Establishment of native tropical timber trees in monoculture and mixed-species plantations: Small-scale effects on tree performance and insect herbivory

Plantations of native timber species established on former pasture are a potential strategy to reduce the logging pressure on remnant natural forests in the tropics. Such plantations may help to mitigate or reverse the negative impacts of land degradation, and they may contribute to the long-term livelihood of livestock farmers. Planting native trees is, however, perceived as a risky activity due to limited knowledge of their performance and due to marked losses of newly established seedlings attributed to insect pests. Our study focuses on the small-scale effects of environmental heterogeneity, tree diversity and insecticide treatment on the performance of three native Central American timber species two years after establishment, and on damage inflicted by insect herbivores. Growth, survival and herbivore leaf damage were quantified for Anacardium excelsum (Anacardiaceae), Cedrela odorata (Meliaceae) and Tabebuia rosea (Bignoniaceae) planted in an experiment in Panama as (1) monocultures, (2) mixed stands, and (3) mixed stands protected by insecticides. Our study revealed that small-scale effects can have a substantial impact on the success of timber trees planted on former pasture. Growth performance and survival of the three species was strongly affected by small-scale environmental heterogeneity, which was expressed as significant differences in growth and survival among different plantation plots at the same study site. Establishment of trees in mixed stands did not have significant effects on tree survival and growth compared to pure stands, although it reduced herbivore pressure in one of the studied tree species. All tree species grew best and suffered lowest leaf damage when protected by insecticides, indicating a general influence of insect herbivory on growth of newly established trees. In contrast to growth performance, survival was not significantly affected by different management practices or herbivory. The large variability among plots in tree survival and growth, and also in the effects of management practices such as planting design and insecticide treatment, emphasizes the importance of small-scale environmental heterogeneity on tree survival and growth.     

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.     

Mixed-species plantations of Acacia mangium and Eucalyptus grandis in Brazil: 1. Growth dynamics and aboveground net primary production

Nitrogen fertilizer inputs increased sharply over the last decade in Brazilian eucalypt plantations. Due to the economic and potential environmental cost of fertilizers, mixed plantations with N-fixing species might be an attractive option to improve the long-term soil N status. A randomized block design was set up in southern Brazil, including a replacement series and an additive series design, as well as a nitrogen fertilization treatment. The development of mono-specific stands of Eucalyptus grandis (0A:100E) and Acacia mangium (100A:0E) was compared with mixed plantations in proportions of 1:1 (50A:50E), and other stands with different densities of acacia for the same density of eucalypts. The objective was to assess the effect of inter-specific interactions on the early development of the two species. Aboveground biomass was measured 6, 12, 18 and 30 months after planting, sampling 6–10 trees of each species per treatment at each age, and allometric equations were established in 0A:100E, 100A:0E, 50A:50E and 50A:100E. The height and basal area of E. grandis seedlings were enhanced by 12% and 30%, respectively by N fertilization at age 1 year. Inter-specific competition led to a stratified canopy, with suppression in acacia growth earlier for basal area than for height. The mean number of stems per acacia tree at 36 months after planting was significantly higher in pure stands (3.7), than in 50A:50E (2.7) and in the additive series (between 1.6 and 1.8). H/D ratios were highly sensitive to inter-tree competition for the two species. The suppressed acacia understorey in mixed-species stands did not influence biomass production and partitioning within eucalypts. This pattern led to biomass accumulation combining the two species in 50A:100E that was about 10% higher than in 0A:100E, from age 12 months onwards. Aboveground net primary production (ANPP) amounted to 25 Mg ha⁻¹ and 37 Mg ha⁻¹ from age 18 to 30 months in 100A:0E and 0A:100E, respectively. Acacia ANPP in 50A:100E amounted to 2 Mg ha⁻¹ over the same period, as a result of substantial inter-specific competition. An increment in biomass production in these very fast-growing eucalypt plantations was achieved introducing acacia as an understorey and not in the 50A:50E design, as observed in other studies.     

Development of bioavailable pools of base cations and P after afforestation of volcanic soils in Iceland

Few long-term studies have been conducted on changes in soil nutrients after afforestation in Iceland, a country with a young history of forest management. While fertilization was shown to improve survival of seedlings in the first years after planting on the nutrient limited soils, knowledge about the nutrients status of the soils that develop under maturing forest stands is still scarce. In a chronosequence study, the development of base cations and Olsen-phosphorus (Olsen-P) in the mineral soil was followed in six forest stands of two different tree species of increasing age (14–97 years): native birch (Betula pubescens) and introduced Siberian larch (Larix sibirica). A treeless heathland was included to present soil conditions prior to forest establishment. The sites are part of the largest forest area in Iceland, located in the east of the country. Results revealed an effect of stand age on all soil nutrients investigated except for potassium (K). Olsen-P increased in 0–10 cm depth of the mineral soil, indicating a better availability and thus improved P supply in maturing forest stands. Calcium (Ca) and magnesium (Mg) concentrations decreased with stand age in 0–10 and 10–20 cm soil depth, while sodium (Na) decreased only in the upper soil layer. Only Olsen-P and K concentrations were higher in the upper soil layer as compared to 10–20 cm depth. This indicates a higher biotic control as opposed to the geochemical control of the other base cations.     

Understory vegetation in old-growth and second-growth Tsuga canadensis forests in western Massachusetts

We compared the understory communities (herbs, shrubs, and tree seedlings and saplings) of old-growth and second-growth eastern hemlock forests (Tsuga canadensis) in western Massachusetts, USA. Second-growth hemlock forests originated following clear-cut logging in the late 1800s and were 108–136 years old at the time of sampling. Old-growth hemlock forests contained total ground cover of herbaceous and shrub species that was approximately 4 times greater than in second-growth forests (4.02 ± 0.41%/m² versus 1.06 ± 0.47%/m²) and supported greater overall species richness and diversity. In addition, seedling and sapling densities were greater in old-growth stands compared to second-growth stands and the composition of these layers was positively correlated with overstory species composition (Mantel tests, r > 0.26, P < 0.05) highlighting the strong positive neighborhood effects in these systems. Ordination of study site understory species composition identified a strong gradient in community composition from second-growth to old-growth stands. Vector overlays of environmental and forest structural variables indicated that these gradients were related to differences in overstory tree density, nitrogen availability, and coarse woody debris characteristics among hemlock stands. These relationships suggest that differences in resource availability (e.g., light, moisture, and nutrients) and microhabitat heterogeneity between old-growth and second-growth stands were likely driving these compositional patterns. Interestingly, several common forest understory plants, including Aralia nudicaulis, Dryopteris intermedia, and Viburnum alnifolium, were significant indicator species for old-growth hemlock stands, highlighting the lasting legacy of past land use on the reestablishment and growth of these common species within second-growth areas. The return of old-growth understory conditions to these second-growth areas will largely be dependent on disturbance and self-thinning mediated changes in overstory structure, resource availability, and microhabitat heterogeneity.     

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.     

Can Eucalyptus invade native forest fragments close to commercial stands?

Some Eucalyptus species are widely used as a plantation crop in tropical and subtropical regions. One reason for this is the diversity of end uses, but the main reason is the high level of wood production obtained from commercial plantings. With the advancement of biotechnology it will be possible to expand the geographical area in which eucalypts can be used as commercial plantation crops, especially in regions with current climatic restrictions. Despite the popularity of eucalypts and their increasing range, questions still exist, in both traditional planting areas and in the new regions: Can eucalypts invade areas of native vegetation, causing damage to natural ecosystems biodiversity?The objective of this study it was to assess whether eucalypts can invade native vegetation fragments in proximity to commercial stands, and what factors promote this invasive growth. Thus, three experiments were established in forest fragments located in three different regions of Brazil. Each experiment was composed of 40 plots (1 m² each one), 20 plots located at the border between the forest fragment and eucalypts plantation, and 20 plots in the interior of the forest fragments. In each experimental site, the plots were paired by two soil exposure conditions, 10 plots in natural conditions and 10 plots with soil exposure (no plant and no litter). During the rainy season, 2 g of eucalypts seeds were sown in each plot, including Eucalyptus grandis or a hybrid of E. urophylla × E. grandis, the most common commercial eucalypt species planted in the three region. At 15, 30, 45, 90, 180, 270 and 360 days after sowing, we assessed the number of seedlings of eucalypts and the number of seedlings of native species resulting from natural regeneration. Fifteen days after sowing, the greatest number of eucalypts seedlings (37 m⁻²) was observed in the plots with lower luminosity and exposed soil. Also, for native species, it was observed that exposed soil improved natural germination reaching the highest number of 163 seedlings per square meter. Site and soil exposure were the factors that have the greatest influence on seed germination of both eucalypt and native species. However, 270 days after sowing, eucalypt seedlings were not observed at any of the three experimental sites. The result shows the inability of eucalypts to adapt to condition outside of their natural range. However, native species demonstrated their strong capacity for natural regeneration in forest fragments under the same conditions where eucalypts were seeded.     

Effects of bamboo stands on seed rain and seed limitation in a rainforest

Bamboos often negatively affect tree recruitment, survival, and growth, leading to arrested tree regeneration in forested habitats. Studies so far have focused on the effects of bamboos on the performance of seedlings and saplings, but the influence of bamboos on forest dynamics may start very early in the forest regeneration process by altering seed rain patterns. We tested the prediction that the density and composition of the seed rain are altered and seed limitation is higher in stands of Guadua tagoara (B or bamboo stands), a large-sized woody bamboo native from the Brazilian Atlantic Forest, compared to forest patches without bamboos (NB or non-bamboo stands). Forty 1 m² seed traps were set in B and NB stands, and the seed rain was monitored monthly for 1 year. The seed rain was not greatly altered by the presence of bamboos: rarefied seed species richness was higher for B stands, patterns of dominance and density of seeds were similar between stands, and differences in overall composition were slight. Seed limitation, however, was greater at B stands, likely as a resulted of reduced tree density. Despite such reduced density, the presence of trees growing amidst and over the bamboos seems to play a key role in keeping the seeds falling in B stands because they serve as food sources for frugivores or simply as perches for them. The loss of such trees may lead to enhanced seed limitation, contributing ultimately to the self-perpetuating bamboo disturbance cycle.     

Stand and landscape-level factors related to bird assemblages in exotic pine plantations: Implications for forest management

Plantations cover a substantial amount of Earth's terrestrial surface and this area is expected to increase dramatically in the coming decades. Pinus plantations make up approximately 32% of the global plantation estate. They are primarily managed for wood production, but have some capacity to support native fauna. This capacity likely varies with plantation management. We examined changes in the richness and frequency of occurrence of bird species at 32 plots within a Pinus radiata plantation (a management unit comprising multiple Pinus stands) in south-eastern Australia. Plots were stratified by distance to native forest, stand age class and thinning regime. We also assessed the landscape context of each plot to determine relationships between bird assemblages and stand and landscape-level factors. Bird species richness was significantly higher at plots ≥300 m from native forest and in mature (∼20 years since planting) and old (∼27 years since planting) thinned pine stands. We were able to separate the often confounding effects of stand age and thinning regime by including old stands that had never been thinned. These stands had significantly fewer species than thinned stands suggesting thinning regime, not age is a key factor to improving the capacity of pine plantations to support native species (although an age × thinning interaction may influence this result). At the landscape level, species richness increased in pine stands when they were closer to native riparian vegetation. There were no significant differences in species composition across plots. Our study indicates the importance of stand thinning and retention of native riparian vegetation in improving the value of pine plantations for the conservation of native fauna.     

Differences in soil properties in adjacent stands of Scots pine, Norway spruce and silver birch in SW Sweden

Soil properties were compared in adjacent 50-year-old Norway spruce, Scots pine and silver birch stands growing on similar soils in south-west Sweden. The effects of tree species were most apparent in the humus layer and decreased with soil depth. At 20-30 cm depth in the mineral soil, species differences in soil properties were small and mostly not significant. Soil C, N, K, Ca, Mg, and Na content, pH, base saturation and fine root biomass all significantly differed between humus layers of different species. Since the climate, parent material, land use history and soil type were similar, the differences can be ascribed to tree species. Spruce stands had the largest amounts of carbon stored down to 30 cm depth in mineral soil (7.3 kg C m⁻²), whereas birch stands, with the lowest production, smallest amount of litterfall and lowest C:N ratio in litter and humus, had the smallest carbon pool (4.1 kg C m⁻²), with pine intermediate (4.9 kg C m⁻²). Similarly, soil nitrogen pools amounted to 349, 269, and 240 g N m⁻² for spruce, pine, and birch stands, respectively. The humus layer in birch stands was thin and mixed with mineral soil, and soil pH was highest in the birch stands. Spruce had the thickest humus layer with the lowest pH.     

Relative susceptibility of four species of African mahogany to the shoot borer Hypsipyla robusta (Lepidoptera: Pyralidae) in the moist semideciduous forest of Ghana

We examined the relative susceptibility of four mahogany species, Khaya ivorensis, Khaya anthotheca, Entandrophragma angolense, and E. utile, to Hypsipyla robusta attack. Seeds were obtained from one to three parent trees for each species. The research was conducted in the moist semideciduous forest zone in Ghana and used a randomized complete block design. Tree height and diameter and height to first branch were measured until 24 months after out-planting in the field. H. robusta damage was assessed by counting the numbers of shoots attacked, branches, and dead shoots. Khaya spp. grew better but experienced more attack than Entandrophragma spp. The relative susceptibility to H. robusta attack, from most to least, of the four species was: K. anthotheca > K. ivorensis > E. angolense > E. utile. At 24 months, the mean number of shoots attacked per tree ranged from 1.0 for an E. utile seed source to 3.6 on for a K. anthotheca seed source. At 15 months, K. anthotheca and K. ivorensis started branching at about 1.5 m, but height of clear trunk increased over time due to self-pruning. As K. anthotheca grew taller, the number of H. robusta attacks per tree declined. This suggested that selection of genotypes and species that are tolerant of H. robusta attack based on infestation of young plants may not be appropriate. Genetic factors more completely reflecting the response of different species and genotypes to H. robusta attack may manifest themselves at later growth stages.     

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).     

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.     

Stand structure interacts with previous defoliation to influence herbivore fitness

The intensity and duration of Neodiprion abietis outbreaks have recently increased in forests of North America that were precommercially thinned more than a decade earlier. We tested the hypotheses that changes in stand structure following thinning increase the fitness (i.e., survival rate × fecundity) of N. abietis by either (a) increasing foliar availability and/or quality (i.e., increased availability of primary metabolites and/or reduced foliar defenses) or (b) by reducing any negative effects on foliar quality and/or availability resulting from herbivory that occurred during the preceding season(s). Effects of thinning and previous herbivory on N. abietis and its host plant (Abies balsamea) were determined through (i) a manipulative field experiment that evaluated the effects of experimental defoliation on N. abietis in a thinned stand, (ii) a manipulative field experiment that examined the effects of thinning on N. abietis in undamaged and naturally defoliated stands, and (iii) a field survey to estimate survival of N. abietis in natural populations.Defoliation caused reductions in the availability of different-aged foliage available to larvae and in the fitness of a subsequent N. abietis generation feeding on defoliated branches, but decreases in fitness were smaller in thinned than unthinned stands. In thinned stands, defoliation was associated with increases in foliage production and foliar contents of monoterpenes and nitrogen, as well as with a decrease in foliar contents of water. Conversely, only small changes in plant growth and foliar contents of nutrients and secondary chemicals were observed in defoliated unthinned stands. This suggests that deleterious effects of defoliation on sawfly fitness were offset by an increase in the foliar content of nitrogen, a primary compound known to improve larval growth in sawflies, which supports the hypothesis that thinning moderates negative effects of previous defoliation on sawfly fitness. The present study demonstrates that forestry practices that alter stand structure by reducing tree density may increase herbivory by affecting the way trees respond to insect attack, even after crown closure, with consequences on the buildup of herbivore populations in attacked trees.     

Red pine harvest debris as a potential source of inoculum of Diplodia shoot blight pathogens

A high incidence of Diplodia shoot blight (site means ranging 85-100%) was observed on recently planted red pine (Pinus resinosa) seedlings where mature red pine stands previously had been clearcut. An investigation of the potential of harvest debris as a source of inoculum of Diplodia pathogens then was conducted. Cones, bark, needles, stems from shoots bearing needles, and stems from shoots not bearing needles (both suspended above the soil and in soil contact) were collected from harvest debris left at sites where clearcutting occurred. Conidia were quantified, and their germination rate was assessed, and Diplodia species were identified using PCR. Conidia of Diplodia species were found at all study sites and conidia counts increased from samples collected from 6 to 18 months after harvest. Germinable conidia were obtained from debris collected 6 months to 5 years after harvest. Fewer conidia were obtained from debris collected at intervals of up to 4-5 years after harvest and the percentage of germinable conidia was lower after longer intervals following harvest. More conidia were obtained and a greater percentage germinated from debris collected above the soil than from debris in soil contact. The host substrate also influenced the number of conidia and the percentage that germinated. Planting red pine seedlings next to debris infested with Diplodia pathogens could provide a persistent source of inoculum. Results should prompt further consideration by land managers and researchers of the potential forest health risks, in addition to benefits, that may be associated with harvest debris.     

Regeneration niches and functional traits of three common species in subtropical dune forest

South African coastal dune forests are young, highly disturbed subtropical communities where conventional models of forest dynamics may be challenged. We tested predictions from the gap-phase regeneration model by comparing seedlings of three common species representing contrasting regeneration strategies: Acacia karroo as a ruderal, Celtis africana as a coloniser of forest gaps, and Diospyros natalensis as a late-successional species. We grew seedlings under contrasting light and nitrogen levels in a greenhouse and in the field for 1 year to compare their growth and survival rates, allocation and photosynthetic traits. Species’ growth rates generally followed the expected order: Acacia > Celtis > Diospyros, but Acacia responded strongly to light and Celtis responded strongly to nitrogen, leading to cross-overs in growth rates. The plasticity of allocation and photosynthesis did not clearly differentiate the strategies, although it was greater in the light-demanding species. Acacia and Celtis tended to survive better in Acacia stands than in forest plots. Leaf-level light compensation points (LCPs) were similar for the three species in most conditions, but auxiliary data suggest Diospyros has a lower whole-plant LCP than Acacia. Growth rates and LCPs were lower than most of those reported for primary-forest species in the literature, suggesting an unusual degree of shade-tolerance in this habitat. We discuss reasons why variation in shade-tolerance may be less important here than in the prevailing model for forest regeneration and suggest other biotic factors that may help differentiate regeneration niches.     

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.     

Snag longevity of Douglas-fir,western hemlock,and western redcedar from permanent sample plots in coastal British Columbia

Snags are important both as structural components and as animal habitat in forests, but abundance is often low and their dynamics poorly understood in young, managed stands. Using a large data set of 19,622 snags from permanent plots in second-growth forests of coastal British Columbia, we modeled snag longevity (time from tree mortality to snag fall) for three species: Douglas-fir (Pseudotsuga menziesii), western hemlock (Tsuga heterophylla), and western redcedar (Thuja plicata). Snag longevity was strongly related to species and snag size (diameter): the median snag longevity was 16 years for Douglas-fir, 11 years for hemlock and 5 years for redcedar. Western redcedar was predominantly in the subcanopy and its rapid fall rate was related to the small size of its snags. In addition to diameter, other attributes (height to diameter ratio, height, and live crown ratio before death) contributed significantly to models for one or two of the species. However, site level variables did not contribute significantly to any of the models. Snags greater than 50 cm diameter, especially Douglas-fir snags, have the potential for persistence well beyond 20 years in these second-growth forests, and could be important for wildlife.     

Sap flow measurements reveal influence of temperature and stand structure on water use of Eucalyptus regnans forests

We examined water use by maturing Eucalyptus regnans, growing with or without an mid-storey stratum of Acacia spp. (Acacia dealbata or A. melanoxylon), for >180 consecutive days. Study sites were located in the Upper Yarra catchment area in south-eastern Australia. Depending on their contribution to stand basal area, mid-storey Acacia spp. increased total stand water use by up to 30%. Monthly water use in such stands reached more than 640,000 L ha⁻¹ (compared to 545,000 L ha⁻¹ in stands where acacias were absent) in early spring. Water use was curvilinearly related to sapwood area of Acacia spp. and logistically related to sapwood area of E. regnans. Water use of all three species showed a strong relation to daily maximum air temperatures. Distinct and simple relationships provide clear guides to the likely impacts of climate change and forest management on water yield. We compared a traditional up-scaling approach, from individual tree water use to stand water use, to a new approach that incorporates variation in temperature. Development of this approach can lead to greater precision of stand water use estimates – and in turn catchment water yield – under current climate change scenarios, which predict a rise in air temperatures of 0.6–2.5 °C by 2050 for the study area. Our temperature-dependent approach suggests that under conditions of non-limiting water availability, stand water use will rise by 2% for every 0.25 °C increase in maximum air temperatures during winter, and possibly more than that during summer.     

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.