Recovery of photosynthetic capacity in Scots pine: a model analysis of forest plots with contrasting soil temperature

Both aboveground and belowground climate affects net primary production (NNP) and forest growth. Little is known about how above and belowground factors interact. The BIOMASS-model was tested to simulate photosynthetic recovery over a wide range of soil temperatures created by snow cover manipulations on tree-scale plots in a 20-year-old Scots pine stand in northern Sweden. The differences in timing of soil warming between the plots covered a span of two months. Carbon assimilation in needles, sap flow, needle water potential and climatic parameters were measured in the field. The simulations revealed that an early start of soil warming gave a relatively early photosynthetic recovery and a 7.5% increase of NPP. Late soil warming delayed the photosynthetic recovery and reduced the NPP by 13.7%. This indicated that soil temperature needed to be accounted for, as well as air temperature, when analysing photosynthetic recovery and NPP in boreal environment. The effects of differences in soil temperature were reflected in the simulated photosynthetic recovery. The model did not fully capture the delay of photosynthetic recovery caused by a late soil warming. It was possible to integrate the complexity of the soil climate effects into a threshold date for soil thaw, using sapflow measurements together with information about air temperature and a day degree sum, as long as water availability was not limiting water uptake by roots. Although a more realistic mechanism than that currently in BIOMASS is desirable as climate change shifts the typical patterns of interplay between air and soil temperature dynamics.     

Dynamics and potentials of carbon sequestration in managed stands and wood products in Finland under changing climatic conditions

Carbon (C) sequestration was studied in managed boreal forest stands and in wood products under current and changing climate in Finland. The C flows were simulated with a gap-type forest model interfaced with a wood product model. Sites in the simulations represented medium fertile southern and northern Finland sites, and stands were pure Scots pine and Norway spruce stands or mixtures of silver and pubescent birch.

Changing climate increased C sequestration clearly in northern Finland, but in southern Finland sequestration even decreased. Temperature is currently the major factor limiting tree growth in northern Finland. In southern Finland, the total average C balance over the 150 year period increased slightly in Scots pine stands and wood products, from 0.78 Mg C ha⁻¹ per year to 0.84 Mg C ha⁻¹ per year, while in birch stands and wood products the increase was larger, from 0.64 Mg C ha⁻¹ per year to 0.92 Mg C ha⁻¹ per year. In Norway spruce stands and wood products, the total average balance decreased substantially, from 0.96 Mg C ha⁻¹ per year to 0.32 Mg C ha⁻¹ per year. In northern Finland, the total average C balance of the 150 year period increased under changing climate, regardless of tree species: in Scots pine stands and wood products from 1.10 Mg C ha⁻¹ per year to 1.42 Mg C ha⁻¹ per year, in Norway spruce stands and wood products from 0.69 Mg C ha⁻¹ per year to 0.99 Mg C ha⁻¹ per year, and in birch stands and wood products from 0.43 Mg C ha⁻¹ per year to 0.60 Mg C ha⁻¹ per year.

C sequestration in unmanaged stands was larger than in managed systems, regardless of climate. However, wood products should be included in C sequestration assessments since 12–55% of the total 45–214 Mg C ha⁻¹ after 150 years' simulation was in products, depending on tree species, climate and location. The largest C flow from managed system back into the atmosphere was from litter, 36–47% of the total flow, from vegetation 22–32%, from soil organic matter 25–30%. Emissions from the production process and burning of discarded products were 1–6% of the total flow, and emissions from landfills less than 1%.     

Regional assessment of boreal forest productivity using an ecological process model and remote sensing parameter maps

An ecological process model (BIOME-BGC) was used to assess boreal forest regional net primary production (NPP) and response to short-term, year-to-year weather fluctuations based on spatially explicit, land cover and biomass maps derived by radar remote sensing, as well as soil, terrain and daily weather information. Simulations were conducted at a 30-m spatial resolution, over a 1205 km(2) portion of the BOREAS Southern Study Area of central Saskatchewan, Canada, over a 3-year period (1994-1996). Simulations of NPP for the study region were spatially and temporally complex, averaging 2.2 (+/- 0.6), 1.8 (+/- 0.5) and 1.7 (+/- 0.5) Mg C ha(-1) year(-1) for 1994, 1995 and 1996, respectively. Spatial variability of NPP was strongly controlled by the amount of aboveground biomass, particularly photosynthetic leaf area, whereas biophysical differences between broadleaf deciduous and evergreen coniferous vegetation were of secondary importance. Simulations of NPP were strongly sensitive to year-to-year variations in seasonal weather patterns, which influenced the timing of spring thaw and deciduous bud-burst. Reductions in annual NPP of approximately 17 and 22% for 1995 and 1996, respectively, were attributed to 3- and 5-week delays in spring thaw relative to 1994. Boreal forest stands with greater proportions of deciduous vegetation were more sensitive to the timing of spring thaw than evergreen coniferous stands. Similar relationships were found by comparing simulated snow depth records with 10-year records of aboveground NPP measurements obtained from biomass harvest plots within the BOREAS region. These results highlight the importance of sub-grid scale land cover complexity in controlling boreal forest regional productivity, the dynamic response of the biome to short-term interannual climate variations, and the potential implications of climate change and other large-scale disturbances.     

Effects of irrigation and coppicing on above-ground growth, physiology, and fine-root dynamics of two field-grown hybrid poplar clones

To determine the effects of irrigation and coppicing on above- and below-ground growth dynamics, a plantation of Populus × euramericana cv. ‘Eugenei’ and Populus tristis × Populus balsamifera cv. ‘Tristis 1’ was established in May 1984 on a level site in East Lansing, MI, USA. Supplemental water in the form of drip irrigation was applied to half the trees beginning the first growing season. All trees were cut down in March 1988 and the stumps allowed to coppice. Pre- and post-coppice height and diameter growth of ‘Eugenei’ always exceeded that of ‘Tristis’, and the former clone showed a positive response to irrigation, whereas the latter did not. The greater growth of ‘Eugenei’ was primarily due to its full use of the growing season. Post-coppice rates of photosynthesis were not affected by irrigation in either clone, but stomatal conductances were reduced in non-irrigated trees. Analysis of microvideo images taken in minirhizotron tubes buried in the soil close to the trees showed that ‘Tristis’ produced a greater length and number of fine roots in the top 30 cm of soil than ‘Eugenei’, regardless of treatment. Irrigated trees consistently produced more fine roots at 0–30 cm soil depth than non-irrigated trees only in ‘Eugenei’, but both clones showed greater fine-root production in non-irrigated trees at 30–100 cm. Both clones also showed substantial fine-root production in the spring immediately following coppicing, with no evidence of a shock-induced dieback of roots. The root systems of these two poplar genotypes apparently contain sufficient carbon and nitrogen reserves to fuel a spring flush of fine-root growth, even though the tops have been severed during the dormant season.     

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.     

Coarse woody debris dynamics in a post-fire jack pine chronosequence and its relation with site productivity

The long-term relationships between coarse woody debris (CWD) dynamics, soil characteristics and site productivity have, so far, received little attention. The objectives of the study were to describe CWD dynamics along a post-fire chronosequence (43–86 years after fire) in jack pine (Pinus banksiana Lamb.) stands, assess the importance of buried CWD in terms of soil available water holding capacity (AWHC), and investigate relationships between CWD, AWHC, nutrient retention and site productivity.

Twelve jack pine stands on sandy, mesic sites of glaciolacustrine origin were surveyed. Buried wood volume within the forest floor varied between 1 and 57 m³ ha⁻¹ (4–92% of total site CWD volume) and showed no relationship with time. Downed log mass accumulation followed a “U shaped” successional pattern with time since fire. Buried wood AWHC was negligible compared with that of the 0–20 cm mineral soil layer. The most productive sites were characterised by higher forest floor dry weight, effective CEC and water holding capacity in the mineral soil. Path analyses of relationships between organic matter content, CWD and forest floor CEC showed that CEC was conditioned by forest floor organic matter and buried wood content.     

Changes in microclimate after stand conversion in two northern hardwood stands

Changes in air temperature, soil temperature, and soil moisture were monitored for 5 years in two northern hardwood stands after whole-tree harvesting and conversion to red pine (Pinus resinosa Ait.) plantations. Soil temperatures at a depth of 5 cm and maximum air temperatures 2 m above the soil surface were increased 5–25% after stand conversion. Soil moisture content at a depth of 5 cm was increased by 10–20% in one stand but not in the other. Differences in stand, soil, and topographical characteristics between the two stands did not have any apparent effect on the magnitude of air or soil temperature changes after stand conversion. However, higher initial stand density and soil water holding capacity appeared to be related to increased soil moisture content at one of the sites. The increased soil temperatures after conversion were not only a result of the removal of the northern hardwood canopy but also the removal and redistribution of the forest floor caused by whole-tree harvesting. Five years after stand conversion air temperature, soil temperature, and soil moisture showed no evidence of recovering from initial post-harvest levels.     

Nutrient accessions to the forest floor in karri (Eucalyptus diversicolor F. Muell.) forests of varying age

Nutrient accessions to the forest floor in rainfall, throughfall and stemflow from the understorey and tree strata were measured in karri (Eucalyptus diversicolor F. Muell.) forest of south-west Western Australia. These data, for stands aged 2, 6, 9, and 40 years and for mature forest, were compared with annual accessions of nutrients in litterfall.

With increasing age of the stands, the concentration and amounts of Ca, K, Mg, Na and Cl in throughfall increased. The amounts of Mg, Na and Cl in stemflow from the overstorey were also greatest in the older stands. Although concentrations of nutrients in stemflow from the tree stratum were higher than from the understorey, the quantities of nutrients transported to the forest floor in stemflow from the understorey were greater because of its greater volume.

Rainfall is the major source of Na (64–91%) and Cl (51–79%) and it also contributes significantly to Mg (19–50%) accessions. Of the nutrient pathways between canopy and forest floor, litterfall accounts for the largest proportion of N (89–93%), Ca (80–87%) and P (67–79%) recycled in the karri forest. About equal amounts of K are transferred in litterfall, throughfall and stemflow with approximately 80% of stemflow K originating from the understorey strata.     

Forest floor vegetation plays an important role in photosynthetic production of boreal forests

We estimated gross photosynthetic production (GPP) of the forest floor vegetation in a 40-year-old Scots pine stand in southern Finland with three different methods: measurements of CO₂ exchange of single leaves of field and ground layer species, measurement campaigns of forest floor net CO₂ efflux at different irradiances with a manually operated soil chamber, and continuous measurements of forest floor net CO₂ efflux with an automatic transparent chamber system. We upscaled the measured light response curves from the manual soil chambers using the biomass distribution of the forest floor species, a modelled seasonal pattern of photosynthetic capacity and a model of light extinction down the canopy. Leaf gas exchange measurements as well as measurements of net CO₂ efflux with the manual chamber indicated saturation of photosynthesis at relatively low (50–400 μmol m⁻² s⁻¹) light levels. Leaf and patch level measurements gave similar rates of photosynthetic CO₂ fixation per unit leaf biomass suggesting that reduction in photosynthetic production due to within-patch shading was small. Upscaling of photosynthetic production to the stand level and continuous measurements with the automatic soil chambers indicated that momentary photosynthetic production by the forest floor vegetation in the summer was typically about 2 μmol m⁻² (ground) s⁻¹. Cumulative upscaled GPP over the period of no snow (from 20 April to 20 November) in year 2003 was 131 g C m⁻². Continuous measurements with the automatic soil chamber system were in line with the upscaling, the cumulative GPP being 83 g C m⁻² and the seasonal pattern of photosynthetic rate similar to that of the upscaled photosynthesis.     

The ratio of NPP to GPP: evidence of change over the course of stand development

Using Scots pine (Pinus sylvestris L.) in Fenno-Scandia as a case study, we investigate whether net primary production (NPP) and maintenance respiration are constant fractions of gross primary production (GPP) as even-aged mono-specific stands progress from initiation to old age. A model of the ratio of NPP to GPP is developed based on (1) the classical model of respiration, which divides total respiration into construction and maintenance components, and (2) a process-based model, which derives respiration from processes including construction, nitrate uptake and reduction, ion uptake, phloem loading and maintenance. Published estimates of specific respiration and production rates, and some recent measurements of components of dry matter in stands of different ages, are used to quantify the two approaches over the course of stand development in an average environment. Both approaches give similar results, showing a decrease in the NPP/GPP ratio with increasing tree height. In addition, we show that stand-growth models fitted under three different sets of assumptions-(i) annual specific rates of maintenance respiration of sapwood (mW) and photosynthesis (sC) are constant; (ii) m(W) is constant, but sC decreases with increasing tree height; and (iii) total maintenance respiration is a constant fraction of GPP and s(C) decreases with increasing tree height-can lead to nearly identical model projections that agree with empirical observations of NPP and stand-growth variables. Remeasurements of GPP and respiration over time in chronosequences of stands may be needed to discern which set of assumptions is correct. Total (construction + maintenance) sapwood respiration per unit mass of sapwood (kg C (kg C year)-1) decreased with increasing stand age, sapwood stock, and average tree height under all three assumptions. However, total sapwood respiration (kg C (ha year)-1) increased over the course of stand development under (i) and (ii), contributing to a downward trend in the time course of the NPP/GPP ratio after closure. A moderate decrease in mW with increasing tree height or sapwood cross-sectional area had little effect on the downward trend. On the basis of this evidence, we argue that a significant decline in the NPP/GPP ratio with tree size or age seems highly probable, although the decline may appear insignificant over some segments of stand development. We also argue that, because stand-growth models can give correct answers for the wrong reasons, statistical calibration of such models should be avoided whenever possible; instead, values of physiological parameters should come from measurements of the physiological processes themselves.     

Interannual variability of net ecosystem production for a broadleaf deciduous forest in Sapporo, northern Japan

To estimate net ecosystem production (NEP), ecosystem respiration (R _E), and gross primary production (GPP), and to elucidate the interannual variability of NEP in a cool temperate broadleaf deciduous forest in Sapporo, northern Japan, we measured net ecosystem exchange (NEE) using an eddy covariance technique with a closed-path infrared gas analyzer from 2000 to 2003. NEP, R _E, and GPP were derived from NEE, and data gaps were filled using empirical regression models with meteorological variables such as photosynthetic active radiation and soil temperature. In general, NEP was positive (CO₂ uptake) from May to September, either positive or negative in October, and negative (CO₂ release) from November to the following April. NEP rapidly increased during leaf expansion in May and reached its maximum in June or July. The four-year averages (±?standard deviation) of annual NEP, GPP, and R _E were 443?±?45, 1,374?±?39, and 931?±?11?g?C?m^?2?year^?1, respectively. The lower annual NEP and GPP in 2000 may have been caused by lower solar radiation in the foliated season. During the foliated season, monthly GPP varied from year to year more than monthly R _E. Variations in the amount of incoming solar radiation may have caused the interannual variations in the monthly GPP. Additionally, in May, the timing of leaf expansion had a large impact on GPP. Variations in GPP affected the interannual variation in NEP at our site. Thus, interannual variation in NEP was affected by the incoming solar radiation and the timing of leaf expansion.     

Comparison between old-growth stands and secondary stands regenerating after clear-felling in warm-temperate forests of Yakushima, southern Japan

In order to infer successional changes in structure, species composition and diversity of warm-temperate forest, we compared secondary stands regenerating after clear-felling (41–64-years old) with old-growth stands at altitudes between 300 and 800 m on Yakushima Island, southern Japan. Stem density and maximum stem diameter differed between secondary and old-growth stands, but basal area and aboveground biomass did not. At lower altitudes, the dominant species in old-growth stands with a strong sprouting capacity (Castanopsis cuspidata) also dominated secondary stands, and species composition of secondary and old-growth stands was similar. At higher altitudes, by contrast, the dominant species in old-growth stands (Distylium racemosum) had little sprouting capacity and was poorly represented in diverse secondary stands, which were dominated by Castanopsis or other less abundant species. Secondary stands had greater species diversity (Shannon–Wiener index) than old-growth stands, particularly at higher altitudes. This was due to greater species richness resulting from higher stem density per area, but not to greater evenness. We grouped the component species that share ecologically similar traits into four guilds (fagaceous, primary evergreen, secondary evergreen and deciduous species). Secondary stands were characterized by greater numbers of deciduous and secondary evergreen species. We concluded that different sprouting capacities of dominant species and different regeneration traits among guilds are responsible for the change in species composition and diversity during succession.     

Changes in net ecosystem productivity with forest age following clearcutting of a coastal Douglas-fir forest: testing a mathematical model with eddy covariance measurements along a forest chronosequence

We hypothesized that changes in net ecosystem productivity (NEP) during aging of coastal Douglas-fir (Pseudotsuga menziesii Mirb. Franco) stands could be explained by (1) changing nutrient uptake caused by different time scales for decomposition of fine, non-woody and coarse woody litter left after harvesting, (2) declines in canopy water status with lengthening of the water uptake pathway during bole and branch growth, and (3) increases in the ratio of autotrophic respiration (R (a)) to gross primary productivity (GPP) with phytomass accumulation. These hypotheses were implemented and tested in the mathematical model ecosys against eddy covariance (EC) measurements of forest CO(2) and energy exchange in a post-clearcut Douglas-fir chronosequence. Hypothesis 1 explained how (a) an initial rise in GPP observed during the first 3 years after clearcutting could be caused by nutrient mineralization from rapid decomposition of fine, non-woody litter with lower C:N ratios (assart effect), (b) a slower rise in GPP during the next 20 years could be caused by immobilization during later decomposition of coarse woody litter, and (c) a rapid rise in GPP between 20 and 40 years after clearcutting could be caused by nutrient mineralization with further decomposition of coarse woody litter and of its decomposition products. During periods (a) and (b), heterotrophic respiration (R (h)) from decomposition of fine and coarse litter greatly exceeded net primary productivity (NPP = GPP - R (a)) so that Douglas-fir stands were large sources of CO(2). During period (c), NPP exceeded R (h) so that these stands became large sinks for CO(2). Hypothesis 2 explained how declines in NPP during later growth in period (c) could be caused by lower hydraulic conductances in taller trees that would force lower canopy water potentials and hence greater sensitivity of stomatal conductances and CO(2) uptake to vapor pressure deficits. Enhanced sensitivity to vapor pressure deficits was also apparent in the EC measurements over the post-clearcut chronosequence. Hypothesis 3 did not contribute to the explanation of forest age effects on NEP.     

Logging history (1820–2000) of a heavily exploited southern boreal forest landscape: Insights from sunken logs and forestry maps

Over the last two centuries, logging has caused major, but unquantified, compositional and structural changes in the southern portion of the North American boreal forest. In this study, we used a series of old forest inventory maps coupled with a new dendrochronological approach for analyzing timber floating histories in order to document the long-term transformation (1820–2000) of a southern boreal landscape (117 000 ha) in eastern Quebec, Canada, in response to logging practices. Landscape exploitation became increasingly severe throughout this time period. During the ninetieth century (1820–1900) of limited industrial capacity, selective logging targeted pine and spruce trees and excluded balsam fir, a much abundant species of the forest landscape. Logging intensity increased during the first half of the twentieth century, and targeted all conifer species including balsam fir. After 1975, dramatic changes occurred over the landscape in relation to clear-cutting practices, plantations, and salvage logging, which promoted the proliferation of regenerating areas and extensive plantations of the previously uncommon black spruce. Overall, logging disturbance resulted in an inversion in the forest matrix, from conifer to mixed and deciduous, and from old to regenerating stands, thus creating significant consequences on forest sustainability. If biodiversity conservation and sustainable forestry are to be management goals in such a heavily exploited forested landscape, then restoration strategies should be implemented in order to stop the divergence of the forests from their preindustrial conditions.     

Does canopy mean nitrogen concentration explain variation in canopy light use efficiency across 14 contrasting forest sites?

The maximum light use efficiency (LUE?=?gross primary production (GPP)/absorbed photosynthetic photon flux density (aPPFD)) of plant canopies has been reported to vary spatially and some of this variation has previously been attributed to plant species differences. The canopy nitrogen concentration [N] can potentially explain some of this spatial variation. However, the current paradigm of the N-effect on photosynthesis is largely based on the relationship between photosynthetic capacity (A(max)) and [N], i.e., the effects of [N] on photosynthesis rates appear under high PPFD. A maximum LUE-[N] relationship, if it existed, would influence photosynthesis in the whole range of PPFD. We estimated maximum LUE for 14 eddy-covariance forest sites, examined its [N] dependency and investigated how the [N]-maximum LUE dependency could be incorporated into a GPP model. In the model, maximum LUE corresponds to LUE under optimal environmental conditions before light saturation takes place (the slope of GPP vs. PPFD under low PPFD). Maximum LUE was higher in deciduous/mixed than in coniferous sites, and correlated significantly with canopy mean [N]. Correlations between maximum LUE and canopy [N] existed regardless of daily PPFD, although we expected the correlation to disappear under low PPFD when LUE was also highest. Despite these correlations, including [N] in the model of GPP only marginally decreased the root mean squared error. Our results suggest that maximum LUE correlates linearly with canopy [N], but that a larger body of data is required before we can include this relationship into a GPP model. Gross primary production will therefore positively correlate with [N] already at low PPFD, and not only at high PPFD as is suggested by the prevailing paradigm of leaf-level A(max)-[N] relationships. This finding has consequences for modelling GPP driven by temporal changes or spatial variation in canopy [N].     

Seasonal variations in photosynthesis of Picea morrisonicola growing in the subalpine region of subtropical Taiwan

From January 1999 to May 2001, we investigated seasonal variations in the photosynthetic capacity of Taiwan spruce (Picea morrisonicola Hay.) growing in the subalpine region of subtropical Taiwan (23 degrees 29' N, 120 degrees 53' E, 2600 m a.s.l.). Photosynthetic capacity (near light-saturated net photosynthetic rate, Pnsat, chlorophyll fluorescence (Fv/Fm) and soluble protein concentration of needles all increased from mid or late spring to early winter. Even when minimum air temperature of the measuring day dropped to near 0 degrees C, Pnsat remained at about 20% of the highest value observed in winter. There was a curvilinear relationship between Fv/Fm and the minimum or mean air temperature of the measuring day. The increase in Fv/Fm with temperature was slowed when the daily mean air temperature was above 7 degrees C, or the minimum air temperature was above 3 degrees C; however, when air temperatures dropped below these values, Fv/Fm varied sharply. Seasonal variations in Pnsat paralleled those in Fv/Fm and needle soluble protein concentration. In early or mid spring when air temperature and Fv/Fm increased, Pnsat and soluble protein concentration remained low. Multiple regression analysis showed that seasonal variations in Pnsat were affected by Fv/Fm, air temperature and needle soluble protein concentration, and the multiple regression equation could be used to estimate Pnsat in different seasons. We conclude that the decrease in photosynthetic capacity of Taiwan spruce in winter and its subsequent recovery in spring were mainly caused by photoinhibition and its reversal, and changes in needle soluble protein concentration. Another possible explanation for the delayed recovery of photosynthetic capacity in spring may be associated with the slow increase in needle soluble protein concentration.     

Long-term responses of ecosystem components to stand thinning in young lodgepole pine forest: IV. Relative habitat use by mammalian herbivores

Pre-commercial thinning (PCT) is a silvicultural practice that can provide diverse understory and overstory vegetation conditions. We tested the hypothesis that relative habitat use by snowshoe hare (Lepus americanus), mule deer (Odocoileus hemionus), and moose (Alces alces) would increase in response to enhanced abundance of herbs and shrubs, and species diversity and structural diversity of conifers, in heavily thinned (≤1000 stems/ha) stands, at 12–15 years post-thinning. Replicate study areas were located near Penticton, Kamloops, and Prince George in south-central British Columbia, Canada. Each study area had three young pine stands thinned to densities of 500 stems/ha (low), 1000 stems/ha (medium), and 2000 stems/ha (high), with an unthinned young pine and old-growth pine stand for comparison.

Relative habitat use, based on counts of fecal pellets and pellet-groups, was similar among the five treatment stands for hares (P = 0.24), deer (P = 0.23), and moose (P = 0.16). However, low-density stands (500 stems/ha) had ca. 3–20 times as many deer pellet-groups, and ca. 2–4 times as many moose pellet-groups, than other stands. Low-density stands had significantly greater canopy openness, volume of shrubs <2 m, and horizontal hiding cover <1.6 m than other treatments. Relative habitat use by deer and moose was positively related to understory characteristics such as enhanced abundance of forage and security cover. These results support our hypothesis that deer and moose responded positively to enhanced volume of herbs and shrubs as well as to species diversity and structural diversity of conifers and overall vegetation in heavily thinned (≤1000 stems/ha) stands at 12–15 years post-thinning. Our results suggest that ungulate management would be enhanced if greater emphasis was placed on forage enhancement throughout the year, which differs from current management recommendations which tend to focus on winter range and snow-interception cover.     

Pasture production in a silvopastoral system in relation with microclimate variables in the atlantic coast of Spain

Grasses and legumes of high productivity and nutritional quality are a good alternative as pasture supplements in rangelands of low quality forage. Orchardgrass (Dactylis glomerata L. cv. `Artabro') and white clover (Trifolium repens L. cv. `Huia') are known as shade tolerant and low flammability species that have been successfully used in agroforestry systems in Galicia, both diminishing fire hazard compared with natural shrublands. In this study, annual and seasonal production of a grass mixture of both species was quantified during 3 years in a pinewood under different tree canopy covers. Regardless of cover, pasture production increased in summer, and decreased from fall to spring. We obtained a significant correlation between annual pasture production and light transmission through the tree canopy (R² = 0.96, P<0.05). Light transmittance through a maritime pine canopy (Pinus pinaster Ait.) was higher than through a Scots pine canopy (P. sylvestris L.), corresponding to 36–57% and 16–21% of full sunlight respectively. The highest herbage production was obtained in no tree stands and the lowest under a P. sylvestris canopy. Fluctuations inlight transmission, temperature and PAR (Photosynthetically Active Radiation) under tree canopy were less apparent compared with no tree stands. Variation in seasonal production was more pronounced in stands without trees, and appeared more uniform when percentage of light intercepted by tree canopy increased.This revised version was published online in November 2005 with corrections to the Cover Date.     

Irradiance in thinned Norway spruce (Picea abies) stands and the possibilities to prevent suckers of broadleaved trees

Both incoming shortwave radiation (R_g) and photosynthetically active radiation (PAR) in percentage of full daylight were measured at the same time by point and strip sampling in four plots (0.1 ha) of Picea abies (L.) Karst. The standard deviations (%) of R_g and PAR were, respectively, 11.1 and 9.8 at 64 points, 15.7 and 13.9 at 32 points, and 24.7 and 23.8 at 16 points per plot.

A period of at least 40 s per strip (30 m min⁻¹) gives a CV (coefficient of variation) of 30%. There is no significant difference between relative irradiance (RI) estimated by the point method (64 points) and by the strip method (8 strips). Curves of RI (R_g and PAR) and basal area (m² ha⁻¹), diameter sum (m ha⁻¹) and density (stems ha⁻¹) of fifteen trials with different thinning programmes are presented. Irradiance (R_g) in heavily thinned stands was 3–14% of irradiance on an open place. The irradiance, R_g, in extra-heavily thinned stands is 12–27%, and in unthinned stands, 1–3% that of an open place. The R_g curve lies above the PAR curve in all cases. Some practical implications of the study are presented. Heavy thinning of Norway spruce stands gives RI (R_g) values 10% at basal area of 25m² ha⁻¹ which is necessary to minimize development of suckers of broadleaved trees.     

Mountain pine beetle‐associated blue‐stain fungi in lodgepole × jack pine hybrids near Grande Prairie,Alberta (Canada)

The mountain pine beetle (MPB), the most serious pest of lodgepole pine in mountainous western Canada, spread northeastward into lodgepole × jack pine hybrids in the boreal forest of Alberta in 2006. The MPB vectors three species of blue‐stain fungi, which contribute to the success of the beetles. These fungi were isolated from MPB larvae and galleries in several lodgepole × jack pine stands in the Grande Prairie region of northwestern Alberta in autumn 2006 and winter and spring 2007. Fungi were recovered from more than 95% of gallery systems. The three fungi were similarly prevalent but Ophiostoma montium was the most frequently isolated fungus at each sampling point, isolated from 72% to 90% of gallery systems compared with 63% to 78% for Grosmannia clavigera, and 61% to 86% for Leptographium longiclavatum. Ophiostoma montium and G. clavigera were isolated from more larvae than gallery samples, with the opposite true for L. longiclavatum. Most gallery systems contained multiple fungi with three fungi per gallery system being more common in autumn and winter and two fungi more common in the spring. The combination of G. clavigera and L. longiclavatum was less common among gallery systems with two fungi than either of the pairwise combinations containing O. montium. Fungal prevalence was the same above and below snow level. The prevalence of the three fungi did not differ significantly among stands sampled in the spring but stands with more G. clavigera tended to have less L. longiclavatum. The winter of 2006–2007 was colder than average throughout Alberta with temperatures below ?30°C in November, January and February, and all three fungi were present after the cold winter while most larvae had died, suggesting that overwintering mortality of the fungi will not limit persistence and spread of MPB in the boreal forest.