Nitrogen budgets for Scots pine and Norway spruce ecosystems 12 and 7 years after the end of long-term fertilisation

The magnitude of nitrogen storage and its temporal change in forest ecosystems are important when analysing global change. For example, the accelerated growth of European forests has been linked to increased nitrogen deposition, but the changes in the N inputs that cause long-term changes in ecosystems have not yet been identified. We used two Swedish forest optimum nutrition experiments with Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) to study the long-term fate of N applied to these forest ecosystems. In the pine experiment, in addition to fertiliser (NPK) application, soil acidity was manipulated by application of lime and dilute sulphuric acid. From the spruce experiment, we selected treatments with similar fertiliser doses as in the pine experiment and with and without lime addition.We quantified various terms in the N budget 12 years (pine) and 7 years (spruce) after the last N addition. In the pine stand the NPK-treatment was the only treatment to produce a significant increase in N in the tree biomass (97% above control), whereas in the spruce stand the N additions increased tree N in all treatment combinations (207% above control). In the pine stand the relative distribution of nitrogen between trees and soil did not vary across treatments, with trees containing around 12% of ecosystem N and humus containing around 44% of soil N. The increases in N stocks in the pine stands were mainly in the soil. In contrast, in the spruce ecosystem trees accumulated most of the added N and the increase in the soil was restricted to the humus layer.In the pine ecosystem, large losses of added N (between 254 and 738 kg ha⁻¹ out of 1040 kg ha⁻¹ added as fertiliser) occurred, whereas in the spruce ecosystem we recovered more N than could be accounted for by inputs (between 250 and 591 kg ha⁻¹). There was no clear pattern in the interaction between acidification/liming and N additions.     

Dead trees and protected polypores in unmanaged north-temperate forest stands of Lithuania

The availability of coarse woody debris (CWD) and distribution of dead trees into categories of mortality (dead standing, broken and uprooted) were investigated in north-temperate forests of central Europe (Lithuania). The studied area comprised 188.7 ha and included 18 different stands 40–130 years of age with a variety of tree species (spruce (Picea abies (L.) Karst.), pine (Pinus sylvestris L.), alder (Alnus glutinosa (L.) Gaertn.), birch (Betula pendula Roth and B. pubescens Ehrh.), aspen (Populus tremula L.), oak (Quercus robur L.), forest types (caricus-sphagnum, vaccinium-myrtillus, oxalis, myrtillus-oxalis, caricus-calamagrostis) and edaphic conditions (peaty, sandy, loamy soils of different moisture). The stands were excluded from wood harvesting for at least 30 years. A total of 11 365 dead trees (over 10 cm in DBH) or 6160.7 m³ of dead wood was found (60.2 trees/ha and 32.6 m³/ha). The volume of CWD per hectare was larger in older stands (r_S=0.78, P<0.01). Tree mortality during the last 2 years consisted of 482 trees and 381 m³, or 1.28 trees/ha×year and 1.01 m³/ha×year. In 25–33% of cases it was wind-related. Uprooted and broken trees were of larger DBH than dead standing. The distribution into the categories of mortality was strongly dependent on tree species (chi-square test, d.f.=10,P=0). Dead standing dominated in CWD of pine and alder. Broken trees comprised almost a half in CWD of aspen, and about one-third in birch, alder and oak. Uprooting most often occurred in spruce, aspen and birch. Edaphic conditions and stand age had a pronounced impact on distribution into mortality categories for spruce (chi-square test, d.f.=20, P<0.00001) and pine (d.f.=8, P≤0.0003). On peat soil, only a minority of trees of both pine and spruce was uprooted, and standing dead prevailed. In CWD of spruce and pine, the proportions of both dead standing and broken decreased and that of uprooted trees increased on mineral soils of higher moisture and bulk density in older stands. By contrast, uprooting in birch and alder occurred less often on more wet sites, where the proportions of standing snags were higher. A total of 41 species of wood-decomposing polypores were found in the study area. Among those, 10 (24%) were of conservation value.     

Long-term trends in loblolly pine productivity and stand characteristics in response to thinning and fertilization in the West Gulf region

Two levels each of thinning and fertilization were applied to a 7-year-old loblolly pine (Pinus taeda L.) plantation on a nitrogen- and phosphorus-deficient West Gulf Coastal Plain site in Louisiana. Levels of thinning were no thinning, or thinning applied 7 and 14 years after stand initiation. Levels of fertilization were no fertilization or broadcast fertilization with diammonium phosphate at age 7 years plus refertilization with urea, monocalcium phosphate, and potash at age 14 years. Long-term measurements of climate, stand development and productivity, projected leaf area index, and foliar nutrition were initiated at age 11 years. We found that by age 17 years, thinning increased mean live-crown length from 4.2 to 7.8 m, and mean tree diameter from 15.0 to 21.8 cm compared to the unthinned treatment. After rethinning at age 14 years, stand basal area increased 1.2 and 19.2% between ages 15 and 17 years on the unthinned and thinned plots, respectively. Refertilization at age 14 years reestablished foliar N, P and K sufficiency, which increased leaf area index from 4.2 to 6.0 m² m⁻² on the unthinned plots and from 3.2 to 3.8 m² m⁻² on the thinned plots, and subsequently, increased gross stand biomass from 114 to 141 Mg ha⁻¹ on the unthinned plots and from 78 to 95 Mg ha⁻¹ on the thinned plots by age 17 years. Leaf area was an important factor controlling loblolly pine productivity. At our study site, however, competition for light and water and nutrition-limited foliage growth influenced the variability and scope of this relationship. Our results suggest that a positive and linear relationship between leaf area and loblolly pine productivity does not universally occur on loblolly pine sites.     

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.     

Tree and forest encroachment into fescue grasslands on the Cypress Hills plateau,southeast Alberta,Canada

Tree encroachment into rough fescue (Festuca campestris) grassland has been identified as an ecological concern on the Cypress Hills plateau in southeastern Alberta, Canada. A combination of field sampling (109 transects), a dendrochronological assessment (1361 trees), and a time series analysis of remotely sensed images from five different time periods (1950–2002) were used to determine the extent and rate of tree encroachment and forest development. Tree cover increased by 768 ha (～51%) between 1950 and 2007, representing 10% of the study area, from 1502 ha of pre-1950 forest. Post-1950 tree invasion also created an ingress zone of 750 ha (～10% of study area) based on field transects. Forest cover increased at a consistent rate of 14.3 ha/year. Lodgepole pine (Pinus contorta var. latifolia) was the most common tree species associated with encroachment. Invasion based on the number of established trees occurred at an exponential rate of 3.1%/year after 1890, with density increased by filling spaces adjacent to and between trees within the grassland vegetation. The rate of tree establishment increased to 4.4%/year after 1980, suggesting a change occurred in environmental conditions. Annual atmospheric temperatures increased 0.55 °C from 1929 to 2005 (P < 0.001), with winter (December–February) and spring (March–May) temperatures accounting for most of the increase (P < 0.001), whereas summer (June–August) temperatures slightly decreased (P < 0.050, 0.34 °C) and precipitation increased (P < 0.005, 30 mm). Cattle and elk (Cervus elaphus) grazing was not considered a primary factor for explaining tree encroachment. Based on multidimensional scaling, lodgepole pine establishment was associated with warmer spring temperatures and greater fall (October–November) precipitation. A landscape devoid of wildfires, combined with greater moisture availability, and a longer frost-free season is likely conducive to the sustained establishment of coniferous trees and forest development within the Cypress Hills fescue grassland ecosystem. Long-term conservation of the fescue grasslands could be possible by reintroducing fire.     

Effects of experimental acidification,nitrogen addition and liming on ground vegetation in a mature stand of Norway spruce (Picea abies (L.) Karst.) in SE Sweden

During the period 1976–1991, a combined experiment of acidification, liming and nitrogen addition in a mature spruce stand was conducted at Farabol in south-east Sweden. The aim of this study was to investigate the effects of these treatments on the ground vegetation 0, 1, 5 and 15 years after experimental establishment. The treatment regimes were nitrogen (200 kg N ha⁻¹, repeated three times at 4–5-year intervals, totally 600 kg N ha⁻¹), sulphur powder (50 and 100 kg S ha⁻¹ a⁻¹, totally 600 and 1200 kg ha⁻¹), sulphur plus nitrogen (600+600 kg ha⁻¹) and limestone (500 kg ha⁻¹ a⁻¹, i.e. totally 6000 kg ha⁻¹). The results showed that nitrogen addition and liming promoted the abundance of the grass Deschampsia flexuosa, while acidification had a negative effect on D. flexuosa and herbs in the field layer. There was a negative reaction giving immediate damage to the bryophytes in connection with additions of nitrogen, sulphur powder and lime. The magnitude of damage and the capacity to recover varied among species as well as among treatments. The recovery from immediate damage after liming was much faster than after the treatments with sulphur powder and/or nitrogen. A negative interaction between sulphur powder and nitrogen was found for herbs and mosses where the combined effects were stronger than the effects of a single treatment alone. Acidification also had a negative effect on the total number of species. The results of this study showed that acidification and nitrogen deposition could negatively influence forest vegetation by changing the nutrient availability in the soils. Liming led to an improved growth of the forest ground vegetation and the flora changed towards a more nitrophilic species composition.     

Growth and carbon stocks of a spruce forest chronosequence in central Europe

Human induced changes in global environmental conditions are expected to influence or, as it is hypothesised in this study, have already influenced the biomass and growth of forest ecosystems. In this study, we reconstruct the history of tree growth and quantify the standing biomass along a chronosequence of six Norway spruce stands (Picea abies [L.] Karst; 16–142 years old) on acid soils in a mountainous region with high nitrogen deposition. The inventories of the study sites, as well as the historical stem growth of the sample trees were compared with common yield tables, representing growing conditions before 1960, to find out if and when significant changes in growth of trees had occurred. The growth at tree level (0.003–0.030 m³ yr⁻¹) was about 150–350% higher than predicted by the yield tables, independent of tree age. Because of low stand densities due to early thinning, the increase of stem growth at stand level (90% higher than yield table predictions) and the stand volume (35% higher than yield table predictions) were not as high as the increase of growth at tree level. Total biomass at stand level (including stems, branches, twigs, needles and roots) ranged between 35 and 180 t C ha⁻¹. Net primary productivity varied between 6 and 13 t C ha⁻¹ yr⁻¹. Intensive tree thinning activities probably stimulated growth of remaining trees, but the observed growth rates were beyond what would be expected from these activities exclusively. Thus it is assumed that the fertilisation effects of increased nitrogen deposition and CO₂ concentration, and improved climatic conditions due to ongoing climate change, have contributed to the observed changes in stem growth and that the thinning activities were synergetic with changing environmental conditions. The implications for carbon sinks as accountable under the Kyoto Protocol are probably small, because changes in environmental conditions are not accountable under the Kyoto Protocol and most of the observed changes in growth took place before 1990, the baseline for the Kyoto Protocol. Additionally, it is assumed that impacts on the carbon balance of forest stands due to changes in the thinning regime after 1990, which would be accountable according to article 3.4 of the Kyoto Protocol, are very small without any synergetic changes in environmental conditions.     

Stand development and production dynamics of loblolly pine under a range of cultural treatments in north-central Florida USA

The objectives of this study were to examine the effects of stand development and soil nutrient supply on processes affecting the productivity of loblolly pine (Pinus taeda L.) over a period approximately equal to a pulpwood rotation (18 years). The experiment consisted of a 2×2 factorial combination of complete and sustained weed control and annual fertilization treatments (C: control treatment, F: fertilization, W: weed control, FW: combined fertilization and weed control), located on a Spodosol in north-central Florida, USA. The reduction of soil nutrient limitations through fertilization or control of competing vegetation resulted in dramatic increases in almost every measure of productivity investigated, including height (19.7 m in the FW treatment versus 12.5 m in the C treatment at age 18 years), basal area (FW=44.2 m² ha⁻¹, F=39.6 m² ha⁻¹, W=36.6 m² ha⁻¹, C=19.9 m² ha⁻¹ at age 16 years), stemwood biomass accumulation (114 Mg ha⁻¹ in FW versus 42.8 Mg ha⁻¹ in C at age 18 years), foliar nitrogen concentration (1.53% in plots receiving fertilization versus 1.06% in unfertilized plots at age 17 years) and leaf area index (age 16-year peak projected of approximately 3.3 at age 9–10 years in F and FW plots, 2.5 in the W treatment and 1.5 in the C plots). Cultural treatments also decreased the growth ring earlywood/latewood ratio, and accelerated the juvenile wood to mature wood transition. While soil nutrient supply was a major determinant of productivity, production changes that occurred within treatments over the course of stand development were equally dramatic. For example, between age 8 and 15 years, stemwood PAI in the FW treatment declined by 275%; similarly large reductions occurred in the F and W treatments over the same time period. The reductions in PAI in the treated plots were linearly related to stand BA, suggesting the decline in productivity was associated with the onset of inter-tree competition. Responses of stemwood PAI to re-fertilization treatments at age 15 years suggests that the declines in growth and growth efficiency with time were partially attributable to nutrient limitations.     

Production,allocation, and stemwood growth efficiency of Pinus taeda L. stands in response to 6 years of intensive management

Loblolly pine (Pinus taeda L.) is a highly plastic species with respect to growth responses to forest management. Loblolly pine is the most planted species across the southern United States, a region with the most expansive and intensively managed forest plantations in the world. Management intensity, using tools such as site preparation and fertilization, is increasing greatly in scope over time. To better define to the productive potential of loblolly pine under intensive management, the influence of 6 years of management with weed control (W), weed control plus irrigation (WI), weed control plus irrigation and fertigation (irrigation with a fertilizer solution) (WIF), or weed control plus irrigation, fertigation, and pest control (WIFP) since plantation establishment on stand productivity in loblolly pine was examined. The site is located near Bainbridge, GA (30°48′N latitude and 84°39′W longitude) and is of medium quality (site index=18 m, base age 25). Increasing management intensity greatly accelerated stand development and biomass accumulation. At age 6 total production (above plus belowground) was nearly doubled from 50 to 93 Mg ha⁻¹ in WIFP stands compared to W stands, and standing stem biomass increased from 24 Mg ha⁻¹ in W stands to 48 Mg ha⁻¹ in response to WIFP treatment. Stem current annual increment (CAI) peaked at age 5 in the WIF and WIFP stands at 17–18 Mg ha⁻¹ per year at a basal area between 18 and 21 m² ha⁻¹. Year to year variation in CAI was better explained by previous-year leaf area index (LAI) than current-year LAI. Maximum stemwood production in loblolly pine was achieved through large increases in LAI and small decreases in allocation to woody roots (tap+coarse roots) versus woody shoots (stem+branches) associated with intensive treatments.     

Height growth of planted conifer seedlings in relation to solar radiation and position in Scots pine shelterwood

Seedlings of different provenances of Scots pine (Pinus sylvestris L.), lodgepole pine (Pinus contorta Dougl., var. latifolia Engelm.) and Norway spruce (Picea abies (L.) Karst.) were planted in three Scots pine shelterwoods (125, 65 and 43 stems ha⁻¹) and a clear-cut, all in northern Sweden. The sites were mounded and planting took place during 2 consecutive years (1988 and 1989). The solar radiation experienced by the individual seedlings was determined using a simulation model. Height development of the seedlings was examined during their first 6 years after planting. During the final 3 years of the study, height growth of Norway spruce was relatively poor, both in the shelterwoods and the clear-cut area. Height growth of lodgepole pine was significantly greater than that of Scots pine, both in the shelterwoods and the clear-cut. In contrast to Norway spruce, Scots pine and lodgepole pine displayed significantly greater height growth in the clear-cut than in the shelterwoods. For all three species in the shelterwoods, regression analyses showed that height growth was more strongly correlated with the distance to the nearest tree than with the amount of radiation reaching the ground, i.e. growth was reduced in the vicinity of shelter trees. Therefore, we conclude that the significant reduction in height growth of seedlings of Scots pine and lodgepole pine in Scots pine shelterwoods was partially caused by factors associated with the distance to the nearest shelter tree. Because the substrate was a nitrogen-poor sandy soil, we suggest that root competition for mineral nutrients, especially nitrogen, accounts for the reduction in height growth.     

Genotype × environment interactions in selected loblolly and slash pine plantations in the Southeastern United States

Few studies have quantified the combined effects of silvicultural treatments and genetic improvement on unit area production of full-sib family blocks of loblolly (Pinus taeda L.) and slash pine (P. elliottii Engelm. var. elliotttii). Efficient operational deployment of genetic materials requires an understanding of possible site and silvicultural treatment interactions to maximize yield potential. We examined genotype (family) by environmental interactions (G × E) through age 5 years using a factorial experiment consisting of silvicultural treatment intensity (operational versus intensive), planting density (1334 versus 2990 trees ha⁻¹) and families (seven elite full-sib loblolly and six elite full-sib slash pine families). In January of 2000, randomized complete block, split-plot experiments were installed at two locations for each species in southeast Georgia and northeast Florida. Five years after planting, both loblolly and slash pine demonstrated significant interactions among several factors: genotype × location (p < 0.028 and p < 0.016, respectively), genotype × silvicultural treatment intensity (p < 0.055 and p < 0.059), and silvicultural treatment intensity × density (p < 0.002 and p < 0.001) for basal area (BA) and standing stem volume (VOL). Genotype × silvicultural treatment interactions were positive, with the best overall performing families responding the greatest to intensive treatment. There were changes in slash pine family rankings between locations, which were partly explained by reductions in growth associated with a combination of fusiform rust infection [Cronartium quercum (Berk.) Miyabe ex Shirai f. sp. fusiforme] and wind damage from the 2004 hurricane season. No three-way interactions, which included family, were evident and all genetic sources were stable across the contrasting planting densities. At age 5, loblolly pine outperformed slash pine (p < 0.0001), especially under the intensive silvicultural intensity. While loblolly performance was similar whether deployed in mixtures or pure family blocks, slash pine tended to be more productive in intimate mixtures than when grown in pure family blocks (p = 0.0754 for aboveground biomass).     

Stand and landscape level effects of a major outbreak of spruce beetles on forest vegetation in the Copper River Basin,Alaska

From 1989 to 2003, a widespread outbreak of spruce beetles (Dendroctonus rufipennis) in the Copper River Basin, Alaska, infested over 275,000 ha of forests in the region. During 1997 and 1998, we measured forest vegetation structure and composition on one hundred and thirty-six 20-m × 20-m plots to assess both the immediate stand and landscape level effects of the spruce beetle infestation. A photo-interpreted vegetation and infestation map was produced using color-infrared aerial photography at a scale of 1:40,000. We used linear regression to quantify the effects of the outbreak on forest structure and composition. White spruce (Picea glauca) canopy cover and basal area of medium-to-large trees [≥15 cm diameter-at-breast height (1.3 m, dbh)] were reduced linearly as the number of trees attacked by spruce beetles increased. Black spruce (Picea mariana) and small diameter white spruce (<15 cm dbh) were infrequently attacked and killed by spruce beetles. This selective attack of mature white spruce reduced structural complexity of stands to earlier stages of succession and caused mixed tree species stands to lose their white spruce and become more homogeneous in overstory composition. Using the resulting regressions, we developed a transition matrix to describe changes in vegetation types under varying levels of spruce beetle infestations, and applied the model to the vegetation map. Prior to the outbreak, our study area was composed primarily of stands of mixed white and black spruce (29% of area) and pure white spruce (25%). However, the selective attack on white spruce caused many of these stands to transition to black spruce dominated stands (73% increase in area) or shrublands (26% increase in area). The post-infestation landscape was thereby composed of more even distributions of shrubland and white, black, and mixed spruce communities (17–22% of study area). Changes in the cover and composition of understory vegetation were less evident in this study. However, stands with the highest mortality due to spruce beetles had the lowest densities of white spruce seedlings suggesting a longer forest regeneration time without an increase in seedling germination, growth, or survival.     

Effect of burnt wood removal on the natural regeneration of Pinus halepensis after fire in a pine forest in Tus valley (SE Spain)

To determine the effect of burnt tree removal on post-fire natural regeneration of Pinus halepensis, two 2 500 m² areas were selected six months after the fire in a totally destroyed mature (>70 years) pine forest. In one area, all the trees were cut down and removed 10 months after the fire and, in the other, all the trees were left standing (control). In each area, 20 permanent plots of 20 m² each were randomly placed, and all seedlings emerging within them labeled by individual numbered plastic tags. Emergence, mortality, density and growth (height) of 6649 P. halepensis seedlings were monitored during the first four post-fire years. Seedling emergence was concentrated in the first post-fire autumn–winter period. No positive effect on seedling emergence was detected as a consequence of burnt trunk dragging and subsequent turning over of soil. Wood removal produced an immediate average seedling mortality of 33%, and notably increased seedling mortality during the subsequent summer, probably due to increased exposure of seedlings to sunlight and the possible debilitation of many individuals by mechanical contact during burnt wood removal. A negative correlation of pine seedling mortality with height was detected, which increased significantly on wood removal in the third post-fire year. That is, short seedlings (<10 cm) in treated plots were the most likely to die during this period. In spite of the detrimental effect of wood removal on sapling survival, seedling density four years after fire in the cleared area was large (3.3 seedlings/m²). Wood removal treatment reduced seedling growth: seedling height was significantly higher in the control stand, and differences in seedling height growth rate became particularly noticeable in the fourth post-fire year. The results denote that traditional wood removal practices do not threaten natural post-fire P. halepensis re-establishment if initial seedling density is large enough. However, further studies focused on wood removal effects on the final tree development level and other ecological aspects are necessary to choose adequate post-fire forest management.     

Impacts of atmospheric deposition on New Jersey pine barrens forest soils and communities of ectomycorrhizae

A gradient of increasing N deposition was identified in a southwestern to northeastern transect through the New Jersey pine barrens. The effect of this change in N deposition rate on soil chemistry and ectomycorrhizal morphotype community of pitch pine was studied by sampling from the field under mature pine trees, by planting bait seedlings into the field and in a greenhouse study where seedlings were given differential rates of N applications (0, 35, 140 kg ha⁻¹ equivalent). The field transect showed a significant but small increase in N deposition from 0.35 to 0.72 kg N ha⁻¹ (during the ca. 6 months of the study) equating to 7.84 ± 0.50 kg ha⁻¹ year⁻¹ at the northernmost site, 5.31 ± 0.70 at the middle and 3.66 ± 0.61 kg ha⁻¹ year⁻¹ N at the southwestern most site. Along this transect the ectomycorrhizal morphotype abundance and richness declined significantly under pitch pine. The decline in richness was significantly correlated with the N deposition rate. Bait pitch pine seedlings planted into one of the field sites and fertilized with increasing levels of N showed a reduction in ectomycorrhizal morphotype richness with increased N addition. In a greenhouse study, pine seedling biomass was inversely related to N addition. Nitrogen content of plants increased with increasing N supply, but P content of plants decreased, suggesting that P is a limiting nutrient in this ecosystem. Extractable N from the upper soil horizons increased in cores to which tree seedlings had been added as N addition increased. This indicates an approach to a critical loading of N for these oligotrophic soils, where N supply exceeds seedling N demand. In treeless cores N supply appears to exceed microbial immobilization potential even when no exogenous N is applied. As N supply to greenhouse seedlings increased, ectomycorrhizal morphotype richness declined. By combining data from the field and greenhouse studies, specific ectomycorrhizal morphotype groups were identified by their response to added N. Cortinarius- and Lactarius-like morphotypes were restricted to low levels of N availability. Suilloid- and Ascomycete-like morphotypes were more abundant as soil N availability increases, whereas Russula-like types showed an inverse relationship to N availability. We discuss the results from these oligotrophic sandy soils in comparison with European data derived from richer soils, where mycorrhizal fungal community responses appear to occur only at much higher levels of exogenous N. We attribute these differences to the evolved adaptations of pitch pine and their symbionts to growth in highly oligotrophic environments.     

Regeneration dynamics of Sitka spruce in artificially created forest gaps

This study examined the variation in the development of naturally regenerated and planted seedlings of Sitka spruce (Picea sitchensis (Bong.) Carr.) within gaps cut in a 32-year-old stand of the same species. The circular gaps were 20 m in diameter and designed to allow sunlight into only half of the gap floor at midsummer given the latitude of 56°45′N. Eight plots (8 m × 3 m) were laid out along a north–south transect through each gap (four within the gap and two each under the closed canopy north and south of the gap). Each plot was sub-divided and seedlings were planted into one part and the other part was left to naturally regenerate. In subsequent seasons, plots were further subdivided into ‘weed free’ and ‘vegetation left untouched’. Results showed that while the two central plots within the gaps had the highest value of canopy openness, the highest accumulated temperature and lowest soil moisture were recorded in plots that received direct sunlight. However, level of germination was significantly higher in the shaded area of the gap than in the part that received direct sunshine suggesting that higher moisture levels in shaded areas are important to successful germination. Minimal germination was recorded in the plots beneath the canopy. Seedling survival was significantly influenced by the influx of competing vegetation, but only in the part of the gaps that received direct sunlight. The success of Sitka spruce regeneration within gaps appears to depend on sufficient moisture and light to support regeneration and early growth, but not too much light to encourage the development of competing vegetation. The permanently shaded areas of the gaps appeared to offer ground conditions with sufficient moisture and light to ensure successful germination and early growth of seedlings, but without excessive competition from other vegetation.     

Ponderosa pine snag densities following multiple fires in the Gila Wilderness,New Mexico

Fires create and consume snags (standing dead trees), an important structural and ecological component of ponderosa pine forests. The effects of repeated fires on snag densities in ponderosa pine forests of the southwestern USA have not been studied. Line intercept sampling was used to estimate snag densities in areas of the Gila Wilderness that had burned one to three times under Wildland Fire Use for Resource Benefit (WFU), a fire management policy implemented since 1974 aimed at restoring natural fire regimes. Twenty randomly located transects were measured in areas burned since 1946; six in once-burned areas, six in twice-burned areas and eight in thrice-burned areas. The mean density ± standard errors of large (>47.5 cm dbh) snags for areas that burned once, twice and thrice was 7.0 ± 2.7, 4.4 ± 1.1 and 4.1 ± 1.3 snags/ha, respectively. Differences in snag densities between once- and multiple-burned areas were significant (F-test; p < 0.05). There was no significant difference in density of large snags between twice- and thrice-burned areas. Proportions of type 1 snags (recently created) were higher in once- and twice-burned areas than in areas that burned three times, likely reflecting high tree mortality and snag recruitment resulting from an initial entry fire. Type 3 snags (charred by previous fire) were more abundant in areas that burned multiple times. The lack of differences in snag densities between areas that burned two and three times suggests that repeated fires leave many snags standing. The increasing proportion of type 3 snags with repeated fires supports this conclusion.     

Dynamics of above- and below-ground biomass and nutrient accumulation in an age sequence of Nothofagus antarctica forest of Southern Patagonia

Nothofagus antarctica (Forster f.) Oersted is a deciduous tree species, which naturally grows on poorly drained or drier eastern sites in the Andes Mountain near Patagonian steppe. Above- and below-ground biomass and nutrients pools were measured in pure even-aged stands at different ages (5–220 years) and crown classes. Functions were fitted for total biomass and nutrients accumulation, and root/shoot ratio of individual trees against age. Total biomass accumulated for mature dominant trees was eight times greater than mature suppressed trees. Biomass root/shoot ratio decreased with age from 1.8 to a steady-state of 0.5. All nutrients concentration (except Ca) decreased with age and varied according to the degree of crown suppression classes. Nutrient concentrations varied between biomass pool components following the order leaves > bark > small branches > fine roots > medium roots > rooten wood > coarse roots > sapwood > heartwood. Total nutrient accumulation followed the order dominant > codominant > intermediate > suppressed trees and its accumulation rate varied over time, e.g. P accumulation rate of dominant trees increased from 0.17 g tree⁻¹ year⁻¹ during regeneration to 1.39 g tree⁻¹ year⁻¹ in mature trees. Nutrients uptake reached a peak during the period of maximum biomass production, and root/shoot ratio of nutrients decreased from its maximum value at 5 years of age (0.6, 4.0, 0.9, 1.5, 1.0 and 2.6 for N, P, K, Ca, S and Mg, respectively) to a steady-state asymptote beyond 50 years of age. Thus, accumulation of nutrients in roots was greater during the regeneration phase of stand development, and nutrient accumulation increased in above-ground over time. Also, nutrient use efficiency increased in mature trees (111–220 years) and decreased in suppressed crown classes. The equations developed for individual trees have been used to estimate stand biomass and nutrient accumulation from forest inventories data. Total stand biomass varied from 62.5 to 133.4 t ha⁻¹ and total nutrients accumulation ranged from 3 kg Mg ha⁻¹ to 1235 kg Ca ha⁻¹. Proposed equations can be used for practical purposes such as to estimate pasture nutrients requirement in a silvopastoral system based on nutrients supply from leaf litter returns, or to determine amelioration practices like debarking stems before harvesting.     

Litter production and organic matter accumulation in exclosures of the Tigray highlands,Ethiopia

To determine annual litter production of regenerating forest areas in the Tigray highlands of northern Ethiopia monthly litter production was monitored over a two-year period in areas with varying degree of vegetation cover restoration. Total annual litter production varied from 30 to 425 g m⁻² and increased significantly where areas were closed for a longer time. Litter production was depending on vegetation cover through an exponential relation and was influenced also by soil fertility. Leaf litter typically constituted between 70 and 85% of total litter production, while contributions of woody and reproductive litter varied according to species composition. Strong seasonality in litterfall was explained by pronounced seasonal variation in rainfall. Standing crop of litter built up once an area was closed for grazing, increasing from around 20 g m⁻² in degraded grazing lands to nearly 600 g m⁻² in an old exclosure. Litter accumulation was mainly determined by litter input, but was also influenced by litter quality, species composition and microclimate development in the restoring forest areas. A detailed study of nine dominant shrub and tree species revealed three distinct litter production patterns, corresponding to drought-deciduous species, evergreen species and (semi-)evergreen Acacia species respectively.     

Quantifying economic losses from wildfires in black pine afforestations of northern Spain

We implemented a fire risk assessment framework that combines spatially-explicit burn probabilities, post-fire mortality models and public auction timber prices, to estimate expected economic losses from wildfires in 155 black pine stands covering about 450 ha in the Juslapeña Valley of central Navarra, northern Spain. A logit fire occurrence model was generated from observed historic fires to provide required fire ignition input data. Wildfire likelihood and intensity were estimated by modeling 50,000 fires with the minimum travel time algorithm (MTT) at 30 m resolution under 97^th percentile fire weather conditions. Post-fire tree mortality due to burning fire intensity at different successional stages ranged from 0.67% in the latest stages to 9.22% in the earliest. Stands showed a wide range of potential economic losses, and intermediate successional stage stands presented the highest values, with about 124 € ha^− 1 on average. A fire risk map of the target areas was provided for forest management and risk mitigation purposes at the individual stand level. The approach proposed in this work has a wide potential for decision support, policy making and risk mitigation in southern European commercial conifer forests where large wildfires are the main natural hazard.     

The influence of nutrient and water availability on carbohydrate storage in loblolly pine

We quantified the effects of nutrient and water availability on monthly whole-tree carbohydrate budgets and determined allocation patterns of storage carbohydrates in loblolly pine (Pinus taeda) to test site resource impacts on internal carbon (C) storage. A factorial combination of two nutrient and two irrigation treatments were imposed on a 7-year-old loblolly pine stand in the Sandhills of North Carolina. Monthly collections of foliage, branch, stem, bark, and root tissues were made and total non-structural carbohydrate analyses were performed on samples collected in years 3 and 4 after treatment initiation. Seasonal fluxes of carbohydrates reflected the hypothesized use and storage patterns. Starch concentrations peaked in the spring in all tissues measured; however, minimum concentrations in aboveground tissue occurred in late winter while minimum concentrations in below ground tissue occurred in late fall. Increased nutrient availability generally decreased starch concentrations in current year tissue, while increasing starch in 1-year-old woody tissue. Irrigation treatments did not significantly impact carbohydrate flux. The greatest capacity for starch storage was in below ground tissue, accounting for as much as 400 kg C/ha per year, and more than 65% of the total stored starch C pool. The absolute amount of C stored as starch was significantly increased with increased nutrient availability, however, its relative contribution to the total annual C budget was not changed.