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.     

Tree mortality in drought-stressed mixed-conifer and ponderosa pine forests, Arizona, USA

We monitored tree mortality in northern Arizona (USA) mixed-conifer and ponderosa pine (Pinus ponderosa Dougl. ex Laws) forests from 1997 to 2007, a period of severe drought in this area. Mortality was pervasive, occurring on 100 and 98% of 53 mixed-conifer and 60 ponderosa pine plots (1-ha each), respectively. Most mortality was attributable to a suite of forest insects, mediated by drought stress. The number of trees dying from 2002 to 2007 was more than 200% greater than the number dying from 1997 to 2002 in mixed-conifer forest and 74% greater in ponderosa pine forest. Extent of mortality was spatially variable in both forest types. Median cumulative mortality (the ratio of dead to live trees) increased by approximately 53 and 65% in mixed-conifer and ponderosa pine forests, respectively, from 2002 to 2007. Median mortality rates from 2002 to 2007 were approximately 2.0% year⁻¹ in mixed-conifer forest (range = 0-28.5%) and 0.4% year⁻¹ in ponderosa pine forest (range = 0-13.6%). Mortality rates generally were not strongly related to either elevation or stand density. Mortality was nonrandom with respect to tree size classes and species. Proportions of trees dying were greatest in the largest size classes, particularly in mixed-conifer forest, where mortality in the largest size class exceeded 22% from 2002 to 2007. Mortality in mixed-conifer forest was particularly pronounced for quaking aspen (85%) and white fir (28%), the least drought tolerant species present. These results provide an early glimpse of how these forest types are likely to respond to predicted climate changes in the southwestern USA. They suggest that these forests are not resilient to climate change, and that treatments to increase resilience to climate change may be appropriate. Research on causes of spatial heterogeneity in extent of mortality might suggest valuable approaches to aid in increasing resilience.     

Bark beetle-caused mortality in a drought-affected ponderosa pine landscape in Arizona,USA

Extensive ponderosa pine (Pinus ponderosa Dougl. ex Laws.) mortality associated with a widespread severe drought and increased bark beetle (Coleoptera: Curculionidae, Scolytinae) populations occurred in Arizona from 2001 to 2004. A complex of Ips beetles including: the Arizona fivespined ips, Ips lecontei Swaine, the pine engraver beetle, Ips pini (Say), Ips calligraphus (Germar), Ips latidens (LeConte), Ips knausi Swaine and Ips integer (Eichhoff) were the primary bark beetle species associated with ponderosa pine mortality. In this study we examine stand conditions and physiographic factors associated with bark beetle-caused tree mortality in ponderosa pine forests across five National Forests in Arizona. A total of 633 fixed-radius plots were established across five National Forests in Arizona: Apache-Sitgreaves, Coconino, Kaibab, Prescott, and Tonto. Prior to the bark beetle outbreak, plots with mortality had higher tree and stocking compared with plots without pine mortality. Logistic regression modeling found that probability of ponderosa pine mortality caused by bark beetles was positively correlated with tree density and inversely related with elevation and tree diameter. Given the large geographical extent of this study resulting logistic models to estimate the likelihood of bark beetle attack should have wide applicability across similar ponderosa pine forests across the Southwest. This is particularly true of a model driven by tree density and elevation constructed by combining all forests. Tree mortality resulted in significant reductions in basal area, tree density, stand density index, and mean tree diameter for ponderosa pine and for all species combined in these forests. Most of the observed pine mortality was in the 10–35 cm diameter class, which comprise much of the increase in tree density over the past century as a result of fire suppression and grazing practices. Ecological implications of tree mortality are discussed.     

Spatial point-pattern analysis for detecting density-dependent competition in a boreal chronosequence of Alberta

In the boreal forest of Alberta, fire and wind often open gaps in the canopy where late-successional species can establish and over time cause a shift in the species distribution from deciduous (e.g., trembling aspen) dominated to mixedwood, to shade-tolerant conifer (e.g., white spruce) dominated stands. This study attempted to understand the change of density-dependent competition in a boreal chronosequence and the role of tree competition in affecting stand structure and mortality. Four 1-ha stem-mapped plots were established to represent a chronosequence comprised of aspen dominated, mixedwood, and spruce dominated stands in Alberta. Second order spatial point-pattern analysis using Ripley's K(t) function showed that intraspecific competition is a prevailing force causing conspecific tree mortality and thus shaping the stand structure. The results of bivariate K(t) function analysis did not reveal sufficient evidence of interspecific competition. This suggested that competitive interaction among heterospecific trees was not strong enough to cause significant tree mortality, but the analysis of marked correlation function revealed that interspecific competition could have a negative impact on tree growth. This study highlights the importance of density-dependent competition in understanding stand dynamics of boreal forests over succession.     

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.     

Influence of fire and mountain pine beetle on the dynamics of lodgepole pine stands in British Columbia,Canada

An outbreak of the mountain pine beetle (Dendroctonus ponderosae Hopkins; MPB), currently affecting over 10.1 million hectares of lodgepole pine forests (Pinus contorta Dougl.) in British Columbia, Canada, is the largest in recorded history. We examined the dynamics of even-aged lodgepole pine forests in southern British Columbia, which were undergoing this MPB outbreak. Using dendroecology and forest measurements we reconstructed the stand processes of stand initiation, stand disturbances, tree mortality, and regeneration, and explained the current stand structure and the potential MPB impacts in selected stands. Our results indicate that stand-replacing fires initiated even-aged seral lodgepole pine stands in this region. In the absence of fire in the 20th century, multiple MPB disturbances, which each resulted in partial canopy removal, modified the simple one-layer structure of the fire-origin stands by the initiation of post-MPB disturbance regeneration layers, transforming the stands into complex, multi-aged stands. Despite high overstory mortality due to the current MPB outbreak, regeneration layers, which are likely to survive the current outbreak, will provide important ecological legacies and will contribute to mid-term timber supply.     

Tree regeneration and future stand development after bark beetle infestation and harvesting in Colorado lodgepole pine stands

In the southern Rocky Mountains, current mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreaks and associated harvesting have set millions of hectares of lodgepole pine (Pinus contorta var. latifolia Engelm. ex Wats.) forest onto new stand development trajectories. Information about immediate, post-disturbance tree regeneration will provide insight on dynamics of future stand composition and structure. We compared tree regeneration in eight paired harvested and untreated lodgepole pine stands in the Fraser Experimental Forest that experienced more than 70% overstory mortality due to beetles. New seedlings colonized both harvested and untreated stands in the first years after the beetle outbreak. In harvested areas the density of new seedlings, predominantly lodgepole pine and aspen, was four times higher than in untreated stands. Annual height growth of pine and fir advance regeneration (e.g., trees established prior to the onset of the outbreak) has doubled following overstory mortality in untreated stands. Growth simulations based on our regeneration data suggest that stand basal area and stem density will return to pre-beetle levels in untreated and harvested stands within 80-105 years. Furthermore, lodgepole pine will remain the dominant species in harvested stands over the next century, but subalpine fir will become the most abundant species in untreated areas. Owing to terrain, economic and administrative limitations, active management will treat a small fraction (<15%) of the forests killed by pine beetle. Our findings suggest that the long-term consequences of the outbreak will be most dramatic in untreated forests where the shift in tree species composition will influence timber and water production, wildfire behavior, wildlife habitat and other forest attributes.     

Secondary forest floristic composition, structure, and spatial pattern in subtropical China

Secondary evergreen broadleaved forests are precious remnants for biodiversity conservation and templates for sustainable management of natural forests in subtropical China. Floristic composition, size structure, and spatial pattern of dominant tree species have been investigated for a subtropical secondary evergreen broadleaved forest in the Huitong Yingzuijie National Forest Reserve, Hunan, China. The location of all trees greater than 4 cm in diameter at breast height (DBH) were mapped within a 0.96-ha plot in which species, DBH, and total tree height were recorded. Ripley’s K(t) function was used to analyze spatial patterns and associations. The secondary forest consisted of 74 tree species and 1,596 stems per hectare. A reverse-J shaped DBH classes distribution was observed for all stems and trees of later seral species whereas trees of earlier successional species were distributed irregularly. Significant aggregated spatial patterns were observed for all trees within the forest and for conspecific trees of each dominant species. This result, and a repulsive spatial pattern for interspecific trees of Choerospondias axillaries and Cyclobalanopsis glauca against other dominant tree species, support segregation hypothesis. Contributions of seed dispersal, topographic heterogeneity, and competition to spatial patterns of conspecific trees vary depending on tree species. Attractive spatial pattern among interspecific trees of Liquidambar fortunei, Liquidambar formosana, and Pinus massoniana reflects stochastic colonization of pioneer tree species and a facilitation relationship. Although deciduous species are long-lived and persist over long successional processes, they will eventually be replaced by late seral evergreen species within the secondary forest if no disturbance events occur.     

Recruitment limitation in forests: Lessons from an unprecedented mountain pine beetle epidemic

Since the mid-1990s the forests of central British Columbia have undergone an unprecedented Mountain Pine Beetle (Dendroctonus ponderosae Hopkins) (MPB) epidemic that has resulted in extensive mortality of canopy lodgepole pine (Pinus contorta var. latifolia Engelm.). This study investigated how seed-source availability, seedbed substrate, overstory structure, and time since MPB attack interact to affect post-MPB seedling recruitment of the dominant tree species of these forests. In addition to post-MPB recruitment, these forests may be regenerated by trees established in the understory prior to MPB disturbance. Accordingly, we examined abundance and patterns of all regeneration less than 130 cm tall. We found post-MPB recruitment was sparse. Subalpine fir (Abies laciocarpa (Hook.) Nutt.) comprised the majority of the post-MPB recruitment. It increased with local parent tree basal area and increased strongly with proximity to a major seed source. This resulted in a patchy distribution for subalpine fir post-MPB regeneration. Lodgepole pine post-MPB recruitment was limited by overstory shading. Recruitment of pine decreased as the total overstory basal area increased. Interior spruce (Picea glauca × engelmannii) post-MPB recruitment was similarly limited by total overstory basal area. Seedbed substrates were uniform and dominated by moss. Substrate type distribution did not change as time since MPB disturbance increased. The overall low post-MPB recruitment observed was likely due to a lack of disturbance to the moss-dominated forest floor. Moss is known to be a poor substrate in northern forests. The distribution of all regeneration less than 130 cm tall showed the same trends as the post-MPB regeneration. We believe the post-MPB seedling recruitment dynamics of these forests was not substantially changed from conditions prior to MPB disturbance. There was no pulse of regeneration up to 10 years post-MPB disturbance. Unless this changes, future stand structure will be dominated by the seedling bank established prior to the MPB epidemic. Subalpine fir dominated the seedling bank (68%) and post-MPB recruitment (94%). This suggests that MPB-disturbed forests are undergoing a substantial shift in landscape-level species composition.     

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.     

Stand development patterns as a consequence of the mortality in Austrocedrus chilensis forests

The forests of Austrocedrus chilensis (D. Don) Pic. Sern. et Bizarri in Argentina suffer decline and mortality throughout their natural distribution known as ‘mal del ciprés’. While several aspects of this, to date, temporally unpredictable process of overstory tree mortality have been the focus of detailed studies, there has been little research on tree growth and stand dynamics in symptomatic forests nor stand development patterns and prediction of future stand structure. We studied 12 stands in northern Patagonia (Province of Río Negro) using stand reconstruction studies to examine the changes in stand structure over time as a consequence of overstory mortality and the implications of these structural changes on the establishment and growth of the residual overstory. Dendrochronological analyses were used to reconstruct stand establishment and structure over time, and to study past diameter growth patterns. Mortality in A. chilensis forests was variable in time among stands. As expected, overstory mortality led to the successful establishment of trees in the understory in all stands; however, the response of residual overstory trees was variable. Understory establishment was low in some stands and high in others depending on the density of the overstory. While overstory trees in almost all stands released after the onset of the mortality, the pattern was not distinctive, varying in time, number and magnitude. In some stands, growth releases occurred after single or multiple tree deaths suggesting a relationship between processes, while in others this was not the case. Even when the patterns of recruitment in the understory and the overstory response varied greatly among stands, when examined together, some general patterns emerged. This study is the first to intensively explore the dynamics of A. chilensis forests affected by ‘mal del ciprés’. Additionally, this study showed that arbitrarily categorizing disturbances as discrete or chronic masks the true process of release of growing space and the resulting stand dynamics.     

Spatial distribution,advanced regeneration and stand structure of Nepalese Sal (Shorea robusta) forests subject to disturbances of different intensities

We investigated the spatial distribution, advanced regeneration and stand structure of five Shorea robusta-dominated forests in 25 1-ha plots subject to disturbances of different intensities. We aim to elucidate the relationships of advanced regeneration and spatial patterns of the tree species with degree of disturbance magnitude. Sixty-seven tree species were recorded in the forest plots; 41 species were found in the least disturbed forests, while only 10 species were found in the heavily disturbed forests. We found 5320 trees with >1.5 cm diameter at breast height, in total, and found that moderately disturbed forests contained the highest advanced regeneration (sapling)/pole densities. No significant differences were observed in stem basal area among forests. The overall stand density changed quadratically across the disturbance gradient. A strong inverse relationship was found between the overall stand density and diameter class in the least disturbed and moderately disturbed forests. Ten species showed variation in their dispersion patterns across the disturbance gradient. Most of the socio-economically important tree species analyzed showed little or no regeneration in the least and most heavily disturbed forests. Individual species showed different responses to disturbance ranging from ‘tolerant’ (Shorea robusta, Lagerstroemia parviflora and Symplocos spp.) to ‘sensitive’ (Trewia nudiflora, Adina cardifolia and Terminalia alata). We concluded that moderate disturbance intensity not only ensures high stand density, but also enhances the advanced regeneration of socio-economically important tree species and affects their dispersion patterns. Future management strategy must balance the consumptive needs of the local community with those of species conservation by allowing regulated access to the forests.     

Thinning and chipping small-diameter ponderosa pine changes understory plant communities on the Colorado Front Range

Novel fire mitigation treatments that chip harvested biomass on site are increasingly prescribed to reduce the density of small-diameter trees, yet the ecological effects of these treatments are unknown. Our objective was to investigate the impacts of mechanical thinning and whole tree chipping on Pinus ponderosa (ponderosa pine) regeneration and understory plant communities to guide applications of these new fuel disposal methods. We sampled in three treatments: (1) unthinned forests (control), (2) thinned forests with harvested biomass removed (thin-only), and (3) thinned forests with harvested biomass chipped and broadcast on site (thin + chip). Plots were located in a ponderosa pine forest of Colorado and vegetation was sampled three to five growing seasons following treatment. Forest litter depth, augmented with chipped biomass, had a negative relationship with cover of understory plant species. In situ chipping often produces a mosaic of chipped patches tens of meters in size, creating a range of woodchip depths including areas lacking woodchip cover within thinned and chipped forest stands. Thin-only and thin + chip treatments had similar overall abundance and species richness of understory plants at the stand scale, but at smaller spatial scales, areas within thin + chip treatments that were free of woodchip cover had an increased abundance of understory vegetation compared to all other areas sampled. Relative cover of non-native plant species was significantly higher in the thin-only treatments compared to control and thin + chip areas. Thin + chip treated forests also had a significantly different understory plant community composition compared to control or thin-only treatments, including an increased richness of rhizomatous plant species. We suggest that thinning followed by either chipping or removing the harvested biomass could alter understory plant species composition in ponderosa pine forests of Colorado. When considering post-treatment responses, managers should be particularly aware of both the depth and the distribution of chipped biomass that is left in forested landscapes.     

Effects of the lack of forest management on spatiotemporal dynamics of a subalpine Pinus cembra forest

Knowledge about the stand structure and dynamics of subalpine forests is crucial to preserve their multifunctionality. In the present study, we reconstructed the spatiotemporal dynamics of a subalpine Pinus cembra forest in the eastern Italian Alps in response to natural disturbances and forest management. We adopted a concurrent point pattern, dendroecological and growth dominance (GD) analysis. We mapped and measured all trees of Pinus cembra and Larix decidua in a 1?ha plot. We analyzed intra- and interspecific spatial patterns and spatial autocorrelation of tree size and age. We explored establishment dynamics and shifts in competition by analyzing growth suppression/release patterns and GD trends. Results showed a clumped, uneven-aged, multilayered structure where pine was dominant. The synergic action of ecological and human-induced factors is discussed to explain the prevalence of pine over time. Spatial pattern and autocorrelation analyses suggest a different colonization strategy of the two species, in which pine established after small-scale perturbations and experienced a stronger inter- and intra-specific competition. The interruption of tree establishment and shift in GD toward large trees resulting from the lack of forest management are the most important findings of this research. This highlights the importance of an active management to avoid the homogenization of the forest structure that is generally associated with a reduction in biodiversity and protective ability of forests.     

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.     

Temperate-like stand dynamics in relict Mediterranean-fir (Abies pinsapo, Boiss.) forests from southern Spain

Introduction and statement of the research questions

Gap dynamics have been widely studied in forests of Abies spp. from temperate and boreal regions. The local microclimate and competition for light have been identified as the main factors controlling changes in species composition and canopy structure, however little is known on dynamics of such forests in Mediterranean.

Experimental design and aims

We studied forest structure and dynamics of Abies pinsapo stands in southern Spain, in contrasting habitats and successional status. In addition past regeneration patterns and their relationship to canopy structure, disturbances and forest-use history were investigated.

Results

Stands structure attributes were within the range described for temperate conifer biomes. The age structure revealed two main cohorts comprised of a few > 100 year-old trees and abundant younger trees established in a single recruitment event after the stands were protected in the 1950s. Initial growth-rate analyses indicated that A. pinsapo regenerated mainly in small canopy gaps, while only 15% recruited from the forest understorey. For the last ten years, basal area increment was lower than 10 cm² y^?1 in 91% of studied trees and growth rate differences between trees narrowed.

Conclusion

Stand dynamics in A. pinsapo forests maintain general features of temperate fir forests. Tree establishment over time and current stand structure fit to known changes in forest use. Widespread growth decline trends might be linked to stand stagnation and global warming.     

Physiological responses of ponderosa pine in western Montana to thinning, prescribed fire and burning season

Low-elevation ponderosa pine (Pinus ponderosa Dougl. ex. Laws.) forests of the northern Rocky Mountains historically experienced frequent low-intensity fires that maintained open uneven-aged stands. A century of fire exclusion has contributed to denser ponderosa pine forests with greater competition for resources, higher tree stress and greater risk of insect attack and stand-destroying fire. Active management intended to restore a semblance of the more sustainable historic stand structure and composition includes selective thinning and prescribed fire. However, little is known about the relative effects of these management practices on the physiological performance of ponderosa pine. We measured soil water and nitrogen availability, physiological performance and wood radial increment of second growth ponderosa pine trees at the Lick Creek Experimental Site in the Bitterroot National Forest, Montana, 8 and 9 years after the application of four treatments: thinning only; thinning followed by prescribed fire in the spring; thinning followed by prescribed fire in the fall; and untreated controls. Volumetric soil water content and resin capsule ammonium did not differ among treatments. Resin capsule nitrate in the control treatment was similar to that in all other treatments, although burned treatments had lower nitrate relative to the thinned-only treatment. Trees of similar size and canopy condition in the three thinned treatments (with and without fire) displayed higher leaf-area-based photosynthetic rate, stomatal conductance and mid-morning leaf water potential in June and July, and higher wood radial increment relative to trees in control units. Specific leaf area, mass-based leaf nitrogen content and carbon isotope discrimination did not vary among treatments. Our results suggest that, despite minimal differences in soil resource availability, trees in managed units where basal area was reduced had improved gas exchange and growth compared with trees in unmanaged units. Prescribed fire (either in the spring or in the fall) in addition to thinning, had no measurable effect on the mid-term physiological performance and wood growth of second growth ponderosa pine.     

Age-related changes in ecosystem structure and function and effects on water and carbon exchange in ponderosa pine

As forests age, their structure and productivity change, yet in some cases, annual rates of water loss remain unchanged. To identify mechanisms that might explain such observations, and to determine if widely different age classes of forests differ functionally, we examined young (Y, approximately 25 years), mature (M, approximately 90 years) and old (O, approximately 250 years) ponderosa pine (Pinus ponderosa Dougl. ex P. Laws.) stands growing in a drought-prone region of central Oregon. Although the stands differed in tree leaf area index (LAIT) (Y = 0.9, M = 2.8, O = 2.1), cumulative tree transpiration measured by sap flow did not differ substantially during the growing season (100-112 mm). Yet when water was readily available, transpiration per unit leaf area of the youngest trees was about three times that of M trees and five times that of O trees. These patterns resulted from a nearly sixfold difference in leaf specific conductance (KL) between the youngest and oldest trees. At the time of maximum transpiration in the Y stand in May-June, gross carbon uptake (gross ecosystem production, GEP) was similar for Y and O stands despite an almost twofold difference in stand leaf area index (LAIS). However, the higher rate of water use by Y trees was not sustainable in the drought-prone environment, and between spring and late summer, KL of Y trees declined fivefold compared with a nearly twofold decline for M trees and a < 30% reduction in O trees. Because the Y stand contained a significant shrub understory and more exposed soil, there was no appreciable difference in mean daily latent energy fluxes between the Y stand and the older stands as measured by the eddy-covariance technique. These patterns resulted in 60 to 85% higher seasonal GEP and 55 to 65% higher water-use efficiency at the M and O stands compared with the Y stand.