Response of old-growth conifers to reduction in stand density in western Oregon forests

The positive growth response of healthy young trees to density reduction is well known. In contrast, large old trees are usually thought to be intrinsically limited in their ability to respond to increased growing space; therefore, density reduction is seldom used in stands of old-growth trees. We tested the null hypothesis that old-growth trees are incapable of responding with increased growth following density reduction. The diameter growth response of 271 Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), ponderosa pine (Pinus ponderosa Dougl. ex Laws) and sugar pine (Pinus lambertiana Dougl.) trees ranging in age from 158 to 650 years was examined 20 to 50 years after density reduction. Density reduction involved either light thinning with removal of less vigorous trees, or shelterwood treatments in which overstory trees were not removed. Ratios of basal area growth after treatment to basal area growth before treatment, and several other measures of growth, all indicated that the old trees sometimes benefited and were not harmed by density reduction. Growth increased by 10% or more for 68% of the trees in treated stands, and nearly 30% of trees increased growth by over 50%. This growth response persisted for at least 20 years. During this 20-year period, only three trees in treated stands (1.5%) exhibited a rapid decrease in growth, whereas growth decreased in 64% of trees in untreated stands. The length of time before a growth response to density reduction occurred varied from 5 to 25 years, with the greatest growth response often occurring 20 to 25 years after treatment. These results have important implications both for the basic biology of aging in woody plants as well as for silvicultural practices in forests with old-growth trees.     

The contribution of competition to tree mortality in old-growth coniferous forests

Competition is a well-documented contributor to tree mortality in temperate forests, with numerous studies documenting a relationship between tree death and the competitive environment. Models frequently rely on competition as the only non-random mechanism affecting tree mortality. However, for mature forests, competition may cease to be the primary driver of mortality.We use a large, long-term dataset to study the importance of competition in determining tree mortality in old-growth forests on the western slope of the Sierra Nevada of California, U.S.A. We make use of the comparative spatial configuration of dead and live trees, changes in tree spatial pattern through time, and field assessments of contributors to an individual tree's death to quantify competitive effects.Competition was apparently a significant contributor to tree mortality in these forests. Trees that died tended to be in more competitive environments than trees that survived, and suppression frequently appeared as a factor contributing to mortality. On the other hand, based on spatial pattern analyses, only three of 14 plots demonstrated compelling evidence that competition was dominating mortality. Most of the rest of the plots fell within the expectation for random mortality, and three fit neither the random nor the competition model. These results suggest that while competition is often playing a significant role in tree mortality processes in these forests it only infrequently governs those processes. In addition, the field assessments indicated a substantial presence of biotic mortality agents in trees that died.While competition is almost certainly important, demographics in these forests cannot accurately be characterized without a better grasp of other mortality processes. In particular, we likely need a better understanding of biotic agents and their interactions with one another and with competition.     

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.     

Carbon and nitrogen storage in agroforests, tree plantations, and pastures in western Oregon, USA

Pastures store over 90% of their carbon and nitrogen below-ground as soil organic matter. In contrast, temperate conifer forests often store large amounts of organic matter above-ground in woody plant tissue and fibrous litter. Silvopastures, which combine managed pastures with forest trees, should accrete more carbon and nitrogen than pastures or timber plantations because they may produce more total annual biomass and have both forest and grassland nutrient cycling patterns active. This hypothesis was investigated by conducting carbon and nitrogen inventories on three replications of 11 year-old Douglas-fir (Pseudotsuga menziesii)/perennial ryegrass (Lolium perenne)/subclover (Trifolium subterraneum) agroforests, ryegrasss/subclover pastures, and Douglas-fir timber plantations near Corvallis, Oregon in August 2000. Over the 11 years since planting, agroforests accumulated approximately 740 kg ha^–1 year ^–1 more C than forests and 520 kg ha^–1 year^–1 more C than pastures. Agroforests stored approximately 12% of C and 2% of N aboveground compared to 9% of C and 1% of N above ground in plantations and less than 1% of N and C aboveground in pastures. Total N content of agroforests and pastures, both of which included a nitrogen-fixing legume, were approximately 530 and 1200 kg ha^–1 greater than plantations, respectively. These results support the proposition that agroforests, such as silvopastures, may be more efficient at accreting C than plantations or pasture monocultures. However, pastures may accrete more N than agroforests or plantations. This apparent separation of response in obviously interrelated agroecosystem processes, points out the difficulty in using forest plantation or pasture research results to predict outcomes for mixed systems such as agroforests. This revised version was published online in June 2006 with corrections to the Cover Date.     

Avian foraging and nesting use of created snags in intensively-managed forests of western Oregon,USA

Snags are critical structural features for managing biological diversity in forests of the Pacific Northwest, USA. However, commercial forests in this region often contain reduced numbers of snags compared to unmanaged forests and managers require effective methods to augment snag numbers in harvest units. Therefore, we created snags by topping live trees with a mechanical harvester and studied foraging and nesting use by cavity-nesting birds of these snags in clearcuts in Douglas-fir (Pseudotsuga mensezii) forests along the west slope of the Cascade Mountain Range and east slope of the Coast Range in Oregon, USA. We used a completely randomized design to assign 6 different treatments (single or scattered distribution by 3 different densities) to 31 different harvest units. We created 1111 snags from February 1997 through April 1999 and monitored them from 2–5 years after harvest (1999–2002). Fraction of created snags with nest cavities in harvest units was generally low across all treatments and years of the study, although some individual stands demonstrated increased nesting use with snag age. While the highest fractions of snags with nest cavities were found in units with low density and scattered snags, the mean fraction of snags used for nesting did not differ among treatments. Treatment type, distribution of snags (i.e., scattered or clumped), and associated interactions did not influence fraction of snags used for foraging. However, fraction of created snags used for foraging in all harvest units increased with snag age. Fraction of snags used for foraging was greatest in the low density treatments. While this technique provides managers with a relatively economical option for creating snags, mechanical harvesters cannot be used to create tall, large snags upon which several cavity-dependent species rely and provides only a partial solution to a critical forest management issue.     

Nitrogen dynamics across silvicultural canopy gaps in young forests of western Oregon

Silvicultural canopy gaps are emerging as an alternative management tool to accelerate development of complex forest structure in young, even-aged forests of the Pacific Northwest. The effect of gap creation on available nitrogen (N) is of concern to managers because N is often a limiting nutrient in Pacific Northwest forests. We investigated patterns of N availability in the forest floor and upper mineral soil (0–10 cm) across 6–8-year-old silvicultural canopy gaps in three 50–70-year-old Douglas-fir forests spanning a wide range of soil N capital in the Coast Range and Cascade Mountains of western Oregon. We used extractable ammonium (NH₄⁺) and nitrate (NO₃⁻) pools, net N mineralization and nitrification rates, and NH₄⁺ and NO₃⁻ ion exchange resin (IER) concentrations to quantify N availability along north-south transects run through the centers of 0.4 and 0.1 ha gaps. In addition, we measured several factors known to influence N availability, including litterfall, moisture, temperature, and decomposition rates. In general, gap-forest differences in N availability were more pronounced in the mineral soil than in the forest floor. Mineral soil extractable NH₄⁺ and NO₃⁻ pools, net N mineralization and nitrification rates, and NH₄⁺ and NO₃⁻ IER concentrations were all significantly elevated in gaps relative to adjacent forest, and in several cases exhibited significantly greater spatial variability in gaps than forest. Nitrogen availability along the edges of gaps more often resembled levels in the adjacent forest than in gap centers. For the majority of response variables, there were no significant differences between northern and southern transect positions, nor between 0.4 and 0.1 ha gaps. Forest floor and mineral soil gravimetric percent moisture and temperature showed few differences along transects, while litterfall carbon (C) inputs and litterfall C:N ratios in gaps were significantly lower than in the adjacent forest. Reciprocal transfer incubations of mineral soil samples between gap and forest positions revealed that soil originating from gaps had greater net nitrification rates than forest samples, regardless of incubation environment. Overall, our results suggest that increased N availability in 6–8-year-old silvicultural gaps in young western Oregon forests may be due more to the quality and quantity of litterfall inputs resulting from early-seral species colonizing gaps than by changes in temperature and moisture conditions caused by gap creation.     

Effects of shelter tubes on hardwood tree establishment in western Oregon silvopastures

Honey locust (Gleditsia triacanthos), black locust (Robinia pseudoacacia), and honey mesquite (Prosopis glandulosa) are warm season forage trees with potential to efficiently share site resources with cool season pasture plants in Pacific Northwest silvopastures. Establishment of hardwood trees can be difficult, however, because of feeding damage from wildlife and livestock. This study compared establishment and growth of trees planted in 88 cm tall solid plastic shelter tubes to 88 cm tall plastic mesh tubes, used to protect trees from animal damage. Three replications were established in May 1995 for each of the three tree species on a hill pasture near Corvallis, Oregon. Initial tree survival during the first summer and winter following planting was higher in shelter tubes than in mesh tubes. At the end of the third growing season, 58% of black locust and 94% of honey locust trees in shelter tubes were still alive compared to only 14% of black locust and 47% of honey locust in mesh tubes. Few honey mesquite trees survived regardless of tube type used. Average three-year total height growth for black locust was increased by 650% and basal diameter growth by 380% within shelter tubes, while honey locust height growth was increased by 300% and diameter growth was increased by 150% compared to trees in mesh tubes. However, shelter tube trees tended to be taller relative to their diameter and had difficulty standing upright if tubes were removed. This revised version was published online in June 2006 with corrections to the Cover Date.     

Potential impacts of carbon taxes on carbon flux in western Oregon private forests

This study considers a carbon tax system as a policy tool for encouraging carbon sequestration through modification of management in existing forests and examines its welfare impacts and costs of the carbon sequestered. The simulated carbon tax leads to reduced harvest and increased carbon stock in the standing trees and understory biomass. Changes in the level of silvicultural investments vary by owner, depending on the nature of their initial inventory. In general investment under the tax is concentrated in regimes that establish faster growing plantations. Average rotation age increases, varying in extent across ownerships and site qualities. The carbon tax reduces both consumer and producer surpluses in regional timber markets. Producers are compensated by the carbon subsidies, except at low carbon tax levels. Not all rates of carbon tax will attract interest from private owners if participation is voluntary. Estimates of the marginal cost of sequestering carbon in western Oregon private forests are shown to be within the range of costs for projects considering afforestation alone in some eastern regions of the United States.     

Structural and compositional controls on transpiration in 40- and 450-year-old riparian forests in western Oregon, USA

Large areas of forests in the Pacific Northwest are being transformed to younger forests, yet little is known about the impact this may have on hydrological cycles. Previous work suggests that old trees use less water per unit leaf area or sapwood area than young mature trees of the same species in similar environments. Do old forests, therefore, use less water than young mature forests in similar environments, or are there other structural or compositional components in the forests that compensate for tree-level differences? We investigated the impacts of tree age, species composition and sapwood basal area on stand-level transpiration in adjacent watersheds at the H.J. Andrews Forest in the western Cascades of Oregon, one containing a young, mature (about 40 years since disturbance) conifer forest and the other an old growth (about 450 years since disturbance) forest. Sap flow measurements were used to evaluate the degree to which differences in age and species composition affect water use. Stand sapwood basal area was evaluated based on a vegetation survey for species, basal area and sapwood basal area in the riparian area of two watersheds. A simple scaling exercise derived from estimated differences in water use as a result of differences in age, species composition and stand sapwood area was used to estimate transpiration from late June through October within the entire riparian area of these watersheds. Transpiration was higher in the young stand because of greater sap flux density (sap flow per unit sapwood area) by age class and species, and greater total stand sapwood area. During the measurement period, mean daily sap flux density was 2.30 times higher in young compared with old Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) trees. Sap flux density was 1.41 times higher in young red alder (Alnus rubra Bong.) compared with young P. menziesii trees, and was 1.45 times higher in old P. menziesii compared with old western hemlock (Tsuga heterophylla (Raf.) Sarg.) trees. Overall, sapwood basal area was 21% higher in the young stand than in the old stand. In the old forest, T. heterophylla is an important co-dominant, accounting for 58% of total sapwood basal area, whereas P. menziesii is the only dominant conifer in the young stand. Angiosperms accounted for 36% of total sapwood basal area in the young stand, but only 7% in the old stand. For all factors combined, we estimated 3.27 times more water use by vegetation in the riparian area of the young stand over the measurement period. Tree age had the greatest effect on stand differences in water use, followed by differences in sapwood basal area, and finally species composition. The large differences in transpiration provide further evidence that forest management alters site water balance via elevated transpiration in vigorous young stands.     

Medicinal plants in old-growth, degraded and re-growth forests of NW Pakistan

Many old-growth forest stands in northwest Pakistan have been structurally transformed as a consequence of logging and livestock grazing, some of which are thereafter left to secondary succession. These forests represent an important resource for local inhabitants who gather and sell medicinal plants as part of their livelihood. With this in mind, the main objectives of our study were: (1) to assess differences in the structure of the tree layer and the abundance of medicinal plants among differently transformed forests, (2) to evaluate the recovery potential of medicinal plants under re-growth forests, and (3) to assess relationships between tree stand structural characteristics and the occurrence of medicinal plants.The first step of the study involved creating an approximate map covering an area of 90 km² for five forest-use types (old-growth forest, forest degraded by logging, derived woodland, agroforest and re-growth forest). Fifteen plots per forest-use type were randomly allocated at altitudes ranging from 2200 m to 2400 m asl, within which the abundance of 10 locally important medicinal herb species was assessed.The study stands differed greatly in tree basal area, which was highest in old-growth forest (48 m² ha⁻¹), lowest in agroforest areas (6 m² ha⁻¹) and intermediate in re-growth forest (20 m² ha⁻¹). All ten medicinal plant species were encountered in old-growth and in re-growth forests, but only five of these species also occurred on agroforest plots. The mean coverage of study medicinal plants was highest in old-growth forest (7%), low in forest degraded by logging, derived woodland and agroforest (0.3-2%), and intermediate in re-growth forest (4%). The Jaccard abundance based similarity index indicates a considerable similarity (0.6) between re-growth and old growth forest for both trees and medicinal plants. The overall abundance of medicinal plants increased with increasing tree basal area and canopy cover. The abundance of some particular species decreased; however, the most sought-after medicinal species Bergenia ciliata, Valeriana jatamansi and Viola cancescens increased with tree basal area within specific forest-use type and also across forest-use types. In conclusion, our data suggest that anthropogenic forest degradation leads to a reduction in the abundance of economically viable medicinal plants for the study region. It is further indicated that this can be reversed if degraded forests are allowed to regenerate.     

Growth reduction in harvest-age,coniferous forests with residual trees in the western central Cascade Range of Oregon

To evaluate the relationship of overstory residual trees to the growth of unmanaged young-to-mature understory Douglas fir (Pseudotsuga menziesii (Mirb.) Franco) and western hemlock (Tsuga heterophylla (Raf.) Sarg.), the basal area and volume of 14 paired plots with and without residual trees were examined in the Willamette National Forest, Oregon. Residual trees were large survivors of the fires that initiated the understory between 55 and 121 yr ago. Understory stands were naturally regenerated and not managed in any way. High residual tree and understory densities were negatively associated with understory volume. The relation of density of residual trees to total understory and Douglas-fir basal areas and volumes was best described by a negative logarithmic function. The rate of decrease in total understory and Douglas-fir basal areas and volumes per individual residual tree became smaller with increasing residual-tree density. Predicted total understory volume reduction was 23% with five residual trees/ha and 47% with 50 residual trees/ha, averaging 4.6% and 0.9% per residual tree, respectively. After including the estimated volume growth of residual trees since initiation of the understory, stand volume was still 19% lower with five residual trees/ha and 41% lower with 50 residual trees/ha than in stands with no residual trees, averaging a reduction of 38% and 0.8% per residual tree, respectively. In mixed stands of Douglas fir and western hemlock, predicted Douglas-fir basal area and volume declined more rapidly than did total understory basal area and volume when residual-tree densities exceeded about 15 trees/ha. This difference was probably due to the relative shade-intolerance of Douglas fir. Predicted Douglas-fir volume reduction was 13% with five residual trees/ha and 75% with 50 residual trees/ha, averaging 2.6% and 1.5% per residual tree, respectively. The southern aspects had more than 150% the total understory basal area and volume and more than 200% the Douglas-fir volume and basal area of the northern aspects. Lower density and basal area of understory trees, particularly of dominant and codominant Douglas fir, were associated with increasing residual-tree densities. Given the same diameter at breast height (DBH), heights of Douglas fir were not related to residual trees. Regardless of understory age, understory volume was greatest in stands with the lowest understory densities. These results suggest that timber production in unthinned green-tree retention units may be reduced and may depend on the density of leave-trees. Thinning of understory trees is recommended to reduce growth loss from intraspecific competition.     

Impacts of Swiss needle cast on overstory Douglas-fir forests of the western Oregon Coast Range

Tree-ring analysis was applied to assess the impacts of the fungal disease Swiss needle cast on the radial growth of mature Douglas-fir (Pseudotsuga menziesii) forests in the western Oregon Coast Range. Although considered endemic to the Pacific Northwest, Swiss needle cast has significantly lowered productivity in Douglas-fir forests only in the past 20–30 years. To date, studies on Swiss needle cast impacts have almost exclusively involved young (<30 years) plantation trees. To better describe the history of Swiss needle cast and its impacts on older (>80 years) trees, we extracted tree cores from dominant and codominant Douglas-fir and western hemlock (Tsuga heterophylla) in three even-aged stands in western Oregon. In the least affected stand growth rates of both species did not significantly differ, while at the most severely diseased site Douglas-fir radial growth was reduced by as much as 85%. Growth reductions likely associated with Swiss needle cast were dated to as early as 1950, though the most severe impacts occurred after 1984. An index of Swiss needle cast severity significantly (p < 0.01) related to instrumental records of air temperatures such that warm conditions from March through August were associated with reduced radial growth at the most severely affected site. Overall, this study demonstrates that even mature forests of natural origin are susceptible to severe growth reductions by Swiss needle cast, that warmer spring and summer temperatures are associated with Swiss needle cast impacts, and that the disease appears to be increasing in severity.     

Carbon storage in old-growth and second growth fire-dependent western larch (Larix occidentalis Nutt.) forests of the Inland Northwest,USA

There is limited understanding of the carbon (C) storage capacity and overall ecological structure of old-growth forests of western Montana, leaving little ability to evaluate the role of old-growth forests in regional C cycles and ecosystem level C storage capacity. To investigate the difference in C storage between equivalent stands of contrasting age classes and management histories, we surveyed paired old-growth and second growth western larch (Larix occidentalis Nutt)–Douglas-fir (Pseudostuga menziesii var. glauca) stands in northwestern Montana. The specific objectives of this study were to: (1) estimate ecosystem C of old-growth and second growth western larch stands; (2) compare C storage of paired old-growth–second growth stands; and (3) assess differences in ecosystem function and structure between the two age classes, specifically measuring C associated with mineral soil, forest floor, coarse woody debris (CWD), understory, and overstory, as well as overall structure of vegetation. Stands were surveyed using a modified USFS FIA protocol, focusing on ecological components related to soil, forest floor, and overstory C. All downed wood, forest floor, and soil samples were then analyzed for total C and total nitrogen (N). Total ecosystem C in the old-growth forests was significantly greater than that in second growth forests, storing over 3 times the C. Average total mineral soil C was not significantly different in second growth stands compared to old-growth stands; however, total C of the forest floor was significantly greater in old-growth (23.8 Mg ha⁻¹) compared to second growth stands (4.9 Mg ha⁻¹). Overstory and coarse root biomass held the greatest differences in ecosystem C between the two stand types (old-growth, second growth), with nearly 7 times more C in old-growth trees than trees found on second growth stands (144.2 Mg ha⁻¹ vs. 23.8 Mg ha⁻¹). Total CWD on old-growth stands accounted for almost 19 times more C than CWD found in second growth stands. Soil bulk density was also significantly higher on second growth stands some 30+ years after harvest, demonstrating long-term impacts of harvest on soil. Results suggest ecological components specific to old-growth western larch forests, such as coarse root biomass, large amounts of CWD, and a thick forest floor layer are important contributors to long-term C storage within these ecosystems. This, combined with functional implications of contrasts in C distribution and dynamics, suggest that old-growth western larch/Douglas-fir forests are both functionally and structurally distinctive from their second growth counterparts.     

Promoting old-growth characteristics and long-term wood production in Douglas-fir forests

Trade-offs among wood production, wood quality and ecological characteristics in the management of harvested forest stands are explored through model simulation of various silvicultural regimes. Long-term production of merchantable wood, production of various types of high-quality wood, and the level of certain quantitative ecological indicators are projected for coniferous forests of Pacific Northwestern USA. The set of ecological indicators used is based on the species composition and physical structure of old, unlogged forest stands. Simulations are performed with an ecological model of forest stand dynamics that tracks the fate of live and dead trees. Short rotations (<50 years) produce the least amount of high-quality wood over the multi-century simulation period. They also fail to generate ecological attributes resembling those of old forest stands. Production of high-quality wood is moderate to high under all rotations of 80 years or more; however, most ecological indicators require longer rotations unless alternatives to clearcutting are applied. Alternatives examined include retention of 15% cover of live tree canopy at each harvest in combination with artificial thinning between harvests. Thinning from below can expedite the development of large live and dead trees, and canopy height diversity without greatly diminishing wood quantity or quality. Proportional thinning retains understory stems, thereby expediting the recruitment of shade-tolerant trees. A possible drawback to thinning, particularly proportional thinning, is the diminished production of clean-bole wood at rotations of 150 and 260 years. It is concluded that most wood quantity, wood quality and ecological objectives can be met with long rotations (ca. 260 years). Certain objectives can be met with shorter rotations (80–150 years) when treatments of thinning and canopy tree retention are applied.     

Decay progression and classification in two old-growth forests in Atlantic Canada

This paper investigates the relationship between visually apparent stage of decay of coarse woody debris (CWD) and time since death of decaying balsam fir (Abies balsamea L.) and black spruce (Picea mariana [P. Mill]) in old-growth forests in western Newfoundland and in the Cape Breton Highlands (CBH) of Nova Scotia. These sites are two of the least disturbed old-growth forest locations remaining in Atlantic Canada. In Newfoundland, a total of 42 detrital samples were collected from downed logs and standing snags, of which 36 had their mortality dates determined. In the CBH, 50 detrital samples were collected, of which death dates for 44 samples were obtained. For both sites, samples represented all visually discernable classes of decay. In Newfoundland, these visual decay classes were separated by approximately 17 years for a minimum decay time of 85 years. In CBH, a faster rate of decomposition was apparent, with 12-year classes and a minimum decay time of logs of 60 years. Evidence points toward a climate-driven decay regime in both locations, with the longer time frame evident in Newfoundland thought to result from lower temperatures and fewer snow-free days than in CBH.     

Evaluating tree competition indices as predictors of basal area increment in western Montana forests

Fire hazard reduction treatments are commonly applied to mixed-species coniferous forests in western Montana, USA, to modify fuels structures and alter the competitive environments of individual trees. An improved understanding of how competition can be measured and how it conditions individual tree growth is needed for projecting the development of these forests, with and without treatment. Numerous studies have evaluated how competition affects tree growth and many indices have been developed to quantify the competition an individual tree experiences. These studies suggest that no single competition index or a single class of indices is universally superior; indices perform differently according to forest type and forest conditions. We chose several widely used distance-independent and distance-dependent competition indices, and also derived anisotropic distance-dependent indices from estimates of light interception by tree crowns. We evaluated the effectiveness of these competition measures for predicting basal area increment (BAI) of Pinus ponderosa, Pseudotsuga menziesii, and Larix occidentalis in western Montana. The best distance-dependent competition indices explained a larger proportion of growth variation than the best distance-independent indices (64% vs. 56%). This result indicates that competition is an important growth determinant in these forests and that competition varies locally, with variable tree densities and relatively complex stand structures creating heterogeneous neighborhood conditions. Competition indices derived from light interception models were only weakly correlated with other indices and performed poorly in terms of predicting tree growth. This result accords with previous observations that competition for light is not the primarily growth limitation for trees in the semi-arid conditions of western Montana. More sophisticated light availability models could be used to better assess variability in light interception and its marginal contribution to predictive accuracy of radial tree growth. Diameter and distance-dependent BAI models were developed for growth prediction at the species level and for all species combined.     

Tree diameter, height and stocking in even-aged forests

? Empirical observations suggest that in pure even-aged forests, the mean diameter of forest trees (D, diameter at breast height, 1.3 m above ground) tends to remain a constant proportion of stand height (H, average height of the largest trees in a stand) divided by the logarithm of stand density (N, number of trees per hectare): D = β(H ? 1.3)/ ln(N).

? Thinning causes a relatively small and temporary change in the slope β, the magnitude and duration of which depends on the nature of the thinning.

? This relationship may provide a robust predictor of growth in situations where scarce data and resources preclude more sophisticated modelling approaches.

     

Population structure and regeneration patterns of tree species in climate-sensitive subalpine forests of Indian western Himalaya

The population structure of tree species has been explored in order to elucidate regeneration potential of the subalpine forests of Indian western Himalaya. For this study, the subalpine forest area was divided into three strata, i.e., lower altitude （〈3000 m）; mid-altitude （3000-3200 m）; and high altitude （〉3200m）. Considering the major compositional attributes, an increase in altitude came with a significant decline in tree density and the total basal area for all the sites. However, no such clear trends were observed for recruits （i.e., seedlings and saplings）. Seedling density did not exhibit uniform patterns for sites and altitude strata. In general, overall seedling density was greater at the Pindari site compared to the Lata and Tungnath sites. By comparison, significant variation in seedling density along the altitude strata was recorded for the Tungnath and Pindari sites only. Likewise, sapling density patterns varied across the sites and altitude strata, and significant variation in sapling density along the altitude strata was recorded only for the Lata site. At the Pin- daft site, the continuous increase in sapling density along with increasing altitude was revealing. The Pindari forests of exhibited expanding population structure. In contrast, greater accumulation of individuals in the sapling class and sharp decline toward both higher tree classes and lower seedling classes was generally apparent for the Lata and Tungnath sites. This indicates that the replacement in tree size classes from sapling stage is not proportional and the population may decline in the long-term. Considerable variation in patterns of forest and dominant species popula- tion structure were evident across altitude strata. But in all cases irrespective of sites, we found growth at the high-altitude stratum, in the form of entire forests or dominant species. This trend deserves further investigation to explore its relevance under changing climate scenarios.