for more than 300 years. A substantial amount of the LWD found today in managed forest streams in boreal Sweden most likely derives from the time before extensive human influence and is likely to decrease further in the future. Management of riparian forests to ascertain future supply of long-lived LWD in streams should target to increase the proportion of pine trees.     

Distribution of saproxylic beetles in a recently burnt landscape of the northern boreal forest of Québec

This study evaluated the importance of burned habitat characteristics as well as the likely dispersal from specific habitats in the distribution of saproxylic beetles the same year as a fire occurred, in burned black spruce stands (Picea mariana [Mill] B.S.P.) in the northern boreal forest of Québec. The distribution of early post-fire saproxylic species was mainly driven by burned habitat attributes at the plot scale (0.04 ha), especially fire severity, suggesting that the effect of environment attributes can act at a relatively fine scale. Some xylophagous and most predaceous species were more abundant in severely burned stands whereas fire severity had the opposite effect on several common mycophagous species. The amount of newly fire-killed trees that could be used as breeding substrates in the burned stands had only a weak positive influence on these functional groups. The great majority of early saproxylic species were weakly associated with the distance from unburned forests or other recently burned patches that could act as potential “source habitats”. Indeed, these variables were of lesser importance than the attributes of the burned habitat. Woody debris that were already present in plots before the fire, potentially serving as local of source-populations for early colonizers, had virtually no influence on the local abundance of species. Many saproxylic species, including some true pyrophilous, clearly showed higher abundance as distance from unburned stands increased. This unexpected relation may reflect that dispersal of insects toward the burnt landscape very shortly after fire could be driven by the higher amount of volatiles released by severely burned forests, which are more likely as distance from unburned forest increased.     

Threshold effects of variable retention harvesting on understory plant communities in the boreal mixedwood forest

We studied the effects of six levels of dispersed green-tree retention (GTR) harvesting (clearcut (0%), 10%, 20%, 50%, and 75%, and unharvested reference (100%)) on understory plant communities in the 8th growing season post-harvest in the mixedwood boreal forest in northwestern Alberta. For the partial harvest treatments (10%, 20%, 50%, 75%) sample plots were located in the partially harvested (retention) strips as well as in the intervening machine corridors used by the harvesting equipment. The understory plant community was significantly influenced by the gradient of retention level. The cover of understory vegetation, especially graminoids, increased with increasing harvesting intensity for the retention strips and overall considering both plots types. Species richness was unaffected by retention level but did decrease as tree density increased. Lower levels of retention lead to increased abundance of early successional, shade-intolerant species. The results suggest a threshold in understory response to GTR harvesting between the 10% and 20% retention treatments. In terms of understory cover and composition, machine corridors within partially harvested forests resembled clearcuts. The results suggest that retaining more than 10% during GTR harvesting could have significant benefits in terms of maintaining understory plant communities more similar to unharvested reference forest.     

The effect of red wood ant (<Emphasis Type="Italic">Formica rufa</Emphasis> group) mounds on root biomass,density, and nutrient concentrations in boreal managed forests

Red wood ants (Formica rufa group, RWAs) are common insects in boreal forests in Fennoscandia, and they build large, long-lived mounds as their nests. RWA mounds are enriched with carbon and nutrients, but little information is available about how they affect root distribution and the nutrient uptake of trees. In this study, we investigated the biomass, biomass density, nutrient concentrations, and amounts of fine (<2 mm) and coarse (>2 mm) roots in RWA mounds, and compared them with those of surrounding forest soil in mixed coniferous stands of different age classes in Finland. Neither fine nor coarse root biomasses differed significantly between the aboveground parts of the mounds and the organic layer of the soil. Root biomass density was lower in mounds than in the organic layer. However, fine root biomass and biomass density were higher in the belowground parts of mounds than in the surrounding mineral soil. Macroelement (N, Ca, K, P, S, Mg) and Zn and Cu concentrations in roots in the mounds were significantly higher than those in the organic layer. Root biomass and biomass density did not differ between stands of different age classes. The results of this study indicate that RWA mounds increase heterogeneity in root distribution in forest ecosystems, and also increase the availability of nutrients for plants that extend their roots inside RWA mounds.     

Modeling individual tree height growth of Norway spruce and Scots pine from national forest inventory data in Norway

We developed individual tree height growth models for Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.) in Norway based on national forest inventory data. Potential height growth is based on existing dominant height growth models and reduced due to competition by functions developed in this study. Three spatially explicit and two spatially non-explicit competition indices were tested. Distance effects and diameter ratio effects were estimated from the data simultaneously with parameters of the potential modifier functions. Large height measurement errors in the national forest inventory data caused large residual variation of the models. However, the effects of competition on height growth were significant and plausible. The potential modifier functions show that height growth of dominant trees is largely unaffected by competition. Only at higher levels of competition, height growth is reduced as a consequence of competition. However, Scots pine also reduced height growth at very low levels of competition. Distance effects in the spatially explicit competition indices indicated that the closest neighbors are most important for height growth. However, for Scots pine also competitors at larger distance affected height growth. The five competition indices tested in this study explained similar proportions of the variation in relative height growth. Given that unbiased predictions can only be expected for the same plot size, we recommend a spatially explicit index, which describes the distance function with a negative exponential, for use in growth simulators.     

Planning timber harvest of residual forest stands without compromising bird and small mammal communities in boreal landscapes

Numerous efforts have been invested in designing and configuring residual forest stands in Canadian boreal forest to preserve their overall biodiversity. Now that several landscapes have been partially logged, the next issue in forest management involves the planning of residual forest stand harvesting without compromising wildlife populations. Residual stands can be cut when adjacent regeneration reaches 3 m in height according to current regulations in several Canadian provinces (e.g., Québec, Ontario, Alberta, and British-Columbia). However, little is known on whether such regenerating habitat (RE-3m) can maintain wildlife communities similar to those found in unharvested mature forest (CO). We estimated the relative abundance of small mammals and forest birds in RE-3m and CO habitats and characterized landscape and stand structures. These variables were then compared between the two contrasting successional stages and were used to build habitat use models (HUMs) for 21 species. CO and RE-3m differed with regard to several landscape characteristics and stand structure variables as a result of logging. Snowshoe Hare, Northern Flicker, Alder Flycatcher, Ruby-crowned Kinglet, White-throated Sparrow and Magnolia Warbler were more abundant in RE-3m than CO, while Red-backed Vole, Brown Creeper and Golden-crowned Kinglet exhibited lower abundances in RE-3m. No significant differences in abundance were observed for the 12 other species. Species HUMs were highly significant and explained between 64.3 and 99.1% of the total variability in abundance. Following variance partitioning, stand structure variables accounted for most of the explained variability (54.2%) while landscape characteristics accounted for only 28.7%. No difference in species richness was observed but community evenness was greater in CO than RE-3m. Our results suggest that current regulations may threaten the maintenance of 3 out of 21 censused species for which abundances were significantly lower in regenerating 3 m tall stands. As stand structure explained a large amount of variability in abundance, it should be considered during timber harvest planning in both mature and regenerating stands. Until we know more on whether the current regulations are suitable for maintaining overall biodiversity, our results suggest that some mature forest stands should be maintained within managed landscapes for a complete logging rotation period.     

Fertilization—Effects on deciduous tree growth and browsing by moose

Knowledge of plant–herbivore interactions is fundamental for understanding foraging patterns of herbivores and their effects on the ecosystem as well as the human use of natural resources. The nutrient resources available for plants may determine not only growth rate, but also the amounts of secondary compounds in the plant tissues. As a consequence, browsing pressure on plant communities by large herbivores can be affected by fertilization and nitrogen deposition. We conducted a field fertilization experiment to test the hypothesis that fertilization leads to increased sapling growth and increased browsing by a large herbivore, the moose (Alces alces). The study was conducted on fertilized and unfertilized plots (50 m × 50 m) in northern Sweden. The plots were located in Scots pine (Pinus sylvestris) stands of young succession stages with an intermixture of deciduous species. Fertilization with 600 kg calcium–ammonium–nitrate per ha (=200 kg N ha⁻¹), was conducted in spring on one of the plots in each of 25 paired plots. The other plot in each pair was kept as control. The effect of fertilization on plants and browsing pressure was evaluated on five deciduous tree species in June and August the year after fertilization. Both browsing from preceding winter (bites) and current summer (leaf strips and bites) was recorded. Shoot size and other shoot variables increased with fertilization. Fertilization increased winter browsing for most species, but few differences were statistically significant. The late summer recording (August) showed pronounced differences in browsing pressure (leaf strips and bites) between control and treatment. In June, only a trend of increased browsing (leaf strips) was found for all species on fertilized plots. Generally, the pressure of leaf stripping was low in June, probably due to the short time the annual shoots had been exposed to moose browsing. Number of summer bites (recorded in August) was significantly higher on fertilized plots for the Betula spp., and the trend was similar for the other tree species. Within species, there was no difference in strip length between the treatments. The result indicates that fertilization can be used as a habitat improvement for ungulates within wildlife management. But, fertilization is also likely to lead to an increased risk of browsing damage, which is undesirable if forests are being fertilized for increased forest production.