Understory abundance,species diversity and functional attribute response to thinning in coniferous stands

Alternative strategies for stand density management in even-aged coniferous forests may increase plant species and functional diversity. We examined the effects of fixed and variable density thinning on tree seedling regeneration as well as on abundance (indexed by cover) and richness of understory vascular plants 11 years after harvesting 45- to 66-year old forests dominated by Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) or western hemlock (Tsuga heterophylla (Raf.) Sarg.) at three sites in western Oregon. Each site contained an unthinned control (CON), and thinning treatments selected to enhance overstory structural diversity and spatial variability within stands (HD, high density treatment at 300 trees ha⁻¹; MD, moderate density treatment at 200 trees ha⁻¹; VD300, VD200 and VD100, variable density treatments at 300, 200 and 100 trees ha⁻¹). Leave islands are included in HD and the other thinning treatments contain both leave islands and gap openings. Tree seedling regeneration was highly variable and generally increased with thinning. Cover of all understory species was greater in VD100 than in the control whereas richness was greater in HD and MD. Cover and richness of early seral species were greater in most thinning treatments than in the control. Understory plant communities were overwhelmingly dominated by native species. In general, vegetation dynamics was accelerated by thinning, especially in variable density treatments. Cover of N-fixing understory species was greater in VD200 than in the other treatments, and in MD and VD300 than in the control, whereas richness of understory N-fixing species increased in all thinning treatments. Cover of understory species with intermediate soil water requirements was greater in MD, VD200 and VD100 than in the control, whereas richness of these species increased in VD200 compared to the control, HD and VD300. Thinning promoted higher diversity of understory conditions without reducing density and species richness of crop tree regeneration, and seemed to increase functional effect and response diversity.     

Pre-commercial thinning,birch admixture and sprout management in planted Norway spruce stands in South Sweden

Early management of the regenerated seedlings shapes the future stand properties. To address these issues, pre-commercial thinning (PCT) and control treatments were applied to planted Norway spruce (Picea abies L. Karst) and naturally regenerated birch (Betula pendula Roth., Betula pubescens Ehrh.) stands in forest experiments in southern Sweden (lat. 56–57?N) containing 1.1–5.5?m tall saplings. The treatments were retention of 1000 or 2000?stems?ha^?1 of Norway spruce, with no birch or birch at 1000?stems?ha^?1. Treatments were replicated with and without annual removal of birch sprouts from stumps. The periodic annual increment (PAI) over five years was calculated for total stand volume and individual trees. The mean PAI of dominant trees was significantly higher both following all PCT treatments than controls, and following low rather than high-density PCT. Birch retention did not affect growth of the dominant trees but PAI was lower in plots with uncontrolled sprouting. The PAI of birch was significantly higher in low-density Norway spruce plots than in control plots and the high-density plots. The treatment response was significant even in stands with initial heights of only 1–2?m.     

Stem Quality and Height Development in a Betula-Dominated Stand Seven Years After Precommercial Thinning at Different Stump Heights

Topping may provide an attractive alternative to traditional precommercial thinning, offering several advantages. The main stem quality of birch (Betula pubescens Ehrh. and Betula pendula Roth, species not separated) was examined seven growing seasons after such topping in a dense, even-aged and naturally regenerated stand in northern Sweden. The height development and survival of main and secondary stems of birch and secondary stems of willow (Salix spp.) were also studied. At a main stem height of 1.8 m, the following treatments, leaving 3100 main stems ha^?1, were applied to secondary stems: traditional precommercial thinning (cutting just above ground), top-cutting at either 40% (71 cm above ground) or 70% (120 cm above ground) of main stem mean height, and no precommercial thinning. For main stems, topping resulted in fewer forks, straighter stems, higher live crown height and smaller diameter of the thickest branch compared with traditional precommercial thinning. In these respects, topping at the higher level above ground gave the best quality. The main stems were no longer at risk of becoming overtopped by the secondary stems following any of these treatments (except for no precommercial thinning).     

Timing and intensity of precommercial thinning in Pinus sylvestris stands

The effects of timing and intensity of precommercial thinning were studied in three Scots pine artificially regenerated stands on Vaccinium forest sites in southern Finland. A two-level factorial design (3×3) was used in each stand: thinning at dominant height of 3, 6 and 9 m to 1000, 1600 and 2200 stems ha^?1. The effects of the treatments were analysed after a period of 23–25 yrs when the dominant height was 14–15 m. Early thinning resulted in the highest standing volume and amount of merchantable wood, and also in slightly accelerated height development. Thinning to 1000 stems ha^?1 caused a considerable production loss, but there were no differences between the densities of 1600 and 2200 stems ha^?1. Branches became thicker after early thinning, but the differences between the treatments were negligible for crop trees. Crown ratio was lowest as the result of early or moderate thinning (2200 stems ha^?1).     

Selective removal of paper birch increases growth of juvenile Douglas-fir while minimizing impacts on the plant community

Broadcast weeding of naturally regenerating paper birch (Betula papyrifera) to improve performance of conifer crop trees is common in forest plantations in British Columbia, Canada, but such treatments are costly, reduce biodiversity and may increase disease. To investigate whether alternative treatments can enhance conifer performance with minimal loss of other ecosystem services, we examined effects of selective removal of paper birch in two interior Douglas-fir (Pseudotsuga menziesii var. glauca) plantations on Douglas-fir growth and survival, and cover and height of residual birch and the understorey plant community. Douglas-fir-centred birch-free patches of varying radius (0–2.5?m) were created by single and repeated manual cutting and Triclopyr® application. The “threshold brushing radius,” defined as the birch removal radius at which Douglas-fir performance was optimized was 2.0?m at the more productive site and 1.5?m at the less productive one. Armillaria root disease, the major Douglas-fir mortality agent, spiked in the 1.0?m radius manual cutting treatment at one location. The greatest reductions in birch and understory plant abundance occurred following the most intensive treatments. Selective brushing treatments are an effective alternative to broadcast removal of paper birch for increasing conifer performance, and have less impact on the plant community.     

Stand dynamics and tree quality response to precommercial thinning in a northern hardwood forest of the Acadian forest region: 23 years of intermediate results

In the late 1980s, large forest companies began precommercial thinning (PCT) operations in young northern hardwood cutovers in New Brunswick, Canada. To provide supporting growth and yield information, an industrial experiment was established at residual stand densities of 1300, 1600, 1900, and 2200?stems?ha^?1. Stand responses were examined for measurements recorded at 0 (1987), 5 (1992), 10 (1997), 16 (2003), and 23 (2010) years after establishment. Average diameter at breast height, quadratic mean diameter, stand basal area, and stand total volume growth increased as stem density decreased from PCT. There were significant linear differences for many of these variables between treatments and time periods (year). No significant differences were detected in tree height between treatments. In 2010, the four PCT thinning treatments did not exhibit any differences in potential sawlogs at 2.4?m (8?ft) and 3.6?m (12?ft) lengths. Significant differences were observed for 4.9?m (16?ft) sawlogs that were produced at the least dense spacing (1300?stems?ha^?1). Results from this study and recommendations from the European literature suggest that value-added timber products may be produced from more intense PCT treatments than are currently being practiced on sites dominated by yellow birch (Betula alleghaniensis Britt.).     

Canopy density in stands of Picea abies and Pinus sylvestris after different thinning methods

Altogether 82 plots (261 estimations) of Picea abies (L.) Karst, and 193 plots (360 estimations) of Pinus sylvestris (L.) stands were estimated by a vertical tube. The “crown free projection”, CFP, of stands thinned in three methods with different thinning grades was measured: unthinned, heavily and very heavily thinned, heavily thinned delayed first thinning, extra heavily thinned and thinned from the top. Basal area (m²ha^?1) density (stems ha^?1) and diameter sum (m ha^?1) were plotted against CFP. Basal area was the best practical measure of stand in this study. Generally Scots pine stands have higher CFP and the curves are steeper than in Norway spruce stands. Depending on the grade of thinning, heavily and very heavily thinned spruce stands, delayed first thinning included, have CFP values of 10–15% and stands thinned from the top, 20–40%, compared with 30–80% and 30–60% respectively in pine stands. Extra heavily thinned stands have the highest CFP, 20–80% in spruce and 50–90% in pine stands. The CFP levels after thinning are too high in pine stands for avoidance of sucker and sprout production of aspen and birch. In dense Norway spruce stands thinned from the top or heavily and very heavily thinned, the CFP values are low enough (≤30%) to diminish the production of suckers.     

Effect of wide spacing on tree growth, branch and sapwood properties of young Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] in south-western Germany

The influence of stand density on Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] is conceptually understood, but for wide spacings not well quantified, particularly in Europe. This study used 41 trees from 7 different locations in south-western Germany to compare important tree- and branch-level attributes across three different densities, namely 100, 200, and 1,200 stems ha⁻¹. In general, there were only a few tree and branch attributes that were significantly different between the 100 and 200 ha⁻¹ densities. Crown projection area and diameter of the thickest branches were the most important differences between the 100 and 200 ha⁻¹ densities. The most obvious and significant differences in this study were between 100 and 1,200 ha⁻¹ densities, where nearly every examined tree and branch attributes were statistically significant. However, relative sapwood area, the number of branches, branch angle, and the occurrence of spike knots were insensitive to stand density. Although the two lowest stand densities in this study represent rather extremely wide spacings, these results still have important implications for the development of effective thinning regimes for Douglas-fir in south-western Germany. Important management recommendations from this study include thinned stands should be maintained to at least 200 stems ha⁻¹ to maintain high log quality and stand stability. Furthermore, even at stand densities exceeding more than 1,200 trees ha⁻¹ planted trees, artificial pruning may even be necessary to produce high quality logs.     

Foliar Responses of Understorey Abies lasiocarpa to Different Degrees of Release Cutting in a Betula papyrifera and Conifer Mixed Species Stand

Foliar responses of subalpine fir [Abies lasiocarpa (Hook.) Nutt.] to thinning were studied in a 35-yr-old mixed stand of paper birch (Betula papyrifera Marsh.) and conifers. The stand regenerated naturally after a wildfire with a canopy dominated by paper birch (average height 9.8 m) and an understorey dominated by subalpine fir (average height 1.6 m). The stand was thinned to four densities of birch: 0, 600 and 1200 stems ha-¹ and control (unthinned at 2300-6400 stems ha-¹) in the autumn of 1995. The understorey conifers, mainly subalpine fir, were thinned to 1200 stems ha-¹. The study used a completely randomized split-plot design. Three sample trees were systematically selected from each treatment replicate and each tree stratum (upper, intermediate and lower understorey). One-year-old and older age class needles were collected from one south-facing branch within the fifth whorl from the tree top. Thinning of paper birch significantly (p <0.001) increased leaf area and dry weight per 100 needles for intermediate and short trees except in the 0 birch treatment. Understorey subalpine fir trees in 600 stems ha¹ birch (T3) had the largest leaf area and leaf dry weight per 100 1-yr-old needles. Specific leaf area (SLA) decreased from unthinned (T1) to 0 birch (T4). Lower understorey trees had the largest SLA. One-year-old needles had significantly higher N, P and K concentrations in all the thinning treatments. These responses are consistent with the shade tolerance of subalpine fir. The results suggest that when managing a paper birch-conifers mixed-wood forest it may be of benefit to understorey conifers to leave a birch canopy as a nursing crop.     

Influence of precommercial thinning grade on branch diameter and crown ratio in Pinus sylvestris in southern Sweden

Precommercial thinning is regarded as one of the most important measures for influencing timber quality in stands of Scots pine (Pinus sylvestris L.). The present study considers the influence of both thinning intensity and tree height at the time of thinning on diameter of the thickest branch and crown ratio. A total of 115 plots (either naturally regenerated, planted or sown, and either thinned down to 528–8000 stems?ha^?1 or untreated) included in 20 sites in southern Sweden were analysed. The average tree height after thinning varied from 1.2 to 8.3?m. An increasing number of remaining stems resulted in a reduction in branch diameter, although the reduction appeared to be only minor if the number of stems after thinning was more than 3000 stems?ha^?1. It was found that late thinning reduced the diameter of the thickest branch. The crown ratio decreased with stand height, number of stems after thinning and average height at thinning. The results were consistent for all trees and for the 500 thickest trees per hectare.     

Growth,stem quality and nutritional status of Betula pendula and Betula pubescens in pure stands and mixtures

The growth, technical quality and nutritional status of pure and mixed silver birch (Betula pendula Roth) and downy birch (Betula pubescens Ehrh.) plantations were studied 21 and 22 years after planting on afforested organic soil arable land and on upland forest soil. In mixtures, 50% of both birch species was planted. Silver birch trees grew better, but had higher mortality than downy birch trees on both sites. Mortality of both species was highest, and the difference in their growth smallest, on organic soil. In pure stands on organic soil, downy birch dominant height, diameter and mean volume were 96%, 92% and 82% of those of silver birch and on mineral soil 87%, 84% and 60%, correspondingly. On mineral soil, silver birch had a higher mean annual increment (MAI) (5.8 m³ ha^?1a^?1) than downy birch (3.9 m³ ha^?1a^?1), but on organic soil the MAI of both species was similar (3.3–3.4 m³ ha^?1 a^?1). Planting birches in mixture did not affect the growth of the trees on organic soil. On mineral soil, the mean diameter and mean volume of silver birch trees were higher in mixed than in pure plantations. The technical stem quality of both tree species was low. On mineral soils, pure silver birch is more productive than mixture, but on peat soil the higher growth of silver birch could contribute to increased productivity and downy birch would ensure sufficient survival for future timber production.     

Stand density and fertilization effects on aboveground allocation patterns and stem form of Pinus sylvestris in young stands

The increasing demand for forest biomass for energy generation could be partially met by growing denser stands and use of fertilizer. Before this is done at large scale, more knowledge of the effects of stand density and fertilization on aboveground allocation patterns and stem form is needed. Therefore, effects of pre-commercial thinning (PCT) to 3000 stems ha^?1, an unthinned dense control (C), and PCT combined with two levels of fertilization (100 kg ha^?1 of nitrogen applied either during the establishment of the field experiment (F1) or annually (F2)) were examined in 23- to 26-year-old Scots pine (Pinus sylvestris L.) stands six years after the establishment of the field experiment. In total, 114 sample trees were harvested using destructive biomass sampling. The growth allocation and stem form of trees with diameter at breast height (DBH; 1.3 m height) >5.0 cm were not affected by either the PCT or fertilization. Small trees (DBH < 5 cm) in denser, unthinned control plots had more slender stems (lower DBH/height ratios) and allocated less growth to branches and foliage than trees in PCT plots. Fertilization had little effect on the stem form and growth allocation of the smallest trees. Therefore, effects of stem density and fertilization on stem form and growth allocation to foliage were only found for small suppressed trees, and the treatments had very little influence on dominant and codominant trees.     

Sugar maple and yellow birch regeneration in response to canopy opening,liming and vegetation control in a temperate deciduous forest of Quebec

We examined how the density, growth and survival of sugar maple (Acer saccharum Marsh.) and yellow birch (Betula alleghaniensis Britton) regeneration are influenced by gap size, soil nutrient availability and understory vegetation. We used a factorial combination of (1) three gap sizes (small: <100 m²; medium: 100–300 m²; large: ∼1000 m²); (2) presence/absence of liming (92% CaCO₃ at 500 kg ha⁻¹, 1st year post-harvest); and (3) presence/absence of vegetation control (weeding twice a year; 1st to 3rd year post-harvest). We monitored height increment and survival of 1500 seedlings and saplings of both species from the 3rd to the 6th year post-harvest, and assessed density 6 years post-harvest. Both species exhibited a complex set of density, growth and survival responses across the combination of treatments. Compared to sugar maple, yellow birch had an overall lower density, greater growth, and similar survival rate; the two species attained maximum values in different gap size for density, and similar gap size for growth and survival. Liming had very little or no effect on the species. The growth of yellow birch was slightly but significantly greater when understory vegetation was controlled, particularly in medium and large gaps. These results suggest that a variety of canopy gap sizes can provide the right combination of understory conditions for regenerating these two functionally different tree species.     

Spring frost and decay fungi are implicated in suppressing aspen re-growth following partial cleaning in juvenile stands

? Aspen (Populus tremuloides Michx.) regenerates at high densities following manual cleaning.

? Ten-year-old stands located near Lac La Biche and Peace River, Alberta were manually cleaned to three densities (0, 500 or 1 500 stems ha^?1) at three times (bud set, dormancy or bud flush) to test the hypothesis that maintaining residual aspen reduces regeneration.

? At Lac La Biche up to 98% of the aspen regeneration died in the partially-cleaned plots compared to 67% at Peace River five years post-treatment. A spring frost in the second growing season at Lac La Biche is hypothesized to be the inciting factor predisposing the stump sprouts to infection by decay fungi such as Armillaria root rot, resulting in reduced density and height of the aspen regeneration at Lac La Biche relative to Peace River. Drought and ungulate herbivory provided additional stresses. The high mortality of aspen regeneration at Lac La Biche shifted the understory regeneration from aspen to balsam poplar (Populus balsamifera L.).

? These results indicate that maintaining 1 500 stems ha^?1 of residual aspen will not effectively control the re-sprouting of aspen; however, the vulnerability of aspen regeneration to spring frost and other stressors can nearly eradicate the re-growth of aspen.

     

The effect of scalping on seedling establishment after seed tree cutting of Scots pine stands in drained peatlands in northern Finland

We studied the establishment of natural seedlings after seed tree cutting and scalping in two drained Scots pine peatland stands in northern Finland (Simo and Sievi). Approximately, 50?ha^?1 of Scots pine stems were retained on both sites. During the six subsequent years, five seedling surveys were conducted. The effect of site preparation was analysed using generalized linear mixed models (GLMM). The mean density of pine seedlings increased more quickly in the scalped plots and was ca. 1.1?m^?2 after six years compared to 0.87?m^?2 in the non-scalped plots, on average. Scalping increased the number of pine seedlings 2-fold compared to that in non-treated plots in Simo, however, in Sievi non-treated plots had 30% higher pine seedling density after six years. Downy birch seedling density was 9–10-fold compared to that of non-treated plots in both sites. In terms of seedling density, seed tree cutting led to satisfactory pine regeneration after just two growing seasons in drained pine peatlands, even without scalping. Scalping significantly increased the number of pine seedlings, but it also enhanced the germination of birch seedlings strongly.     

Economic feasibility of and vegetation responses to biomass harvest in managed loblolly pine plantations

We evaluated two biomass harvest methods, (1) Whole Tree Thinning (WTT; third-row thinning), and (2) Whole Tree Thinning with Fuelchips (WTTF; third-row thinning plus remove all accessible hardwood stems >2.5 cm diameter at breast height and understory shrubs in thinned area of stand) in a 21-year-old loblolly pine (Pinus taeda L.) plantation in northwest Florida for their harvesting yields and productivities, costs, and effects on groundcover. Both WTT and WTTF produced similar quantities of roundwood (~70 Mg ha^?1) and chips (8.43–13.12 Mg ha^?1) without significantly added operational time (15.77–28.12 Mg h^?1). On-board costs of chip production (US$6.93 to 10.60 Mg^?1) and total cost of roundwood and chip production (~22.5 US$ Mg^?1) of the two harvest methods also did not differ significantly. Following either WTT or WTTF, overall percent groundcover recovered within 6 months. While shrub and grass cover were similar, forb cover in WTT increased significantly following six months of harvests. Importantly, the study suggests that biomass removal is an attractive option that could be integrated with traditional silvicultural thinning methods to manage vegetation and alleviate hazardous fuel and wildfire conditions, leading to improved forest health.     

Biomass storage in low timber productivity Mediterranean forests managed after natural post-fire regeneration in south-eastern Spain

Despite the low timber productivity of Mediterranean Pinus halepensis Mill. forests in south-eastern Spain, they are a valuable carbon sequestration source which could be extended if young stands and understories were considered. We monitored changes in biomass storage of young Aleppo pine stands naturally regenerated after wildfires, with a diachronic approach from 5 to 16 years old, including pine and understory strata, at two different quality sites (dry and semiarid climates). At each site, we set 21 permanent plots and carried out different thinning intensities at two ages, 5 and 10 years after fires. We found similar post-fire regeneration capacity at both sites in terms of total above-ground biomass storage ~6 Mg ha^?1 (3 Mg ha^?1 of the above-ground pine biomass plus 3 Mg ha^?1 of the above-ground understory biomass), but with a contrasting pine layer structure. Generally, across the diachronic study, the earlier thinning reduced biomass stocks at both sites, except for the best quality site (the dry site), where the earliest thinning (applied at post-fire year 5) enlarged carbon storage by 11 % as compared to non-thinned plots. We found root:shoot ratios of an average 0.37 for the pine layer and 0.45 for the understory layer. These results provided new information which not only furthers our understanding of carbon sequestration in low timber productivity Mediterranean forests, but will also help to develop new guidelines for sustainable management adapted to the high-risk terrestrial carbon losses of fire-prone areas.     

Predicting average growth and size distributions of Douglas-fir saplings competing with sprout clumps of tanoak or Pacific madrone

The average growth and size distributions of Douglas-fir (Pseudotsuga menziesii) saplings in three plantations were studied for 7 years (1983–1989) after thinning of associated sprout clumps of tanoak (Lithocarpus densiflorus) or Pacific madrone (Arbutus menziesii); in some cases understory vegetation (shrubs and herbs) was experimentally suppressed. Biologically based nonlinear equations explained 66, 90, and 53% of the variation in the average annual increment of Douglas-fir height, diameter-squared, and cover, respectively. Equations for annual increment of cover of hardwood and understory vegetation explained only 10 to 12% of the variation, because these parameters exhibited a high degree of variability. Model simulations demonstrated that, for the same initial levels of cover, tanoak had faster rates of cover growth than madrone and also caused greater limitations in growth of Douglas-fir. Suppression of understory vegetation increased growth of Douglas-fir only when hardwood cover was absent. Weibull functions adequately described size distributions for Douglas-fir in 92% of the individual-tree data sets. Regression functions of hardwood cover and average Douglas-fir size explained 51, 93, and 24% of the variation in the Weibull A, B, and C parameters, respectively. Model simulations with predicted Weibull parameters demonstrated that hardwood competition caused a positive skewing in size distributions for height and stem diameter of Douglas-fir.     

Biomass production and carbon sequestration of dense downy birch stands on cutaway peatlands

The establishment of biomass plantations with short-rotation forestry principles is one of the after-use options for cutaway peatlands. We studied biomass production and carbon sequestration in the above- and below-ground biomass of 25 naturally afforested, 10–30 years old downy birch (Betula pubescens Ehrh.) stands located in peat cutaway areas in Finland. Self-thinning reduced the stand density from 122,000 trees ha^?1 (stand age of 10 years) to 10,000 trees ha^?1 (25–30 years), while the leafless above-ground biomass increased from 17?Mg ha^?1 up to 79–116?Mg ha^?1. The total leafless biomass (including stumps and roots) varied from 46 to 151?Mg ha^?1. The mean annual increment (MAI) of the above-ground biomass increased up to the stand age of 15 years, after which the MAI was on the average 3.2?Mg ha^?1a^?1. With below-ground biomass, the MAI of the stands older than 15 years was 4.7?Mg ha^?1. The organic matter accumulated in the O-layer on the top of the residual peat increased linearly with the stand age, reaching 29.3?Mg ha^?1 in the oldest stand. The O-layer contributed significantly to the C sink, and the afforestation with downy birch converted most of sites into C sinks.