Multipurpose conversion management of Scots pine towards mixed oak–birch stands—A long-term simulation approach

The forest growth model 4C was used to investigate how conversion management of a Scots pine (Pinus sylvestris L.) stand towards a mixed oak–birch stand would affect stand structural development – and hence biodiversity and productivity – in the long term. For this purpose the 4C model was parameterised for natural regeneration of light demanding species and extended for management of multi-layered stands. A series of structural indicators was selected to describe key factors of forest biodiversity at the stand scale. Two consecutive aspects of Scots pine conversion were tested: (1) the choice of conversion strategy between thinning and gap creation and (2) the choice of conversion regime in terms of cutting cycle, thinning type and pine tree retention. Three simulated conversion strategies aim at the gradual removal of the pine canopy but differ in the spatial organisation of pine cuttings and hence result in different light conditions for regeneration. Only the directed gap creation strategy was able to maintain and increase birch admixture to the stand and to approach natural stand structural development. Simulation of 12 conversion regimes for the directed gap creation strategy indicated that thinning type (from above or from below), pine tree retention at final felling (50% of the standing volume or none) and cutting cycle (6, 9 or 12 years) all significantly influence stand structural development. These effects were clearest for oak development. Birch occurred in a few mixed clusters, but tended to disappear when longer cutting cycles were used. Based on a multi-criteria analysis we conclude that the optimal conversion regime – in which both stand productivity and biodiversity objectives can be combined – implies thinning from above, pine tree retention, and cutting cycles of 6 years. The conceptual validity of the model as well as the applicability of the results are discussed.     

Growth of Mixed Birch-Coniferous Stands in Relation to Pure Coniferous Stands at Similar Sites in South-eastern Norway

Mean age, mean and top heights and yield were studied in 20 mixed stands of birch ( Betula pubescens Ehrh. and B. pendula Roth) and Picea abies (L.) Karst. and nine mixed stands of birch and Pinus sylvestris L. in south-eastern Norway. Each mixed stand and the adjacent pure coniferous stand (control) were growing under the same site conditions and had not been commercially thinned. There were no significant differences in mean age at breast height or in top heights between birch and conifers in the mixed stands, while mean height was significantly higher for birch than for spruce. A growth index was calculated based on total volume and age at breast height. For the spruce sites the growth in young mixed stands (birch < 17 m) was superior to that of pure spruce, while the difference was insignificant in older stands. The growth index correlated positively with the ratio between generatively and vegetatively regenerated birch trees, and negatively with the age of the oldest species in the mixture and with site quality. For the pine sites there was no significant difference in the growth index between mixed birch-pine and pure pine stands. A mixture effect of birch on the volume yield of spruce or pine could not be demonstrated.     

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.     

Height-diameter curves with random intercepts and slopes for trees growing on drained peatlands

Models for predicting tree height were constructed for Scots pine (Pinus sylvestris), Norway spruce (Picea abies) and pubescent birch (Betula pubescens). The data consisted of two separate sets of permanent sample plots forming a representative sample of drained peatland stands in the whole country. A logarithmic height-diameter curve with one nonlinear parameter specific to each tree species was applied. It was assumed that the intercept and slope of the curve would vary randomly from stand to stand. Stand characteristics were used to predict the mean intercept and slope. A nonhomogeneous variance of the residual error was modelled as a function of tree diameter. A mixed linear model technique was applied to fit the models. The diameter of the tree of the median basal area, stand basal area, geographical location of the stand, and site quality were used as fixed independent variables in explaining the variation in the intercept. The diameter of the tree of the median basal area and the stand basal area were used in explaining the variation in the slope.     

Impacts of changing climate on the productivity of Norway spruce dominant stands with a mixture of Scots pine and birch in relation to water availability in southern and northern Finland

A process-based ecosystem model was used to assess the impacts of changing climate on net photosynthesis and total stem wood growth in relation to water availability in two unmanaged Norway spruce (Picea abies) dominant stands with a mixture of Scots pine (Pinus sylvestris) and birch (Betula sp.). The mixed stands were grown over a 100-year rotation (2000-99) in southern and northern Finland with initial species shares of 50, 25 and 25% for Norway spruce, Scots pine and birch, respectively. In addition, pure Norway spruce, Scots pine and birch stands were used as a comparison to identify whether species' response is different in mixed and pure stands. Soil type and moisture conditions (moderate drought) were expected to be the same at the beginning of the simulations irrespective of site location. Regardless of tree species, both annual net canopy photosynthesis (P(nc)) and total stem wood growth (V(s)) were, on average, lower on the southern site under the changing climate compared with the current climate (difference increasing toward the end of the rotation); the opposite was the case for the northern site. Regarding the stand water budget, evapotranspiration (E(T)) was higher under the changing climate regardless of site location. Transpiration and evaporation from the canopy affected water depletion the most. Norway spruce and birch accounted for most of the water depletion in mixed stands on both sites regardless of climatic condition. The annual soil water deficit (W(d)) was higher on the southern site under the changing climate. On the northern site, the situation was the opposite. According to our results, the growth of pure Norway spruce stands in southern Finland could be even lower than the growth of Norway spruce in mixed stands under the changing climate. The opposite was found for pure Scots pine and birch stands due to lower water depletion. This indicates that in the future the management should be properly adapted to climate change in order to sustain the productivity of mixed stands dominated by Norway spruce.     

Effect of edge stands on the development of young Pinus sylvestris stands in southern Finland

Abstract

The quantitative effect of pine-dominated edge stands in southern Finland on the density and growth of pine saplings was addressed in the study. The cumulative effect of a mature edge stand on the development of a pine sapling stand with variable cutblock sizes was also estimated. The data were acquired from 10 study sites with a Scots pine sapling stand with a current dominant height of 1–6 m and an adjoining middle-aged to mature edge stand dominated by Scots pine. Tree level models were constructed to describe and simulate the structure and development of the sapling stands. The results indicated that dividing an average regeneration area (2 ha) into smaller cutblock units (1.0–0.5 ha) appreciably increased the edge stand effect. Total stand volume growth was 6% lower on 1 ha cutblocks, and 31% lower on 0.5 ha blocks, than on 2 ha blocks.     

Aboveground biomass and nutrient accumulation dynamics in young black alder, silver birch and Scots pine plantations on reclaimed oil shale mining areas in Estonia

The growth, aboveground biomass production and nutrient accumulation in black alder (Alnus glutinosa (L.) Gaertn.), silver birch (Betula pendula Roth.) and Scots pine (Pinus sylvestris L.) plantations during 7 years after planting were investigated on reclaimed oil shale mining areas in Northeast Estonia with the aim to assess the suitability of the studied species for the reclamation of post-mining areas. The present study revealed changes in soil properties with increasing stand age. Soil pH and P concentration decreased and soil N concentration increased with stand age. The largest height and diameter of trees, aboveground biomass and current annual production occurred in the black alder stands. In the 7-year-old stands the aboveground biomass of black alder (2100 trees ha⁻¹) was 2563 kg ha⁻¹, in silver birch (1017 trees ha⁻¹) and Scots pine (3042 trees ha⁻¹) stands respective figures were 161 and 1899 kg ha⁻¹. The largest amounts of N, P, K accumulated in the aboveground part were in black alder stands. In the 7th year, the amount of N accumulated in the aboveground biomass of black alder stand was 36.1 kg ha⁻¹, the amounts of P and K were 3.0 and 8.8 kg ha⁻¹, respectively. The larger amounts of nutrients in black alder plantations are related to the larger biomass of stands. The studied species used N and P with different efficiency for the production of a unit of biomass. Black alder and silver birch needed more N and P for biomass production, and Scots pine used nutrients most efficiently. The present study showed that during 7 years after planting, the survival and productivity of black alder were high. Therefore black alder is a promising tree species for the reclamation of oil shale post-mining areas.     

Fine root morphological adaptations in Scots pine, Norway spruce and silver birch along a latitudinal gradient in boreal forests

Variability in short root morphology of the three main tree species of Europe's boreal forest (Norway spruce (Picea abies L. Karst.), Scots pine (Pinus sylvestris L.) and silver birch (Betula pendula Roth)) was investigated in four stands along a latitudinal gradient from northern Finland to southern Estonia. Silver birch and Scots pine were present in three stands and Norway spruce was present in all stands. For three fertile Norway spruce stands, fine root biomass and number of root tips per stand area or unit basal area were assessed from north to south. Principal component analysis indicated that short root morphology was significantly affected by tree species and site, which together explained 34.7% of the total variability. The range of variation in mean specific root area (SRA) was 51-74, 60-70 and 84-124 m(2) kg(-1) for Norway spruce, Scots pine and silver birch, respectively, and the corresponding ranges for specific root length were 37-47, 40-48 and 87-97 m g(-1). The range of variation in root tissue density of Norway spruce, Scots pine and silver birch was 113-182, 127-158 and 81-156 kg m(-3), respectively. Sensitivity of short root morphology to site conditions decreased in the order: Norway spruce > silver birch > Scots pine. Short root SRA increased with site fertility in all species. In Norway spruce, fine root biomass and number of root tips per m(2) decreased from north to south. The differences in morphological parameters among sites were significant but smaller than the site differences in fine root biomass and number of root tips.     

Species Interactions in the Dynamics of Even- and Uneven-Aged Boreal Forests

Many boreal tree stands are neither clearly even-aged nor clearly uneven-aged. The stands may undergo a series of stages, during which an even-aged stand is transformed into two-storied mixed stand, and finally to multistoried or uneven-aged stand structure. The species composition often changes during the succession of stand stages. This study developed models for stand dynamics that can be used in different stand structures and species compositions. The model set consists of species-specific individual-tree diameter increment and survival models, and models for ingrowth. Separate models were developed for Scots pine, Norway spruce, and hardwood species. The models were used in a growth simulator, to give illustrative examples on species influences and stand dynamics. Methods to simulate residual variation around diameter increment and ingrowth models are also presented. The results suggest that mixed stands are more productive than one-species stands. Spruce in particular benefits from an admixture of other species. Mixed species improve diameter increment, decrease mortality, and increase ingrowth. Pine is a more beneficial admixture than birch. Simulations showed that uneven-aged management of spruce forests is sustainable and productive, and even-aged conifer stands growing on medium sites can be converted into uneven-aged mixed stands by a series of strong high thinnings.     

The spreading of the S type of Heterobasidion annosum from Norway spruce stumps to the subsequent tree stand

The occurrence of Heterobasidion annosum in stumps and growing trees was investigated on 15 forest sites in southern Finland where the previous tree stand had been Norway spruce (Picea abies) infected by H. annosum, and the present stand was either Scots pine (Pinus sylvestris), lodgepole pine (Pinus contorta), Siberian larch (Larix siberica), silver birch (Betula pendula) or Norway spruce 8–53 years old. Out of 712 spruce stumps investigated of the previous tree stand, 26.3% were infected by the S group and 0.3% by the P group of H. annosum. The fungus was alive and the fruit bodies were active even in stumps cut 46 years ago. In the subsequent stand, the proportion of trees with root rot increased in spruce stands and decreased in stands of other tree species. On average, one S type genet spreading from an old spruce stump had infected 3.0 trees in the following spruce stand, 0.5 trees in lodgepole pine, 0.3 trees in Siberian larch, 0.05 trees in Scots pine and 0.03 trees in silver birch stand. Although silver birch generally was highly resistant to the S type of H. annosum, infected trees were found on one site that was planted with birch of a very northern provenance.     

Integrating the risk of wind damage into forest planning

Wind is the major abiotic risk factor in Finnish forests. Therefore, tools that help managers to assess the risk of wind damage are required. This study developed simple regression models for predicting the critical wind speed needed to uproot Scots pine, Norway spruce and birch trees at the stand edges in Finnish conditions, using the characteristics of the retained forest both downwind and upwind stands as predictors. Using information on the prevailing wind conditions in the region, the critical wind speeds were converted into probabilities of wind damage, from which a mean risk index was calculated. The mean risk index was used as an objective variable in heuristic optimisation. The results of minimizing the mean risk index were compared to other objective variables such as minimal height differences between adjacent stands. The residuals of the regression models of critical wind speeds were small, especially in Scots pine and birch. Increasing tree height of the downwind stand or area of the upwind stand (gap size) decreased the critical wind speed regardless of tree species, whereas increases in the dbh/height ratio of the downwind stand increased the critical wind speed. The shelter effect of upwind stand height was stronger in Norway spruce than in other tree species, whereas the effect of tree height of the downwind stand was larger in Scots pine and birch. Minimization of the mean risk of wind damage within forest landscapes led to smooth and non-fragmented landscape structures in terms of tree height. Incorporating even-flow constraints into the planning model led to a slight increase in the mean risk of wind damage. Of the surrogate methods for risk assessment minimization of height differences between adjacent stands performed well but not equally well as minimization of the mean risk index.     

Optimizing the joint production of timber and bilberries

Berries and mushrooms are increasingly appreciated products of Finnish forests. Therefore, there is a need to integrate them in silvicultural planning. Bilberry (Vaccinium myrtillus L.) is an economically important wild berry that is widely collected for household consumption and sale in North Karelia, Finland. In this study, bilberry yield models developed recently were included in a stand growth simulator and the joint production of timber and bilberry was optimized by maximizing soil expectation value (SEV) with 3% discounting rate, assuming that 75% of the bilberry yield is harvested. The effect of bilberry production on the optimal stand management increased with increasing bilberry price. With high bilberry prices (4–8 € kg⁻¹) it was optimal to manage the mixed stand of Scots pine, Norway spruce and birch, and the pure stand of Norway spruce so as to promote bilberry production. In the Scots pine stand, where bilberry yields are higher, bilberry production affected optimal stand management already with a price of 2 € kg⁻¹. Compared to timber production, joint production led to longer rotation lengths, higher thinning intensities, more frequent thinnings, and higher share of Scots pine in the mixed stand. The contribution of bilberries to the total SEV increased with increasing bilberry price and discounting rate. In the mixed stand and pine stand the SEV of bilberry production, calculated with 3% discounting rate, exceeded the SEV of timber production when bilberry price was 4 € kg⁻¹.With 4% discounting rate this happened already with bilberry price of 2 € kg⁻¹. It was concluded that forest management which promotes bilberry yields is the most profitable in pine stands where the potential bilberry yields are high.     

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.     

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

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

Effect of Betula pendula Admixture on Tree Growth and Branch Diameter in young Pinus sylvestris Stands in Southern Finland

The effects of a birch admixture on the height and diameter growth and maximum branch diameter in planted Scots pine stands was studied using models constructed with a data set from 13 stands of 9–16 yrs of age and 2–8 m dominant height on average sites on mineral soils in southern Finland. The density and height of the birch varied highly between and within stands. Simulated results indicated that the pines were capable of keeping up in height growth with birches that had originated from seed. Even a very high number of birches (10?000 stems ha^?1) had virtually no effect on the height growth of the pines. The number of birches had a pronounced effect on the diameter growth and the maximum branch diameter in pine. Retention of a temporary birch component in young pine stands seems a feasible way of mitigating the adverse effects of low planting densities on the external quality of pine.     

Differences in Agrobacterium infections in silver birch and Scots pine

The aim of this study was to investigate the differences in infections caused by Agrobacterium tumefaciens in a conifer, Scots pine (Pinus sylvestris), and in a non-host deciduous species, silver birch (Betula pendula). All the Agrobacterium tumefaciens strains tested caused crown-gall formation in both tree species, but the infection rates varied remarkably. In Scots pine, the development of galls was rare, and slower than in silver birch. Inoculation into the base of the stem were the most successful in gall induction. Silver-birch galls were large, often surrounding the whole stem, in contrast to Scots pine galls, which were characterized by their small size and neck-like connection with the host plant. In silver birch, no other morphological changes could be seen. In Scots pine, abnormal phenotypes with proliferating short shoots above the galls were observed during the second and third growing season. The results indicate that, of the two non-host tree species, the deciduous one, silver birch, is more susceptible to an A. tumefaciens infection than the conifer, Scots pine. The matrix for A. tumefaciens infection in silver birch differs from that in Scots pine, since the terpene compounds of Scots pine seem either to kill the agrobacteria or to suppress their growth. The differences between the species could be partly caused by their difference in sensitivity to phytohormones. These features reflect evolutionary incompatibility between A. tumefaciens and a gymnosperm.     

Risk of Snow Damage in Unmanaged and Managed Stands of Scots Pine,Norway Spruce and Birch

The aim of this study was to assess the risk of snow damage to trees in unmanaged and managed stands of Scots pine (Pinus sylvestris L.), Norway spruce [Picea abies (L.) Karst.] and birch (Betula spp.) over a rotation. The risk assessment was based on the prediction of critical snow loads in interaction with the windspeed at which trees can be expected to break or be uprooted, and on the frequency of long-term extremes of precipitation and of suitable temperature conditions for the accumulation of snow on the tree crowns. The Scots pine stands were found to be more susceptible to snow damage than the others, and an unmanaged stand of Scots pine to be more susceptible to break and uproot than a managed one. Correspondingly, an unmanaged stand of Norway spruce was more susceptible to stem breakage than a managed one, but less susceptible to uprooting. Neither unmanaged nor managed birch stands were likely to suffer any kind of snow damage. The susceptibility of unmanaged stands is caused by low tapering of the trees. Based on the frequency of long-term extremes in precipitation at the temperatures needed for snow accumulation on tree crowns, critical snow loads of 10-19, 20-29 and 30-39 kg m^-2 occurred 19.3, 3.3 and 1.3 times in a decade in southern Finland. Critical snow loads of 10-19, 20-29, 30-39 and 60-69 kg m^-2 occurred in northern Finland 17.0, 6.3, 1.7 and 0.3 times in a decade.     

Modelling annual individual-tree growth and mortality of Scots pine with data obtained at irregular measurement intervals and containing missing observations

An individual-tree growth model was developed with data from 54 permanent plots of Scots pine (Pinus sylvestris L.) located in Galicia (northwestern Spain). The study involved two model fitting approaches, one considering constant growth and mortality rates in the period between two consecutive inventories, and another considering variable growth and mortality rates in the same period. The individual-tree growth model was based on annual basal area growth, height growth and survival probability. The model included variables from groups pertaining to tree size, competition and age. Weighted regression was used as a tool for dealing with missing height observations in model fitting. Evaluation of the model via simulation of growth and mortality in the period between inventories showed that the variable growth rate approach provided slightly better results than the constant growth rate approach. The final model was consistent with expected diameter growth, height growth, dominant height growth, stand basal area growth and reduction in number of stems per hectare.     

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

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

Lessons from 72 years of monitoring a once-cut pine-hardwood stand on the Crossett Experimental Forest, Arkansas, U.S.A.

The Crossett Experimental Forest was established in 1934 to provide landowners in the Upper West Gulf Coastal Plain with reliable, science-based advice on how to manage their loblolly (Pinus taeda) and shortleaf (Pinus echinata) pine-dominated forests. A key component of this program was the establishment of an unmanaged control, currently known as the Russell R. Reynolds Research Natural Area (RRNA). Originally intended to show how the lack of regulation reduced sawtimber production compared to more intensively managed stands, the once-cut RRNA is now recognized as an increasingly scarce example of an undisturbed, mature pine-hardwood stand. This, in turn, has led to studies on forest succession, coarse woody debris, old-growth stand structure conditions, and biomass accumulation patterns. Long-term (72 years, to date) research has shown, as an example, that the RRNA has sustained >33 m² of basal area and over 240 Mg of aboveground live tree biomass per hectare for decades, values that are near the upper end of temperate forest ecosystems (outside of rainforests). These high levels are made possible by the abundance of large pines; however, pine mortality and natural successional patterns in this undisturbed stand will likely result in declining biomass in the near future. Additional work is possible regarding endangered species habitat and paleoclimate change, and there is potential for studies on invasive species effects on mature, unmanaged forests. Monitoring will continue indefinitely on the RRNA.