The potential of transforming simple structured pine plantations into mixed Mediterranean forests through natural regeneration along a rainfall gradient

The aim was to study the potential for using natural regeneration as a basis for transformation of simply structured conifer plantations into mixed Mediterranean forests. We studied the variation along a rainfall gradient, in the natural regeneration of tree species in the understory of planted 40- to 50-year-old Aleppo pine (Pinus halepensis) forests. The study was conducted within the Mediterranean zone of Israel, which extends from the semiarid northern Negev desert (rainfall ca. 300 mm yr⁻¹) in the south to the humid Upper Galilee in the north (ca 900 mm yr⁻¹). Cover and height, density, and species composition of regenerating trees were measured on south- and north-facing slopes in forest sites of comparable silvicultural history (site preparation methodology, planting density and thinning regime) distributed along the rainfall gradient. Altogether, 12 species of regenerating native broadleaved trees were found in the understory of the various forest sites. Surface cover, density and species richness increased linearly along the entire rainfall gradient, on both north- and south-facing slopes, ranging from zero in the driest forest sites up to 85% cover, 7980 trees ha⁻¹ and 4.5 species per 200 m², respectively, in the most humid ones. Species composition of regenerating trees was also related to rainfall amount, through changes in the relative importance of species along the rainfall gradient. The effect of topographic aspect on tree regeneration was inconsistent, i.e., the interaction Rainfall × Aspect was significant. Nevertheless, the general trend showed better regeneration on north-facing slopes. Most of the regenerating trees in the understory were small, i.e., less than 100 cm in height, with no clear effect of rainfall amount and topographic aspect on the relative abundance of height classes. Regeneration by Aleppo pine was highly variable among and within the different forest sites and ranged from 0 to 1565 trees ha⁻¹, with no clear relationships with rainfall amount and topographic aspect. In light of our results we propose that the future structure of forests should vary with respect to annual rainfall amount within possible silvicultural scenarios.     

Forest thinning and subsequent bark beetle-caused mortality in Northeastern California

The Warner Mountains of northeastern California on the Modoc National Forest experienced a high incidence of tree mortality (2001–2007) that was associated with drought and bark beetle (Coleoptera: Curculionidae, Scolytinae) attack. Various silvicultural thinning treatments were implemented prior to this period of tree mortality to reduce stand density and increase residual tree growth and vigor. Our study: (1) compared bark beetle-caused conifer mortality in forested areas thinned from 1985 to 1998 to similar, non-thinned areas and (2) identified site, stand and individual tree characteristics associated with conifer mortality. We sampled ponderosa pine (Pinus ponderosa var ponderosa Dougl. ex Laws.) and Jeffrey pine (Pinus jeffreyi Grev. and Balf.) trees in pre-commercially thinned and non-thinned plantations and ponderosa pine and white fir (Abies concolor var lowiana Gordon) in mixed conifer forests that were commercially thinned, salvage-thinned, and non-thinned. Clusters of five plots (1/50th ha) and four transects (20.1 × 100.6 m) were sampled to estimate stand, site and tree mortality characteristics. A total of 20 pre-commercially thinned and 13 non-thinned plantation plot clusters as well as 20 commercially thinned, 20 salvage-thinned and 20 non-thinned mixed conifer plot clusters were established. Plantation and mixed conifer data were analyzed separately. In ponderosa pine plantations, mountain pine beetle (Dendroctonus ponderosae Hopkins) (MPB) caused greater density of mortality (trees ha⁻¹ killed) in non-thinned (median 16.1 trees ha⁻¹) compared to the pre-commercially thinned (1.2 trees ha⁻¹) stands. Percent mortality (trees ha⁻¹ killed/trees ha⁻¹ host available) was less in the pre-commercially thinned (median 0.5%) compared to the non-thinned (5.0%) plantation stands. In mixed conifer areas, fir engraver beetles (Scolytus ventralis LeConte) (FEN) caused greater density of white fir mortality in non-thinned (least square mean 44.5 trees ha⁻¹) compared to the commercially thinned (23.8 trees ha⁻¹) and salvage-thinned stands (16.4 trees ha⁻¹). Percent mortality did not differ between commercially thinned (least square mean 12.6%), salvage-thinned (11.0%), and non-thinned (13.1%) mixed conifer stands. Thus, FEN-caused mortality occurred in direct proportion to the density of available white fir. In plantations, density of MPB-caused mortality was associated with treatment and tree density of all species. In mixed conifer areas, density of FEN-caused mortality had a positive association with white fir density and a curvilinear association with elevation.     

Spatial variability in stand structure and density-dependent mortality in newly established post-fire stands of a California closed-cone pine forest

Fire is an important process in California closed-cone pine forests; however spatial variability in post-fire stand dynamics of these forests is poorly understood. The 1995 Vision Fire in Point Reyes National Seashore burned over 5000 ha, initiating vigorous Pinus muricata (bishop pine) regeneration in areas that were forested prior to the fire but also serving as a catalyst for forest expansion into other locales. We examined the post-fire stand structure of P. muricata forest 14 years after fire in newly established stands where the forest has expanded across the burn landscape to determine the important factors driving variability in density, basal area, tree size, and mortality. Additionally, we estimated the self-thinning line at this point in stand development and compared the size-density relationship in this forest to the theorized (−1.605) log-log slope of Reineke’s Rule, which relates maximum stand density to average tree size. Following the fire, post-fire P. muricata density in the expanded forest ranged from 500 to 8900 live stems ha⁻¹ (median density = 1800 ha⁻¹). Post-fire tree density and basal area declined with increasing distance to individual pre-fire trees, but showed little variation with other environmental covariates. Self-thinning (density-dependent mortality) was observed in nearly all stands with post-fire density >1800 stems ha⁻¹, and post-fire P. muricata stands conformed to the size-density relationship predicted by Reineke’s Rule. This study demonstrates broad spatial variability in forest development following stand-replacing fires in California closed-cone pine forests, and highlights the importance of isolated pre-fire trees as drivers of stand establishment and development in serotinous conifers.     

Susceptibility of ponderosa pine, Pinus ponderosa (Dougl. ex Laws.), to mountain pine beetle, Dendroctonus ponderosae Hopkins,attack in uneven-aged stands in the Black Hills of South Dakota and Wyoming USA

Mountain pine beetle, Dendroctonus ponderosae Hopkins can cause extensive tree mortality in ponderosa pine, Pinus ponderosa Dougl. ex Laws., forests in the Black Hills of South Dakota and Wyoming. Most studies that have examined stand susceptibility to mountain pine beetle have been conducted in even-aged stands. Land managers increasingly practice uneven-aged management. We established 84 clusters of four plots, one where bark beetle-caused mortality was present and three uninfested plots. For all plot trees we recorded species, tree diameter, and crown position and for ponderosa pine whether they were killed or infested by mountain pine beetle. Elevation, slope, and aspect were also recorded. We used classification trees to model the likelihood of bark beetle attack based on plot and site variables. The probability of individual tree attack within the infested plots was estimated using logistic regression. Basal area of ponderosa pine in trees ≥25.4 cm in diameter at breast height (dbh) and ponderosa pine stand density index were correlated with mountain pine beetle attack. Regression trees and linear regression indicated that the amount of observed tree mortality was associated with initial ponderosa pine basal area and ponderosa pine stand density index. Infested stands had higher total and ponderosa pine basal area, total and ponderosa pine stand density index, and ponderosa pine basal area in trees ≥25.4 cm dbh. The probability of individual tree attack within infested plots was positively correlated with tree diameter with ponderosa pine stand density index modifying the relationship. A tree of a given size was more likely to be attacked in a denser stand. We conclude that stands with higher ponderosa pine basal area in trees >25.4 cm and ponderosa pine stand density index are correlated with an increased likelihood of mountain pine beetle bark beetle attack. Information form this study will help forest managers in the identification of uneven-aged stands with a higher likelihood of bark beetle attack and expected levels of tree mortality.     

Logging residue removal after thinning in Nordic boreal forests: Long-term impact on tree growth

The aim of this study was to determine the effect of whole-tree harvesting (WTH) on the growth of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) as compared to conventional stem harvesting (CH) over 10 and 20 years. Compensatory (WTH + CoF) and normal nitrogen-based (CH + F or WTH + F) fertilisation were also studied. A series of 22 field experiments were established during 1977-1987, representing a range of site types and climatic conditions in Finland, Norway and Sweden. The treatments were performed at the time of establishment and were repeated after 10-13 years at 11 experimental sites. Seven experiments were followed for 25 years.Volume increment was on average significantly lower after WTH than after CH in both 10-year periods in the spruce stands. In the pine stands thinned only once, the WTH induced growth reduction was significant during the second 10-year period, indicating a long-term response.Volume increment of pine stands was 4 and 8% and that of spruce stands 5 and 13% lower on the WTH plots than on CH during the first and the second 10-year period, respectively. For the second 10-year period the relative volume increment of the whole-tree harvested plots tended to be negatively correlated with the amount of logging residue. Accordingly, the relative volume increment decreased more, the more logging residue was harvested, stressing the importance of developing methods for leaving the nutrient-rich needles on site.If nutrient (N, P, K) losses with the removed logging residues were compensated with fertiliser (WTH + CoF), the volume increment was equal to that in the CH plots. Nitrogen (150-180 kg ha⁻¹) or N + P fertilisation increased tree growth in all experiments except in one very productive spruce stand. Pine stands fertilised only once had a normal positive growth response during the first 10-year period, on average 13 m³ ha⁻¹, followed by a negative response of 5 m³ ha⁻¹ during the second 10-year period. The fertilisation effect of WTH + F and WTH + CoF on basal area increment was both smaller and shorter than with CH + F.     

Multiscale spatial variation of the bark beetle Ips sexdentatus damage in a pine plantation forest (Landes de Gascogne,Southwestern France)

Bark beetles are notorious pests of natural and planted forests causing extensive damage. These insects depend on dead or weakened trees but can switch to healthy trees during an outbreak as mass-attacks allow the beetle to overwhelm tree defences. Climatic events like windstorms are known to favour bark beetle outbreaks because they create a large number of breeding sites, i.e., weakened trees and for this reason, windthrown timber is generally preventively harvested and removed. In December 1999, the southwest of France was struck by a devastating windstorm that felled more that 27 million m³of timber. This event offered the opportunity to study large-scale spatial pattern of trees attacked by the bark beetle Ips sexdentatus and its relationship with the spatial location of pine logs that were temporally stored in piles along stand edges during the post-storm process of fallen tree removal. The study was undertaken in a pure maritime pine forest of 1300 ha in 2001 and 2002. We developed a landscape approach based on a GIS and a complete inventory of attacked trees. During this study more than 70% of the investigated stands had at least one tree attacked by I. sexdentatus  . Spatial aggregation prevailed in stands with n≥15$n\ge 15$ attacked trees. Patches of attacked trees were identified using a kernel estimation procedure coupled with randomization tests. Attacked trees formed patches of 500–700 m² on average which displayed a clumped spatial distribution. Log piles stemming from the sanitation removals were mainly distributed along the large access roads and showed an aggregated spatial pattern as well. The spatial relationship between patches of attacked trees and log pile storage areas was analyzed by means of the Ripley’s statistic that revealed a strong association at the scale of the studied forest. Our results indicated that bark beetle attacks were facilitated in the vicinity of areas where pine logs were stored. The spatial extent of this relationship was >1000 m. Similar results were obtained in 2001 and 2002 despite differences in the number and spatial distribution of attacked trees. The presence of a strong “facilitation effect” suggests that log piles should be removed quickly in order to prevent outbreaks of bark beetles.     

Estimation of biomass and carbon stocks in plants,soil and forest floor in different tropical forests

An accurate characterization of tree carbon (TC), forest floor carbon (FFC) and soil organic carbon (SOC) in tropical forest plantations is important to estimate their contribution to global carbon stocks. This information, however, is poor and fragmented. Carbon contents were assessed in patula pine (Pinus patula) and teak (Tectona grandis) stands in tropical forest plantations of different development stages in combination with inventory assessments and soil survey information. Growth models were used to associate TOC to tree normal diameter (D) with average basal area and total tree height (H_T), with D and H_T parameters that can be used in 6–26 years old patula pine and teak in commercial tropical forests as indicators of carbon stocks. The information was obtained from individual trees in different development stages in 54 patula pine plots and 42 teak plots. The obtained TC was 99.6 Mg ha⁻¹ in patula pine and 85.7 Mg ha⁻¹ in teak forests. FFC was 2.3 and 1.2 Mg ha⁻¹, SOC in the surface layer (0–25 cm) was 92.6 and 35.8 Mg ha⁻¹, 76.1 and 19 Mg ha⁻¹ in deep layers (25–50 cm) in patula pine and teak, respectively. Carbon storage in trees was similar between patula pine and teak plantations, but patula pine had higher levels of forest floor carbon and soil organic carbon. Carbon storage in trees represents 37 and 60% of the total carbon content in patula pine and teak plantations, respectively. Even so, the remaining percentage corresponds to SOC, whereas FFC content is less than 1%. In summary, differences in carbon stocks between patula pine and teak trees were not significant, but the distribution of carbon differed between the plantation types. The low FFC does not explain the SOC stocks; however, current variability of SOC stocks could be related to variation in land use history.     

Can we use shelterwoods in Mediterranean pine forests to promote oak seedling development?

The use of shelterwoods to favour the development of natural or underplanted seedlings is common in temperate forests but rare in the pine forests of the Mediterranean area. Our aim was to assess the use of shelterwoods in Aleppo pine (Pinus halepensis) woodlands in southern France to promote the survival and growth of two co-occurring oak species: the deciduous Quercus pubescens and the evergreen Quercus ilex.Twelve Aleppo pine stands were selected and differentially thinned to create a light shelterwood (basal area = 10 m²/ha, irradiance 52%), a medium shelterwood (basal area = 19 m²/ha, irradiance 33%) and a dense shelterwood (basal area = 32 m²/ha, irradiance 13%). A total of 1248 sowing points, half composed of Q. pubescens and half of Q. ilex, were then set up in these three conditions. Seedling survival and growth were monitored for 3 years. Plant stress was assessed by measuring predawn leaf potential and photosynthetic performance through the F_v/F_m ratio. Soil moisture was also recorded at two depths during two growing seasons.Survival was high for both species in all three conditions due to three consecutive wet years. The lowest survival was recorded for Q. pubescens in the dense shelterwoods. Growth in diameter and height increased from the dense to the light shelterwoods. Shrubs developed more strongly in the light shelterwood, and increasing shrub cover enhanced height growth. Photosynthetic performance was lowest for Q. pubescens in dense shelterwoods and highest in light shelterwoods, whereas the reverse was true for Q. ilex. The lowest predawn potentials were recorded in the dense shelterwoods even though higher soil water content values were measured in this treatment during the summer drought.We show that light shelterwoods were more beneficial to growth than denser ones, indicating control mainly by light availability during the 3 years of the study. However, as lower soil moisture at 30-50 cm depth and faster understorey development were also recorded in this condition, more extended observation is needed to determine whether this benefit persists in subsequent years.     

Specific gravity responses of slash and loblolly pine following mid-rotation fertilization

Wood quality attributes were examined in six stands of slash pine (Pinus elliottii Engelm. var. elliottii) and loblolly pine (P. taeda L.) in the lower Coastal Plain of Georgia and Florida. Several plots comprised each stand, and each plot was divided so that it received three fertilizer treatments: a control treatment with herbaceous weed control at planting and brush control at mid-rotation only (control); 45 kg ha⁻¹ N + 56 kg ha⁻¹ P + herbaceous weed control at planting and 224 kg ha⁻¹ N + 45 kg ha⁻¹ P + brush control at mid-rotation (fertilizer with N at planting); and 56 kg ha⁻¹ P + herbaceous weed control at planting and 224 kg ha⁻¹ N + 45 kg ha⁻¹ P + brush control at mid-rotation (fertilizer without N at planting). Ring width, ring earlywood specific gravity (SG), ring latewood SG, whole ring SG, and ring percent latewood were measured on each of seven trees. Of these measurements, this study focused mainly on the properties related to SG. Examination of the rings showed that latewood SG was significantly lower in trees treated with fertilizers with and without N at planting in the two to three years following fertilization, but that latewood SG gradually returned to a level similar to the control. Fertilizer without N at planting may also have had a brief negative effect on earlywood SG following mid-rotation fertilization, but it was not as clear or lasting as the effect on latewood SG. Additionally, although slash and loblolly pine appear to differ in the developmental patterns of these SG properties, there were no significant differences in how these patterns interacted with treatment. This study demonstrated that fertilization treatments have similar short-term effects on the SG of slash and loblolly pines, particularly in latewood, but the trees will return to a SG pattern consistent with unfertilized trees within two or three years.     

Growth effects of thinning damage in a Corsican pine (Pinus laricio Poiret) stand in central Italy

Damage to residual stand after partial harvesting or thinning may lead to serious economic losses in terms of both timber quality at the final harvest, and tree growth reduction. Logging damages and their effect on tree growth were studied in a long term experiment on Corsican pine in central Italy. Damage frequency, agent (felling, skidding), position (root damage, stem base, between 0.3 and 1 m a.g.l., >1 m a.g.l.) and severity (light, severe) and tree growth were measured after selective thinning from below and at 10 years after the treatment. In detail, we aimed at: monitoring mechanical damages to trees at the end of thinning and after 10 years; and assessing stand stability, growing stock, ring width and basal area at 10 years after the thinning. The thinning removed about 20% of volume, 38% of trees and 26% of basal area. The basal area decreased from 56 m² ha⁻¹ to 42 m² ha⁻¹ but after 10 years it increased again to 56 m² ha⁻¹. Immediately after thinning, 13.6% of the standing trees was damaged, out of these 36.17% showed severe injuries. Damages to standing trees were mainly due to skidding. Ten years after thinning, the percentage of damaged trees was about 17%, out of which 86.67% showed severe wounds. An increase of damaged trees and of trees with severe wounds was observed suggesting that a deeper knowledge on long-term effect of logging damages is needed. This study did not highlight any effect of logging damage on tree growth. In fact, no difference in ring width was recorded between damaged and undamaged trees.     

A comparison of fire hazard mitigation alternatives in pinyon–juniper woodlands of Arizona

Concern over uncontrollable wildfire in pinyon–juniper woodlands has led public land managers in the southwestern United States to seek approaches for mitigating wildfire hazard, yet little information is available concerning effectiveness and ecological responses of alternative treatments. We established a randomized block experiment at a pinyon–juniper site in northern Arizona and tested effects of no treatment (Control), thinning only (Thin), prescribed fire only (Burn), and thinning followed by prescribed fire (Thin + Burn) on overstory structure, hazardous fuels reduction, and woody understory responses. One year after implementation, mean trees per hectare (TPH) of Utah juniper (Juniperus osteosperma) and pinyon pine (Pinus edulis), and basal area (BA) of pinyon, were significantly (P < 0.05) less in Thin and Thin + Burn treatments than Control. Additionally, pinyon TPH was less in Burn than Control. Quadratic mean diameter was significantly greater in Thin and Thin + Burn than in Control and Burn treatments. Thinning shifted diameter distributions from uneven- to even-sized. Crown fuel load (CFL) of both pinyon and juniper was significantly lower in Thin and Thin + Burn compared with Control and Burn treatments. Thin, Burn, and Thin + Burn treatments resulted in significantly greater 1-h surface fuel loads compared with the Control. The Thin treatment resulted in significantly greater mean load of the 1000-h fuel class compared with Burn and Control treatments, but did not differ from Thin + Burn. Forest floor Oi (litter) layer was not significantly affected by the treatments but Oe + Oa (duff) depth was significantly less in the Burn treatment compared with Thin and Control. Live shrubs and tree regeneration showed no differences among treatments. We concluded that thinning and thinning followed by prescribed fire were effective approaches for fuels reduction; however, resulting stand structures may be novel and outside the historical range of variability. Prescribed fire alone had minimal effects on structure and fuels reduction. Woody shrubs and tree regeneration in the understory suggested that these treatments may not have long-term deleterious ecological effects.     

Does tree species composition control soil organic carbon pools in Mediterranean mountain forests?

We compared soil organic carbon (SOC) stocks and stability under two widely distributed tree species in the Mediterranean region: Scots pine (Pinus sylvestris L.) and Pyrenean oak (Quercus pyrenaica Willd.) at their ecotone. We hypothesised that soils under Scots pine store more SOC and that tree species composition controls the amount and biochemical composition of organic matter inputs, but does not influence physico-chemical stabilization of SOC. At three locations in Central Spain, we assessed SOC stocks in the forest floor and down to 50 cm in the mineral in pure and mixed stands of Pyrenean oak and Scots pine, as well as litterfall inputs over approximately 3 years at two sites. The relative SOC stability in the topsoil (0-10 cm) was determined through size-fractionation (53 μm) into mineral-associated and particulate organic matter and through KMnO₄-reactive C and soil C:N ratio.Scots pine soils stored 95-140 Mg ha⁻¹ of C (forest floor plus 50 cm mineral soil), roughly the double than Pyrenean oak soils (40-80 Mg ha⁻¹ of C), with stocks closely correlated to litterfall rates. Differences were most pronounced in the forest floor and uppermost 10 cm of the mineral soil, but remained evident in the deeper layers. Biochemical indicators of soil organic matter suggested that biochemical recalcitrance of soil organic matter was higher under pine than under oak, contributing as well to a greater SOC storage under pine. Differences in SOC stocks between tree species were mainly due to the particulate organic matter (not associated to mineral particles). Forest conversion from Pyrenean oak to Scots pine may contribute to enhance soil C sequestration, but only in form of mineral-unprotected soil organic matter.     

A positive growth response to NaCl applications in Eucalyptus plantations established on K-deficient soils

Contrasting responses of Eucalyptus trees to K fertilizer applications have been reported on soils with low K contents. A complete randomized block experiment was set up in Brazil to test the hypothesis that large atmospheric deposits of NaCl in coastal regions might lead to a partial substitution of K by Na in Eucalyptus physiology and enhance tree growth. Treatments with application of 1.5, 3.0, 4.5 kmol K ha⁻¹ (K_1.5, K_3.0, K_4.5, respectively) as KCl, 3.0 kmol K ha⁻¹ applied as K₂SO₄, 3.0 kmol Na ha⁻¹ (Na_3.0) as NaCl commercialized for cattle feeding, and a mixture of 1.5 kmol K + 1.5 kmol Na ha⁻¹ (K_1.5 + Na_1.5) were compared to a control treatment (C) with no K and Na applications. All the plots were fertilized with large amounts of the other nutrients.     

Effect of species and spacing of fast-growing nurse trees on growth of an indigenous tree, Hopea odorata Roxb., in northeast Thailand

We tested the effects of species and spacing of nurse trees on the growth of Hopea odorata, a dipterocarp tree indigenous to Southeast Asia, in a two-storied forest management system in northeast Thailand. Eucalyptus camaldulensis, Acacia auriculiformis, and Senna siamea were planted as nurse trees in 1987 at spacings of 4 m × 8 m, 2 m × 8 m, 4 m × 4 m, and 2 m × 4 m in the Sakaerat Silvicultural Research Station of the Royal Forest Department, Thailand. Seedlings of H. odorata were planted in the nurse tree stands at a uniform spacing of 4 m × 4 m and in control plots (no nurse trees) in 1990. Stem numbers of some nurse trees were thinned by half in 1994. The stem diameter and height of all trees were measured annually until 1995 and again in 2007. The mean annual increment (MAI) in volume was estimated as 8.2–10.1 m³ ha⁻¹ year⁻¹ for E. camaldulensis and 0.9–1.2 m³ ha⁻¹ year⁻¹ for S. siamea, smaller than reported elsewhere. This suggests that the site properties were not suitable for them. The MAI of A. auriculiformis was 7.9–9.8 m³ ha⁻¹ year⁻¹, within the reported range. Survival rates of H. odorata in the S. siamea stands and the control plots decreased rapidly during the first 2 years but then stayed constant from 1992. In contrast, survival rates of H. odorata in the E. camaldulensis and A. auriculiformis stands were initially high (>70%), but then decreased after 1995. Stem diameter, tree height, and stand basal area of H. odorata were large in both the S. siamea stands and the control plots from then. The growth of H. odorata was largest in the 2 m × 8 m S. siamea stands. In contrast, it was restricted in the E. camaldulensis and A. auriculiformis stands owing to strong shading by their canopies. Thinning by 50% tended to facilitate the growth of H. odorata temporarily in the E. camaldulensis and A. auriculiformis stands. The stand basal areas of nurse trees and of H. odorata showed a trade-off. These results suggest that the growth of H. odorata was maximized in the S. siamea stands. We assume, however, that the growth of H. odorata could be improved even in the E. camaldulensis and A. auriculiformis stands by frequent or heavy thinning.     

Stand characteristics and downed woody debris accumulations associated with a mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreak in Colorado

Lodgepole pine (Pinus contorta Dougl. ex Loud.)-dominated ecosystems in north-central Colorado are undergoing rapid and drastic changes associated with overstory tree mortality from a current mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreak. To characterize stand characteristics and downed woody debris loads during the first 7 years of the outbreak, 221 plots (0.02 ha) were randomly established in infested and uninfested stands distributed across the Arapaho National Forest, Colorado. Mountain pine beetle initially attacked stands with higher lodgepole pine basal area, and lower density and basal area of Engelmann spruce (Picea engelmannii [Parry]), and subalpine fir (Abies lasiocarpa (Hook.) Nutt. var. lasiocarpa) compared to uninfested plots. Mountain pine beetle-affected stands had reduced total and lodgepole pine stocking and quadratic mean diameter. The density and basal area of live overstory lodgepole declined by 62% and 71% in infested plots, respectively. The mean diameter of live lodgepole pine was 53% lower than pre-outbreak in infested plots. Downed woody debris loads did not differ between uninfested plots and plots currently infested at the time of sampling to 3 or 4–7 years after initial infestation, but the projected downed coarse wood accumulations when 80% of the mountain pine beetle-killed trees fall indicated a fourfold increase. Depth of the litter layer and maximum height of grass and herbaceous vegetation were greater 4–7 years after initial infestation compared to uninfested plots, though understory plant percent cover was not different. Seedling and sapling density of all species combined was higher in uninfested plots but there was no difference between infested and uninfested plots for lodgepole pine alone. For trees ≥2.5 cm in diameter at breast height, the density of live lodgepole pine trees in mountain pine beetle-affected stands was higher than Engelmann spruce, subalpine fir, and aspen, (Populus tremuloides Michx.), in diameter classes comprised of trees from 2.5 cm to 30 cm in diameter, suggesting that lodgepole pine will remain as a dominant overstory tree after the bark beetle outbreak.     

Prescribed fire effects on bark beetle activity and tree mortality in southwestern ponderosa pine forests

Prescribed fire is an important tool in the management of ponderosa pine (Pinus ponderosa Dougl. ex Laws.) forests, yet effects on bark beetle (Coleoptera: Curculionidae, Scolytinae) activity and tree mortality are poorly understood in the southwestern U.S. We compared bark beetle attacks and tree mortality between paired prescribed-burned and unburned stands at each of four sites in Arizona and New Mexico for three growing seasons after burning (2004–2006). Prescribed burns increased bark beetle attacks on ponderosa pine over the first three post-fire years from 1.5 to 13% of all trees, increased successful, lethal attacks on ponderosa pine from 0.4 to 7.6%, increased mortality of ponderosa pine from all causes from 0.6 to 8.4%, and increased mortality of all tree species with diameter at breast height >13 cm from 0.6 to 9.6%. On a per year basis, prescribed burns increased ponderosa pine mortality from 0.2% per year in unburned stands to 2.8% per year in burned stands. Mortality of ponderosa pine 3 years after burning was best described by a logistic regression model with total crown damage (crown scorch + crown consumption) and bark beetle attack rating (no, partial, or mass attack by bark beetles) as independent variables. Attacks by Dendroctonus spp. did not differ significantly over bole heights, whereas attacks by Ips spp. were greater on the upper bole compared with the lower bole. Three previously published logistic regression models of tree mortality, developed from fires in 1995–1996 in northern Arizona, were moderately successful in predicting broad patterns of tree mortality in our data. The influence of bark beetle attack rating on tree mortality was stronger for our data than for data from the 1995–1996 fires. Our results highlight canopy damage from fire as a strong and consistent predictor of post-fire mortality of ponderosa pine, and bark beetle attacks and bole char rating as less consistent predictors because of temporal variability in their relationship to mortality. The small increase in tree mortality and bark beetle attacks caused by prescribed burning should be acceptable to many forest managers and the public given the resulting reduction in surface fuel and risk of severe wildfire.     

Effects of prescribed fire and season of burn on direct and indirect levels of tree mortality in Ponderosa and Jeffrey Pine Forests in California,USA

Many forests that historically experienced frequent low-intensity wildfires have undergone extensive alterations during the past century. Prescribed fire is now commonly used to restore these fire-adapted forest ecosystems. In this study, we examined the influence of prescribed burn season on levels of tree mortality attributed to prescribed fire effects (direct mortality) and bark beetles (Coleoptera: Curculionidae, Scolytinae) (indirect mortality) in ponderosa pine, Pinusponderosa Dougl. ex Laws., and Jeffrey pine, Pinusjeffreyi Grev. and Balf., forests in California, USA. A total of 816 trees (9.9% of all trees) died during this 3-yr study. Significantly higher levels of tree mortality (all sources) occurred following early and late season burns compared to the untreated control, but no significant difference was observed between burn treatments. The majority (461 trees) of tree deaths were attributed to direct mortality from prescribed burns and was strongly concentrated (391 trees) in the smallest diameter class (<20.2 cm diameter at breast height, dbh). For the largest trees (>50.7 cm dbh), significantly higher levels of tree mortality occurred on early season burns than the untreated control, most of which resulted from indirect mortality attributed to bark beetle attacks, specifically western pine beetle, Dendroctonus brevicomis LeConte, and mountain pine beetle, D. ponderosae Hopkins. Red turpentine beetle, D. valens LeConte, was the most common bark beetle species found colonizing trees, but tree mortality was not attributed to this species. A total of 355 trees (4.3% of all trees) were killed by bark beetles. Dendroctonus brevicomis (67 trees, 18.9%) and D. ponderosae (56 trees, 15.8%), were found colonizing P. ponderosa; and Jeffrey pine beetle, D. jeffreyi Hopkins, was found colonizing P. jeffreyi (seven trees, 2.0%). We also found pine engraver, Ips pini (Say) (137 trees, 38.6%), and, to a much lesser extent, Orthotomicus (=Ips) latidens (LeConte) (85 trees, 23.9%) and emarginate ips, I. emarginatus (LeConte) (3 trees, 0.8%) colonizing P. ponderosa and P. jeffreyi. Few meaningful differences in levels of indirect tree mortality attributed to bark beetle attack were observed between early and late season burns. The incidence of root and root collar pathogens (Leptographium and Sporothrix spp.), including species known to be vectored by bark beetles, was low (18% of trees sampled). The implications of these and other results to management of P. ponderosa and P. jeffreyi forests are discussed in detail.     

Protection of spruce from colonization by the bark beetle, Ips perturbatus, in Alaska

Field bioassays were conducted in south-central Alaska in a stand of Lutz spruce, Picea × lutzii, to determine whether a semiochemical interruptant (verbenone and trans-conophthorin) and/or a defense-inducing plant hormone (methyl jasmonate, MJ) could be used to protect individual standing trees from bark beetle attack. During two experiments (initiated in May 2004 and 2005, respectively), attacks by Ips perturbatus on standing trees were induced by using a three-component aggregation pheromone (ipsenol, cis-verbenol, and ipsdienol) and prevented by using the interruptant. In 2005, treatments from 2004 were repeated and additional treatments were evaluated by using MJ spray or injection with and without the interruptant. Aggregation began before 3 or 7 June, and attack density was monitored through 3 or 16 August. During both years, tree mortality caused by I. perturbatus was recorded twice (in August, and in May of the following year). In both experiments, attack density was greatest on trees baited with the three-component attractive pheromone, but was significantly reduced by addition of the semiochemical interruptant to trees baited with the attractant. There were no significant differences in attack density between attractant + interruptant-treated trees and unbaited trees. In 2004, mortality was highest among attractant-baited trees, whereas addition of the interruptant significantly reduced the level of initial (10 week post-treatment) and final (54 week post-treatment) mortality. In 2005, no significant reduction in attack density occurred on trees baited with the attractant when MJ was sprayed or injected. The highest initial (10.6 week post-treatment) and final (49.4 week post-treatment) mortality was observed among trees that had been injected with MJ and baited with the attractant. Mortality at the final assessment was significantly lower in all other treatment groups. As in 2004, addition of the interruptant to attractant-baited trees significantly reduced the level of final mortality compared to attractant-baited trees. MJ was not attractive or interruptive to I. perturbatus or associated bark beetles in a flight trapping study. However, MJ-treated trees (sprayed or injected) exuded copious amounts of resin on the bark surface. Anatomical analyses of felled trees from four treatment groups [Tween (solvent)-sprayed, MJ-sprayed, Tween-injected, and MJ-injected + attractant baited] showed that treatment with MJ increased the number and size of resin ducts produced following treatment. These analyses also revealed a reduction in radial growth in MJ-treated trees. Our results show that during both years, treatment with a simple, two-component interruptant system of verbenone and trans-conophthorin significantly reduced I. perturbatus attack density and tree mortality on attractant-baited trees and provided a full year of protection from bark beetle attack.     

Tree biomass estimation in regenerating areas of tropical dry vegetation in northeast Brazil

Allometric equations have been developed for various different vegetation types but have rarely been validated in the field and never for dry tropical forest such as caatinga. In three areas of semi-arid Brazil, with regenerating caatinga vegetation, we measured and weighed twelve hundred individuals of four tree species and used the data to validate equations previously determined in mature caatinga. They and several other equations developed for tropical vegetations overestimate the biomass (B) of trees from the regeneration areas by more than 20%, possibly because these trees have reduced crowns, with lower branch masses. We then determined new allometric equations for them, validating equations for one site against data of the others and pooling the data if they were cross-validated. The best equations were power ones, based on diameter at breast height (D), with little improvement by including height, crown area and/or wood density (Caesalpinia pyramidalis, B = 0.3129D^1.8838; Croton sonderianus, B = 0.4171D^1.5601; Mimosa ophthalmocentra, B = 0.4369D^1.8493; and Mimosa tenuiflora, B = 0.3344D^1.9648 and 0.4138D^1.7718).     

An individual-tree diameter growth model for managed uneven-aged oak-shortleaf pine stands in the Ozark Highlands of Missouri, USA

The Pioneer Forest encompasses more than 60,000 ha in the Ozark Highlands of Missouri, USA and has been managed using single-tree selection since the early 1950s. This paper quantifies the influence of tree size and competitive position, stand density, species composition, and site quality on ten-year (1992-2002) diameter increment within oak (Quercus spp.) and shortleaf pine (Pinus echinata Mill.) stands on the Pioneer Forest. An individual-tree model was developed for each species using mixed-effects regression and 290 inventory plots. Model efficiency (R²) ranged from 0.26 to 0.57 and fit was generally better for oak species. Basal area in larger trees (BAL) and tree diameter were significant predictors for all species and crown competition factor improved prediction for shortleaf pine and hickory (Carya spp.). Effect of species composition and site quality on diameter growth was not consistent across species. Models were evaluated using a subset of data not included in model fitting and the effect of single tree and standwise (1, 3, or 5 sample trees) calibration on model predictions were evaluated. Inclusion of random effects through calibration improved model prediction for all species and fit was best following single tree and 3 tree calibration.