Characterization of radiation regimes in nonrandom forest canopies: theory,measurements, and a simplified modeling approach

We used field measurements and Monte Carlo simulations of canopy gap-size distribution and gap fraction to examine how beam radiation interacts with clumped boreal forest canopies of aspen (Populus tremuloides Michx.), black spruce (Picea mariana (Mill.) B.S.P.) and jack pine (Pinus banksiana Lamb.). We demonstrate that the Beer-Lambert law can be modified to accommodate transmission of radiation through a clumped forest canopy as a function of path length or sun zenith angle. Multiband Vegetation Imager (MVI) measurements and Monte Carlo simulations showed that values of the zenith element clumping index (Omega(e)(0)) are typically between 0.4 and 0.5 in jack pine and black spruce and 0.65 in aspen. Estimates of LAI obtained from MVI measurements of the canopy gap fraction and adjusted for canopy clumping and branch architecture yielded LAI values of 3.0 in jack pine, 3.3 in aspen, and about 6.0 in black spruce. These LAI estimates were within 10-25% of direct measurements made at the same sites. Data obtained with the MVI, along with numerical simulations, demonstrated that assumptions of random foliage distributions in boreal forests are invalid and could yield erroneous values of LAI measured by indirect techniques and false characterizations of atmosphere-biosphere interactions. Monte Carlo simulations were used to develop a general equation for beam radiation penetration as a function of zenith angle in clumped canopies. The essential measurements included stem spacing, crown diameter, crown depth, and within-crown gap fraction.     

Will changes in root-zone temperature in boreal spring affect recovery of photosynthesis in Picea mariana and Populus tremuloides in a future climate?

Future climate will alter the soil cover of mosses and snow depths in the boreal forests of eastern Canada. In field manipulation experiments, we assessed the effects of varying moss and snow depths on the physiology of black spruce (Picea -mariana (Mill.) B.S.P.) and trembling aspen (Populus tremuloides Michx.) in the boreal black spruce forest of western Québec. For 1 year, naturally regenerated 10-year-old spruce and aspen were grown with one of the following treatments: additional N fertilization, addition of sphagnum moss cover, removal of mosses, delayed soil thawing through snow and hay addition, or accelerated soil thawing through springtime snow removal. Treatments that involved the addition of insulating moss or snow in the spring caused lower soil temperature, while removing moss and snow in the spring caused elevated soil temperature and thus had a warming effect. Soil warming treatments were associated with greater temperature variability. Additional soil cover, whether moss or snow, increased the rate of photosynthetic recovery in the spring. Moss and snow removal, on the other hand, had the opposite effect and lowered photosynthetic activity, especially in spruce. Maximal electron transport rate (ETR(max)) was, for spruce, 39.5% lower after moss removal than with moss addition, and 16.3% lower with accelerated thawing than with delayed thawing. Impaired photosynthetic recovery in the absence of insulating moss or snow covers was associated with lower foliar N concentrations. Both species were affected in that way, but trembling aspen generally reacted less strongly to all treatments. Our results indicate that a clear negative response of black spruce to changes in root-zone temperature should be anticipated in a future climate. Reduced moss cover and snow depth could adversely affect the photosynthetic capacities of black spruce, while having only minor effects on trembling aspen.     

Microclimatic and spruce growth gradients adjacent to young aspen stands

Effective utilization of a patch or cluster approach to growing aspen-white spruce mixtures in the boreal forest requires an understanding of how the faster growing aspen patches influence microclimate and spruce growth in adjacent spruce patches. In this paper we examine, how young (11–13 year old) aspen (Populus tremuloides Michx) patches influence microclimatic conditions in adjacent openings and how these are reflected in the growth of white spruce (Picea glauca [Moench] Voss) on three boreal mixedwood sites in west-central Alberta, Canada. Light levels increased continuously across the aspen stand boundary and reached maximum levels within the opening, while soil moisture was highest near the edge of the opening and then decreased with distance into the aspen stand or into the adjacent opening north of the aspen stand. Light levels were reduced over a greater distance when moving north from aspen stand edges compared to openings located south of the young aspen. These young aspen stands provided growing season frost protection within one tree length from the edge. The growth of white spruce was positively related to initial crown size and either light levels or distance from the edge. Stem volume growth was best predicted by initial tree size and the light levels at the midcrown of seedlings.     

Persistent effects of fire severity on early successional forests in interior Alaska

There has been a recent increase in the frequency and extent of wildfires in interior Alaska, and this trend is predicted to continue under a warming climate. Although less well documented, corresponding increases in fire severity are expected. Previous research from boreal forests in Alaska and western Canada indicate that severe fire promotes the recruitment of deciduous tree species and decreases the relative abundance of black spruce (Picea mariana) immediately after fire. Here we extend these observations by (1) examining changes in patterns of aspen and spruce density and biomass that occurred during the first two decades of post-fire succession, and (2) comparing patterns of tree composition in relation to variations in post-fire organic layer depth in four burned black spruce forests in interior Alaska after 10-20 years of succession. We found that initial effects of fire severity on recruitment and establishment of aspen and black spruce were maintained by subsequent effects of organic layer depth and initial plant biomass on plant growth during post-fire succession. The proportional contribution of aspen (Populus tremuloides) to total stand biomass remained above 90% during the first and second decades of succession in severely burned sites, while in lightly burned sites the proportional contribution of aspen was reduced due to a 40-fold increase in spruce biomass in these sites. Relationships between organic layer depth and stem density and biomass were consistently negative for aspen, and positive or neutral for black spruce in all four burns. Our results suggest that initial effects of post-fire organic layer depths on deciduous recruitment are likely to translate into a prolonged phase of deciduous dominance during post-fire succession in severely burned stands. This shift in vegetation distribution has important implications for climate-albedo feedbacks, future fire regime, wildlife habitat quality and natural resources for indigenous subsistence activities in interior Alaska.     

Modeling topographic effects on net ecosystem productivity of boreal black spruce forests

Current regional estimates of net primary productivity (NPP) of boreal black spruce overlook the large variation in NPP caused by small-scale topographic effects on soil water, temperature and nutrient availability. Topographic effects on black spruce NPP could likely be modeled by simulating the lateral and vertical movement of water, and its effects on soil nutrient transformation and uptake, through three-dimensional watersheds defined by aspects and slopes of their topographic positions. To examine this likelihood, the ecosystem model 'ecosys' was run for 120 years on a transect that included upper- and lower-slope positions and a basin in which a basal water table was set 0.5 m below the soil surface. For the run, we used soil properties and weather conditions recorded at the 115-year-old BOREAS Southern Old Black Spruce site. Short-term model performance was tested by comparing diurnal and annual carbon (C) transfers simulated under 1994 weather conditions during the 115th year of the model run with those measured at this site during 1994 by eddy covariance, surface chambers and allometry. After 115 years, annual spruce NPP simulated at the upper-slope positions was twice that at the basin (350 versus 170 g C m-2), whereas accumulated wood C was almost three times as large (6.8 versus 2.4 kg C m-2). In the model, increases in NPP and wood growth in upper-slope positions were caused by lower soil water contents, higher soil temperatures, and more rapid O2 uptake that accelerated heterotrophic respiration and hence nutrient mineralization and uptake. Modeled differences in wood growth with topographic position were quantitatively consistent with measurements of boreal black spruce at several research sites differing in water table depth. Modeled differences also agreed with differences in wood growth rates derived from allometric measurements at boreal black spruce sites differing in productivity indices as a result of differences in subsurface hydrology. The magnitude of these differences clearly indicates the importance of accounting for subsurface hydrology in regional estimates of boreal forest productivity.     

Leaf area dynamics of a boreal black spruce fire chronosequence

Specific leaf area (SLA) and leaf area index (LAI) were estimated using site-specific allometric equations for a boreal black spruce (Picea mariana (Mill.) BSP) fire chronosequence in northern Manitoba, Canada. Stands ranged from 3 to 131 years in age and had soils that were categorized as well or poorly drained. The goals of the study were to: (i) measure SLA for the dominant tree and understory species of boreal black spruce-dominated stands, and examine the effect of various biophysical conditions on SLA; and (ii) examine leaf area dynamics of both understory and overstory for well- and poorly drained stands in the chronosequence. Overall, average SLA values for black spruce (n = 215), jack pine (Pinus banksiana Lamb., n = 72) and trembling aspen (Populus tremuloides Michx., n = 27) were 5.82 +/- 1.91, 5.76 +/- 1.91 and 17.42 +/- 2.21 m2 x kg-1, respectively. Foliage age, stand age, vertical position in the canopy and soil drainage had significant effects on SLA. Black spruce dominated overstory LAI in the older stands. Well-drained stands had significantly higher overstory LAI (P < 0.001), but lower understory LAI (P = 0.022), than poorly drained stands. Overstory LAI was negligible in the recent (3-12 years old) burn sites and highest in the 70-year-old burn site (6.8 and 3.0 in the well- and poorly drained stands, respectively), declining significantly (by 30-50%) from this peak in the oldest stands. Understory leaf area represented a significant portion (> 40%) of total leaf area in all stands except the oldest.     

Photosynthesis and light-use efficiency by plants in a Canadian boreal forest ecosystem

Measurements of the photosynthetic response to midsummer irradiance were made for 11 species representing the dominant trees, understory shrubs, herbaceous plants and moss species in an old black spruce (Picea mariana (Mill.) B.S.P.) boreal forest ecosystem. Maximum rates of photosynthesis per unit foliage area at saturating irradiance, A(max), were highest for aspen (Populus tremuloides Michx.), reaching 16 micromol m(-2) s(-1). For tamarack (Larix laricina (Du Roi) K. Kock) and P. mariana, Amax was only 2.6 and 1.8 micromol m(-2) s(-1), respectively. Values of A(max) for understory shrubs and herbaceous plants were clustered between 9 and 11 micromol m(-2) s(-1), whereas A(max) of feather moss (Pleurozium schreberi (Brid.) Mitt.) reached only 1.9 micromol m(-2) s(-1). No corrections were made for differences in shoot structure, but values of photosynthetic light-use efficiency were similar for most species (70-80 mmol CO2 mol(-1)); however, they were much lower for L. laricina and P. mariana (15 mmol CO2 mol(-1)) and much higher for P. schreberi (102 m;mol CO2 mol(-1)). There was a linear relationship between Amax and foliage nitrogen concentration on an area basis for the broad-leaved species in the canopy and understory, but the data for P. mariana, L. laricina and P. schreberi fell well below this line. We conclude that it is not possible to scale photosynthesis from leaves to the canopy in this ecosystem based on a single relationship between photosynthetic rate and foliage nitrogen concentration.     

Environmental controls over isoprene emission in deciduous oak canopies

In summer 1992, isoprene emission was measured on intact leaves and branches of Quercus alba (L.) at two heights in a forest canopy. Isoprene emission capacity (measured at 30 degrees C and a photosynthetic photon flux density of 1000 micro mol m(-2) s(-1)) was significantly higher in sun leaves than in shade leaves when expressed on a leaf area basis (51 versus 31 nmol m(-2) s(-1); P < 0.01). Because leaf mass per unit area (LMA, g m(-2)) was higher in sun leaves than in shade leaves, emissions of sun and shade leaves expressed on a dry mass basis did not differ significantly (99 versus 89 micro g C g(DW) (-1) h(-1); P = 0.05). Similar measurements in 1995 were consistent with the 1992 data, but data from leaves in more shaded locations demonstrated that isoprene emission capacity decreased with decreasing growth irradiance, irrespective of units of expression. Isoprene emission capacity in leaves of Q. coccinea Muenchh. and Q. velutina Lam. also declined steeply with canopy depth. Emission capacity, on a dry mass basis, showed no obvious pattern with canopy position in Q. prinus L. There was no difference in the temperature response of sun versus shade leaves of Q. alba, but shade leaves exhibited a greater quantum efficiency and saturated at lower irradiance than sun leaves. Rates of isoprene emission measured on branches of Q. alba were approximately 60% of those measured on individual leaves, as a result of self-shading within branch enclosures. It is recommended that within-canopy variation in isoprene emission capacity be incorporated into regional emission models.     

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.     

Temporal variability in basal isoprene emission factor

Seasonal variability in basal isoprene emission factor (&mgr;g C g(-1) h(-1) or nmol m(-2) s(-1), leaf temperature at 30 degrees C and photosynthetically active radiation (PAR) at 1000 &mgr;mol m(-2) s(-1)) was studied during the 1998 growing season at Duke Forest in the North Carolina Piedmont. Emissions from eight upper-canopy white oak (Quercus alba L.) leaves were measured periodically from the onset of isoprene emission on Day of Year (DOY) 119 (April 29) to leaf senescence in late October (DOY 299). Emissions from four leaves were measured under basal conditions with a controlled-environment cuvette system equipped with 10-ml gas-tight syringes and a reduction gas detector. Emissions from the other four leaves were measured under ambient conditions with the same system. Emission rates from the four leaves measured under ambient conditions were adjusted to basal conditions based on the PAR and leaf temperature algorithms of Guenther et al. (1993). The seasonal onset of isoprene emission was in agreement with previous studies where cumulative degree days from the date of the last spring frost were used to estimate bud break, leaf expansion, and increase in basal emission factor (EF). Between DOY 141 (May 21) and 240 (August 28), mean meteorological conditions 6 to 18 h prior to the EF measurements (ambient PAR and temperature) explained up to 78% of the variability in mean basal EF between measurement periods. Summertime mean isoprene emission potential was reached on DOY 141 (May 21) and was maintained until DOY 240 (August 28), when isoprene emission began to decline monotonically as leaf senescence approached. The mean value for leaves measured under ambient conditions and adjusted to basal conditions for DOY 141-240 was 75.6 &mgr;g C g(-1) h(-1) (74.2-79.1), whereas the mean value for leaves measured under basal conditions was 72.9 &mgr;g C g(-1) h(-1) (64.7-88.9). Between DOY 141 and 240, daily mean isoprene EFs varied from 54 to 96 &mgr;g C g(-1) h(-1) (27 to 49 nmol m(-2) s(-1)). In agreement with previous work at this and other sites, basal isoprene emission rates of fully exposed leaves at the crown apex of this tree were about 20% higher than those of the selected leaves. The length of the period prior to measurement of isoprene emission, during which meteorology was correlated with basal EF, appeared to be related to the timing and periodicity of meteorological change, and probably explains quantitative differences in the length of this period among studies. The empirical equation that we derived for this effect explained variability in midday EFs at the study site, but its general applicability remains to be tested. Strong diurnal changes in EF (as high as a factor of 2) are implied in this study, and should be examined further.     

Influences of gap position, vegetation management and herbivore control on survival and growth of white spruce (Picea glauca (Moench) Voss) seedlings

The boreal mixedwood forest type of the Canadian interior boreal is largely comprised of two dominant tree species: white spruce and trembling aspen (Picea glauca and Populus tremuloides). This forest type is expansive, providing important ecosystem services and economic production, yet such mixtures are difficult to establish after harvests. While aspen resprouts and grows vigorously following disturbance, spruce growth is relatively slow and is often limited by intense competition from associated vegetation, including aspen. To improve management, it is important to understand how environmental and vegetative conditions vary in relation to the competitive-facilitative relationship of spruce-aspen mixtures. In this study white spruce was planted across large canopy openings to determine whether survival and height growth is influenced by position within gap and by differing levels of competing vegetation control of aspen and understory plants. In addition, we addressed the issue of herbivory, which can pose a significant threat to planted spruce seedlings. Within each of four sites, linear gaps were created and five gap positions were recognized spanning the southern and northern forest understories, and southern, center and northern positions within each opening. Three different levels of vegetation management were implemented: a brush saw treatment in which all vegetation was cut to ground level, a mixing treatment in which all vegetation and rootstock was ground up, and a control with no vegetation management. The three herbivory treatments excluded large ungulates, small herbivores (rabbits, hares) or had no herbivore exclusion. Growth and survival of white spruce seedlings were measured for four years (1997-2000). Understory survival was significantly lower than within the gap, with the sheltered southern edge position providing the best initial environmental conditions and or ameliorative cover for spruce establishment. However, after four years the shelter effect starts to be inhibitive relative to center and northern gap positions, suggesting the removal of the canopy is necessary before spruce productivity declines. The optimal vegetation management treatment also changed over the study period. The most intensive treatment (mixing) initially showed a negative influence on survival and growth, but by year four, survival converged to approximately 75% for all treatments, and the mixing treatment produced the best height growth. The growth advantage became most evident in the center gap positions, which initially lagged the brushsaw and control treatments. Lastly, some growth losses from herbivory must be expected in boreal mixedwoods, although not enough to merit control. Results have implications for the timing and intensity of silvicultural treatments for harvesting and planting.     

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.     

Online analysis of volatile organic compound emissions from Sitka spruce (Picea sitchensis)

Volatile organic compound (VOC) emissions from Sitka spruce (Picea sitchensis Bong.) growing in a range of controlled light and temperature regimes were monitored online with a proton transfer reaction-mass spectrometer (PTR-MS) operating at a temporal resolution of approximately 1 min. Isoprene emissions accounted for an average of more than 70% of measured VOCs and up to 3.5% of assimilated carbon. Emission rates (E) for isoprene correlated closely with photosynthetic photon flux (PPF) and temperature, showing saturation at a PPF of between 300 and 400 micromol m(-2) s(-1) and a maximum between 35 and 38 degrees C. Under standard conditions of 30 degrees C and 1000 micromol m(-2) s(-1) PPF, the mean isoprene E was 13 microg gdm(-1) h(-1), considerably higher than previously observed in this species. Mean E for acetaldehyde, methanol and monoterpenes at 30 degrees C were 0.37, 0.78 and 2.97 microg gdm(-1) h(-1), respectively. In response to a sudden light to dark transition, isoprene E decreased exponentially by > 98% over about 3 h; however, during the first 7 min, this otherwise steady decay was temporarily but immediately depressed to approximately 40% of the pre-darkness rate, before rallying during the following 7 min to rejoin the general downward trajectory of the exponential decay. The sudden sharp fall in isoprene E was mirrored by a burst in acetaldehyde E. The acetaldehyde E maximum coincided with the isoprene E minimum (7 min post-illumination), and ceased when isoprene emissions resumed their exponential decay. The causes of, and linkages between, these phenomena were investigated.     

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.     

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.     

Carbon and nitrogen dynamics along the log bark decomposition continuum in a mesic old-growth boreal forest

Narrowing the uncertainties in carbon (C) and nitrogen (N) dynamics during decomposition of coarse woody debris (CWD) can significantly improve our understanding of forest ecosystem functioning. We examined C, N and pH dynamics in the least studied CWD component—tree bark in a 66-year-long decomposition chronosequence. The relative C concentration decreased by ca. 32% in pine bark, increased by ca. 18% in birch bark and remained stable in spruce and aspen bark. Nitrogen increased in bark of all tree species. In conifer bark, it increased along with epixylic succession. Over 45 years, the relative C/N ratio in bark decreased by 63 and 45% for coniferous and deciduous species, respectively. Bark pH did not change. Due to bark fragmentation, the total C and N amounts in bark of individual logs of aspen, birch, pine and spruce decreased at average rates of 0.03, 0.02, 0.26 and 0.05 year^?1, and 0.02, 0.02, 0.03 and 0.03 year^?1, respectively. At the forest stand level, the total amounts of C and N in log bark were 853 and 21 kg ha^?1 or 11.2 and 45.5% of the C and N amounts stored in downed logs and ca. 2.3–3.8 and 2.2–2.4%, respectively, of total C and N amounts stored in forest litter. In boreal forests, decomposing log bark may act as a long-term source of N for wood-inhabiting communities.     

Spatial point-pattern analysis for detecting density-dependent competition in a boreal chronosequence of Alberta

In the boreal forest of Alberta, fire and wind often open gaps in the canopy where late-successional species can establish and over time cause a shift in the species distribution from deciduous (e.g., trembling aspen) dominated to mixedwood, to shade-tolerant conifer (e.g., white spruce) dominated stands. This study attempted to understand the change of density-dependent competition in a boreal chronosequence and the role of tree competition in affecting stand structure and mortality. Four 1-ha stem-mapped plots were established to represent a chronosequence comprised of aspen dominated, mixedwood, and spruce dominated stands in Alberta. Second order spatial point-pattern analysis using Ripley's K(t) function showed that intraspecific competition is a prevailing force causing conspecific tree mortality and thus shaping the stand structure. The results of bivariate K(t) function analysis did not reveal sufficient evidence of interspecific competition. This suggested that competitive interaction among heterospecific trees was not strong enough to cause significant tree mortality, but the analysis of marked correlation function revealed that interspecific competition could have a negative impact on tree growth. This study highlights the importance of density-dependent competition in understanding stand dynamics of boreal forests over succession.     

Effects of ungulate browsing on recruitment of aspen and rowan: a demographic approach

Aspen (Populus tremula L.) and rowan (Sorbus aucuparia L.) are important for biodiversity conservation and aesthetic reasons. Intensified levels of ungulate browsing the last decades have raised concerns over the recruitment of aspen and rowan in boreal forests. Demographic data are vital to understand and quantify how the structure of tree populations are affected over time by browsing and other factors but such data are rare. In a five-year study, we quantified vital demographic rates for co-occurring aspen and rowan using exclosures in a productive forest area in eastern-central Sweden. Mean annual browsing rate on small (<1 m) aspen and rowan stems was 20 vs. 38%, whereas the order in browsing intensity was reversed for stems of medium (1–2 m) size (28 vs. 14%). Browsing had a clear negative effect on transition rate from small to medium size class in both species, but had a much smaller impact on transition rate from medium to large (2–3 m) size class. Residence time for small stems and regeneration rate was much higher in aspen than in rowan, but browsing did not affect these dynamics. Transition to ungulate safe height (>3 m) was reduced with a factor of two in rowan and four in aspen by browsing.     

Regeneration dynamics of planted seedling-origin aspen (<Emphasis Type="Italic">Populus tremuloides</Emphasis> Michx.)

Aspen (Populus tremuloides Michx.) is a foundational tree species of the North American boreal forest. After disturbance, clonal aspen stands quickly achieve canopy closure by sending up numerous clonal shoots (root suckers) from their lateral root system. Controlled aboveground disturbance will commonly induce prolific root suckering and thereby increase stem density in clonal aspen stands, but it is unclear if increases in stem density will be observed in planted seedling-origin aspen stands. The objectives of this study were to determine (1) overall root suckering response of planted aspen to aboveground disturbance; (2) if different cut heights of the stem or infliction of root damage impact the number of root suckers produced. We found that planted aspen regenerated readily after disturbance, averaging five root suckers per cut tree. However, individual response was highly variable, ranging from zero to 29 root suckers per root system. Of the cut trees, 75% produced at least one root sucker and 60% produced at least one stump sprout. Cutting trees close to the soil surface produced more root suckers than leaving a 25 cm stump. While root system size (mass and length) was well correlated with aboveground measures of planted aspen, root suckering was not related to root system size. As a result of increased forest reclamation efforts in the boreal forest region the planting of aspen has become a more common practice, necessitating a better understanding of the regeneration dynamics and root suckering potential of these planted and seedling-origin aspen forests.     

Fine root morphological adaptations in Scots pine, Norway spruce and silver birch along a latitudinal gradient in boreal forests

Variability in short root morphology of the three main tree species of Europe's boreal forest (Norway spruce (Picea abies L. Karst.), Scots pine (Pinus sylvestris L.) and silver birch (Betula pendula Roth)) was investigated in four stands along a latitudinal gradient from northern Finland to southern Estonia. Silver birch and Scots pine were present in three stands and Norway spruce was present in all stands. For three fertile Norway spruce stands, fine root biomass and number of root tips per stand area or unit basal area were assessed from north to south. Principal component analysis indicated that short root morphology was significantly affected by tree species and site, which together explained 34.7% of the total variability. The range of variation in mean specific root area (SRA) was 51-74, 60-70 and 84-124 m(2) kg(-1) for Norway spruce, Scots pine and silver birch, respectively, and the corresponding ranges for specific root length were 37-47, 40-48 and 87-97 m g(-1). The range of variation in root tissue density of Norway spruce, Scots pine and silver birch was 113-182, 127-158 and 81-156 kg m(-3), respectively. Sensitivity of short root morphology to site conditions decreased in the order: Norway spruce > silver birch > Scots pine. Short root SRA increased with site fertility in all species. In Norway spruce, fine root biomass and number of root tips per m(2) decreased from north to south. The differences in morphological parameters among sites were significant but smaller than the site differences in fine root biomass and number of root tips.