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

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

Ecosystem production in Douglas-fir plantations: Interaction of red alder and site fertility

Pure stands of Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco) and mixed stands of Douglas-fir and naturally established red alder (Alnus rubra Bong.) were examined on two different sites for soil properties, tree growth and nutrition, and aboveground ecosystem biomass and net primary production. One site (Mt. Benson, Nanaimo, B.C.) was nitrogen (N)-deficient and had a low site index (expected Douglas-fir height of 24 m at 50 years). The other site (Skykomish, western Washington) was N-rich and had a site index of 45 m at 50 years. Soil N accretion on the red alder units was estimated at 65 (Mt. Benson) and 42 (Skykomish) kg ha^?1 year^?1 for 23 years to a soil depth of 50 cm. At the current stage of plantation development, presence of red alder at the infertile Mt. Benson site increased average Douglas-fir diameter but did not affect its basal area and basal area growth rate; including alder stem biomass increased total stand basal area and basal area growth 2.5 fold. Presence of red alder at the fertile Skykomish site decreased average diameter, basal area, and basal area growth of Douglas-fir; including alder biomass left total stand basal area and basal area growth unchanged. Douglas-fir foliar N concentrations on Mt. Benson increased from 0.93 without alder to 1.41% on the red alder unit but were 1.55% for both units at Skykomish. Although alder did not affect Douglas-fir aboveground biomass and net primary production on Mt. Benson, total ecosystem biomass doubled and production tripled when alder biomass was included. Conversely, at Skykomish, Douglas-fir biomass and production decreased, and total ecosystem values were essentially unchanged. Mixing red alder and Douglas-fir seems to have great potential for increasing Douglas-fir growth and ecosystem production on infertile, N-deficient sites but probably has limited value on fertile, N-rich sites.     

Hydraulic architecture and photosynthetic capacity as constraints on release from suppression in Douglas-fir and western hemlock

We compared hydraulic architecture, photosynthesis and growth in Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), a shade-intolerant species, and western hemlock (Tsuga heterophylla (Raf.) Sarg.), a shade-tolerant species, to study the temporal pattern of release from suppressive shade. In particular, we sought to determine whether hydraulic architecture or photosynthetic capacity is most important in constraining release. The study was conducted at two sites with mixed stands of 10- to 20-year-old Douglas-fir and western hemlock. At one site, the stand had been thinned allowing release of the understory trees, whereas at the other site, the stand remained unthinned. Douglas-fir had lower height growth (from 1998-2003) and lower relative height growth (height growth from 1998 to 2003/height in 1998) than western hemlock. However, relative height growth of released versus suppressed trees was higher in Douglas-fir (130%) than in western hemlock (65%), indicating that, although absolute height growth was less, Douglas-fir did release from suppression. Release seemed to be constrained initially by a limited photosynthetic capacity in both species. Five years after release, Douglas-fir trees had 14 times the leaf area and 1.5 times the leaf nitrogen concentration (N (area)) of suppressed trees. Needles of released western hemlock trees had about twice the maximum assimilation rate (A (max)) at ambient [CO(2)] as needles of suppressed trees and exhibited no photoinhibition at the highest irradiances. After release, trees increased in leaf area, leaf N concentration and overall photosynthetic capacity. Subsequently, hydraulic architecture appeared to constrain release in Douglas-fir and, to a lesser extent, in western hemlock. Released trees had significantly less negative foliar delta(13)C values than suppressed trees and showed a positive relationship between leaf area:sapwood area ratio (A (L)/A (S)) and delta(13)C, suggesting that trees with more leaf area for a given sapwood area experienced a stomatal limitation on carbon gain. Nonetheless, these changes had no significant effects on leaf specific conductivities of suppressed versus released trees of either species, but leaf specific root conductance was significantly lower in released Douglas-fir.     

Effects of red alder and paper birch competition on juvenile growth of three conifer species in southwestern British Columbia

We examined the effects of competition from red alder (Alnus rubra Bong.) and paper birch (Betula papyrifera Marsh.) on the growth of western redcedar (Thuja plicata Donn), western hemlock (Tsuga heterophylla Sarg.), and Douglas-fir (Pseudotsuga menziesii Franco) at a site near Maple Ridge, British Columbia, Canada. At this site, the three coniferous species and two broadleaf species had been planted in 1999 as part of a long-term experiment examining effects of broadleaf density on conifer growth. Red alder and paper birch were planted alone and as a 50:50 mixture at four densities (0, 277, 556, and 1150 stems ha⁻¹).     

Predicting average growth and size distributions of Douglas-fir saplings competing with sprout clumps of tanoak or Pacific madrone

The average growth and size distributions of Douglas-fir (Pseudotsuga menziesii) saplings in three plantations were studied for 7 years (1983–1989) after thinning of associated sprout clumps of tanoak (Lithocarpus densiflorus) or Pacific madrone (Arbutus menziesii); in some cases understory vegetation (shrubs and herbs) was experimentally suppressed. Biologically based nonlinear equations explained 66, 90, and 53% of the variation in the average annual increment of Douglas-fir height, diameter-squared, and cover, respectively. Equations for annual increment of cover of hardwood and understory vegetation explained only 10 to 12% of the variation, because these parameters exhibited a high degree of variability. Model simulations demonstrated that, for the same initial levels of cover, tanoak had faster rates of cover growth than madrone and also caused greater limitations in growth of Douglas-fir. Suppression of understory vegetation increased growth of Douglas-fir only when hardwood cover was absent. Weibull functions adequately described size distributions for Douglas-fir in 92% of the individual-tree data sets. Regression functions of hardwood cover and average Douglas-fir size explained 51, 93, and 24% of the variation in the Weibull A, B, and C parameters, respectively. Model simulations with predicted Weibull parameters demonstrated that hardwood competition caused a positive skewing in size distributions for height and stem diameter of Douglas-fir.     

Tall planting stock for enhanced growth and domination of brush in the Douglas-fir region

Two long-term experiments followed development of planted stock of various sizes, origins, and species. In one experiment, multiyear comparisons of container, 2-0 bareroot, and 3-year-old Douglas-fir transplants showed strong positive relationship between initial height and long-term (10–14 years) growth under a range of site conditions with high probability of brush development. In the other experiment, Douglas-fir, western hemlock, and Sitka spruce were planted in salmonberry disturbed by logging 0 and 4 years previously. Half the seedlings were released with glyphosate 6 months after planting. Hemlock and Douglas-fir bare-roots all grew well if planted in a fresh burn, despite rapid regrowth of salmonberry, but virtually all seedlings less than 60 cm tall except Sitka spruce were killed by 4-year-old salmonberry if not released. Release improved growth of seedlings in the fresh burn by 6%, gaining an average of about 0.6 year toward reaching a height of 6 m. Release improved growth of survivors in 4-year-old salmonberry by 51% in height, 72% in diameter, and 325% in volume at age 12. Sitka spruce grew well until damaged by insects. Hemlock growth was equal to or greater than that of Douglas-fir of comparable initial height. In all comparisons, the probability of being overtopped by brush decreased with increasing initial stock height, and the effect of suppression on growth was also inversely related to initial height. Field planting operations may require special logistic measures for the largest stock types.This is Paper 2706 of the Forest Research Laboratory, Oregon State University.     

Stem volume equations and basic density for grey alder and common alder in Sweden

The objective was to determine stem volume models for grey andcommon alders and, based on the models, stand volume for naturallyregenerated grey and common alder stands was summarized. Basicdensity for grey and common alders and mean annual growth forstands was estimated. Net volume accretion data were collectedfrom 24 stands of grey alder (Alnus incana (L.) Moench) and31 stands of common alder (Alnus glutinosa (L.) Gaertner) inSweden. The stands ranged in latitude from 58 to 64° N andfrom 56 to 62° N for grey and common alder, respectively.The mean age of grey and common alder stands was 41 years and48 years, respectively, the mean stand density 1726 stems ha^–1and 1078 stems ha^–1, and the mean diameter at breast height(over bark) was 20 cm and 21 cm. Stem volume equations weredeveloped for grey and common alders. The adopted model forgrey alder was based on diameter at breast height and height.For common alder, crown height was added to diameter and height.Mean standing volume (over bark) for grey and common alder standswas 428 and 374 m³ ha^–1. Mean annual growth for grey andcommon alder stands was 12.0 m³ and 8.4 m³ a^–1 ha^–1,respectively. Basic density (under bark), for grey and commonalder stems was 359 and 427 kg m^–3, respectively. Thebasic density (under bark) for the lowest twigs in the crownand in the lateral part of the crown was 415 and 421 kg m^–3for grey alder and 423 and 423 kg m^–3 for common alder.     

Effects of diversity of tree species and size on forest basal area growth,recruitment, and mortality

The objective of this study was to determine the relationship, or lack thereof, between growth and diversity of tree species and size in conifer stands of western North America. Growth was measured by net basal area growth and its components: survivor growth, recruitment, and mortality. The analysis used inventory data from permanent plots in the Douglas-fir/western hemlock forest type in Oregon and Washington, and in the mixed-conifer forest type in California. The methods consisted of generalized least square regression with spatial autocorrelation, controlling for the effect of other stand characteristics. Other things being equal, in the two forest types under study there was a strong positive relationship between net basal area growth and tree-species diversity. This effect was associated with higher recruitment in stands of higher tree-species diversity. Neither mortality nor growth of survivors was related to tree-species diversity. The relationship between growth and tree-size diversity was less clear. For Douglas-fir/western hemlock, net basal area growth was negatively correlated with tree-size diversity, essentially because recruitment was lower on plots of high tree-size diversity. For mixed conifers, net basal area growth tended also to be lower in plots of high tree-size diversity, but this was mostly because mortality was higher in plots of higher tree-size diversity.     

Modeling top height growth of red alder plantations

Height growth equations for dominant trees are needed for growth and yield projections, to determine appropriate silvicultural regimes, and to estimate site index. Red alder [Alnus rubra Bong.] is a fast-growing hardwood species that is widely planted in the Pacific Northwest, USA. However, red alder dominant height growth equations used currently have been determined using stem analysis trees from natural stands rather than repeated measurements of stand-level top height from plantations, which may cause them to be biased. A regional dataset of red alder plantations was complied and used to construct a dynamic base-age invariant top height growth equation. Ten anamorphic and polymorphic Generalized Algebraic Difference Approach (GADA) forms were fit using the forward difference approach. The Chapman–Richards anamorphic and Schumacher anamorphic model forms were the only ones with statistically significant parameters that yielded biologically reasonable predictions across a full range of the available data. The Schumacher model form performed better on three independent datasets and, therefore, was selected as the final model. The resulting top height growth equations differed appreciably from tree-level dominant height growth equations developed using data from natural stands, particularly at the younger ages and on lower site indices. Both the rate and shape parameters of the Schumacher function were not influenced by initial planting density. However, this analysis indicates that the asymptote, which is related to site index, may be reduced for plantations with initial planting density below 500 trees ha⁻¹. The final equation can be used for predictions of top height (and thus) site index for red alder plantations across a range of different growing conditions.     

Underplanting western hemlock in a red alder thinning: early survival,growth, and damage

Underplanting conifers beneath thinned hardwood stands could shorten conversion to a more valuable crop species. Thinning the hardwoods to final crop-tree spacing could increase growth and marketable volume while releasing the site resources necessary to support conifers planted in the understory. In western Oregon, an experimental thinning of a 14-year-old red alder (Alnus rubra Bong.) stand provided an opportunity to test this concept. Initial efforts were directed toward testing survival and growth of wilding seedlings of western hemlock (Tsuga heterophylla [Raf.] Sarg.) planted in thinned and unthinned alder stands. Survival averaged 78% and 52% after the first and fifth growing seasons. Fifth-year height growth of surviving seedlings averaged from 38 to 49 cm under various thinning regimes but only 9 cm in the unthinned. Wildlife browsing and the pinning of seedlings by falling debris reduced growth and survival.     

Individual-tree height-, diameter- and crown-width increment equations for young Douglas-fir plantations

This work presents the results from the initial model development of a simulator to predict vegetation dynamics in young plantations growing in a Mediterranean environment. The simulator can predict growth dynamics for coniferous crop trees as well as competing hardwoods and shrubs. Model specification included conifer, shrub, and hardwood competition expressed at the plot-level. The system employs water-holding capacity as an indicator of productivity. Growth data were obtained from 109 plantations, ranging in age from 3 to 25, in southern Oregon and northern California. Douglas-fir (Pseudotsuga menziesii Mirb. Franco), the most common conifer species, was observed in 80 of the sampled stands. These observations were used for model development of the primary driving functions, which forecast 2-year basal diameter increment, height increment, and crown width increment. Parameters for all three dynamic expressions for growth were estimated using weighted, nonlinear three-stage least squares. This estimation method provided a predictive model with slight improvements in standard errors for two of the three equations (an average of 3% for height and diameter growth) and no improvement for crown width, when compared with two-stage least squares. The system includes competition from shrubs and hardwoods in predictions of height growth, diameter growth and crown width increment. This allows individual-tree/distance-independent simulator architecture to be extended to young plantations in southern Oregon and northern California.     

Response of branch growth and mortality to silvicultural treatments in coastal Douglas-fir plantations: Implications for predicting tree growth

Static models of individual tree crown attributes such as height to crown base and maximum branch diameter profile have been developed for several commercially important species. Dynamic models of individual branch growth and mortality have received less attention, but have generally been developed retrospectively by dissecting felled trees; however, this approach is limited by the lack of historic stand data and the difficulty in determining the exact timing of branch death. This study monitored the development of individual branches on 103 stems located on a variety of silvicultural trials in the Pacific Northwest, USA. The results indicated that branch growth and mortality were significantly influenced by precommercial thinning (PCT), commercial thinning, fertilization, vegetation management, and a foliar disease known as Swiss needle cast [caused by Phaeocryptopus gaeumannii (T. Rohde) Petr.]. Models developed across these datasets accounted for treatment effects through variables such as tree basal area growth and the size of the crown. Insertion of the branch growth and mortality equations into an individual-tree modeling framework, significantly improved short-term predictions of crown recession on an independent series of silvicultural trials, which increased mean accuracy of diameter growth prediction (reduction in mean bias). However, the static height to crown base equation resulted in a lower mean square error for the tree diameter and height growth predictions. Overall, individual branches were found to be highly responsive to changes in stand conditions imposed by silvicultural treatments, and therefore represent an important mechanism explaining tree and stand growth responses.     

Long-term nutrient cycling patterns in Douglas-fir and red alder stands: a simulation study

Long-term patterns in nutrient cycling in regrowing Douglas-fir (Pseudosuga menziesii Mirb. Franco) and red alder (Alnus rubra Bong.) on native soils plus soils previously occupied by other species were simulated using the nutrient cycling model. Simulations of regrowing stands were also compared with observations of nutrient cycling in mature Douglas-fir and red alder. We hypothesized that (1) prolonged presence of red alder will cause a depletion in soil base cations due to increased nitrification and NO₃⁻ leaching; (2) lower base cation availability under red alder will ultimately cause biomass production to decline; (3) high N availability in red alder soils will favor regrowth of Douglas-fir; (4) higher base cation and P status of the Douglas-fir soils will favor growth of red alder both in the short- and long-term, since N is not limiting to red alder; and (5) in regrowing red alder, NO₃⁻ leaching will increase with time as a result of increased N fixation. All hypotheses were confirmed, but the effect of soil type on biomass production was minimal both for red alder and Douglas-fir. The higher soil organic matter content in the mature red alder stand most likely reflected previous occupation by old-growth Douglas-fir and also a large litter input from the understory vegetation. In general, the nutrient cycling model simulated differences in nutrient cycling patterns at least qualitatively between Douglas-fir and red alder and was helpful in identifying potential gaps in the understanding of biogeochemical cycling as well as uncertainties in the data. The nutrient cycling model did not fully elucidate differences in P cycling between Douglas-fir and red alder and overestimated weathering rates under Douglas-fir. Uncertainties in the data included: (1) temporal patterns in N fixation in the regrowing stands; (2) understory litterfall; and (3) site history and, consequently, presence of pre-existing differences in site conditions.     

兴安落叶松树干去皮直径预测模型

利用兴安落叶松解析木数据,比较了树干去皮直径预测的3种类型模型:Grosenbaugh的比率方程式、回归模型和削度方程。Grosenbaugh的比率方程式有很大的灵活性,没有参数不需要模型拟合。总体评价和模型分段比较表明,回归模型有较小的预测误差,尤其是Cao and Pepper提出的含有带皮直径、树高、相对树高、胸径处的带皮直径和去皮直径变量的模型。由于削度模型不含有带皮直径变量,因此产生较大的去皮直径预测误差。不同类型的模型在森林经营过程中都有一定的适应性。     

The effect of short day treatments on containerized Douglas-fir morphology,physiology and phenology

The effect of short day treatments (blackout) on Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) container seedlings at the time of lift and following cold storage was investigated. Variables measured included height, root collar diameter (RCD), root growth capacity (RGC), photosynthetic efficiency after –18 °C freezing (PEF), and days to terminal bud break (DBB). From one to four blackout dormancy induction treatments were started on three dates (July 12, July 26, and August 10) with 10 or 20 d between multiple blackouts. Increasing the number of blackout treatments resulted in lower RCD, lower DBB in the late winter/early spring, and higher PEF in the early fall. Later blackout start dates decreased PEF in the early fall, and increased overall height and late fall RGC as compared to earlier blackout start dates. Nurseries growing Douglas-fir seedlings from coastal Pacific Northwest provenances should be aware that blackout regimes can decrease RGC in the late fall, and cause quicker dormancy release in the early spring. Coastal Douglas-fir can be lifted and planted in the early fall, when RGC and DBB are relatively high. If planting between February and April is necessary, seedlings given blackout should be cold stored in January to maintain an adequate level of dormancy, RGC and PEF.     

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.     

Distance-independent tree basal area growth models for Norway spruce,Douglas-fir and Japanese larch in Southern Belgium

This paper presents new harmonized distance-independent individual tree basal area growth models for Norway spruce, Douglas-fir and Japanese larch in pure even-aged stands in Southern Belgium. The selected model was originally developed for Norway spruce and Douglas-fir in neighboring France. New formulations are proposed for some of the model components in order to lower the number of fitted parameters and facilitate the fitting procedure. The resulting models integrate the most recent corresponding top-height growth models and use four simple and usually collected explanatory variables: stand age, top-height, total basal area and tree girth at breast height. The modified formulations maintain similar fitting performances and make it easier to interpret the influence of the explanatory variables on tree growth. Parameters estimates were fitted on thousands of growth measurements gathered from several monitoring plots, forest management inventories and silvicultural field experiments that represent the wide range of site conditions and of forest management scenarios applied to coniferous stands in Southern Belgium. Cross-validation of the models revealed no bias and highlighted their consistent behavior over the entire range of girth at breast height, age, top-height, site index and density represented in our dataset. Combining utility and robust performances, these models represent useful forest management tools, purposely ideal for forest simulation software development. Moreover, the flexibility and generic capabilities of the model formulation should make it easily adjustable for other species in even-aged stands.     

Do individual-tree growth models correctly represent height:diameter ratios of Norway spruce and Scots pine?

Height:diameter ratios are an important measure of stand stability. Because of the importance of height:diameter ratios for forest management, individual-tree growth models should correctly depict height:diameter ratios. In particular, (i) height:diameter ratios should not exceed that of very dense stands, (ii) height:diameter ratios should not fall below that of open-grown trees, (iii) height:diameter ratios should decrease with increasing spacing, (iv) height:diameter ratios for suppressed trees should be higher than ratios for dominant trees. We evaluated the prediction of height:diameter ratios by running four commonly used individual-tree growth models in central Europe: BWIN, Moses, Silva and Prognaus. They represent different subtypes of individual-tree growth models, namely models with and without an explicit growth potential and models that are either distance-dependent (spatial) or distance-independent (non-spatial). Note that none of these simulators predict height:diameter ratios directly. We began by building a generic simulator that contained the relevant equations for diameter increment, height increment, and crown size for each of the four simulators. The relevant measures of competition, site characteristics, and stand statistics were also coded. The advantage of this simulator was that it ensured that no additional constraint was being imposed on the growth equations, and that initial conditions were identical. We then simulated growth for a 15- and 30-year period for Austrian permanent research plots in Arnoldstein and in Litschau, which represent stands at different age-classes and densities. We also simulated growth of open-grown trees and compared the results to the literature. We found that the general pattern of height:diameter ratios was correctly predicted by all four individual-tree growth models, with height:diameter ratios above that of open-grown trees and below that of very dense stands. All models showed a decrease of height:diameter ratios with age and an increase with stand density. Also, the height:diameter ratios of dominant trees were always lower than that of mean trees. Although in some cases the observed and predicted height:diameter ratios matched well, there were cases where discrepancies between observed and predicted height:diameter ratios would be unacceptable for practical management predictions.     

Basal area growth models for individual trees of Norway spruce, Scots pine, birch and other broadleaves in Norway

Distance-independent individual tree growth models based on about 30,000 observations from the National Forest Inventory and the Norwegian Forest Research Institute have been developed for the main tree species in Norway. The models predict 5-year basal area increment over bark for trees larger than 5 cm at breast height. Potential input variables were of four types: size of the tree, competition indices, site conditions, and stand variables including species, mixtures and layers. The squared correlation coefficient (R²) varied from 0.26 to 0.55. The accuracy of the models was tested by comparing the individual tree models with Norwegian diameter increment models. The accuracy is similar, but individual tree models forecast diameter distributions directly. The inclusion of species mixture and layer as variables increases the reliability of the models in mixed and in uneven-aged stands.     

Branch characteristics of widely spaced Douglas-fir in south-western Germany: Comparisons of modelling approaches and geographic regions

Models of Douglas-fir branch and whorl characteristics were developed from contrasting spacing experiments in southwest Germany. The dataset was based on 100 young (20–30 years old), unpruned and partially pruned trees from a 100, 200, and 1200 stems ha⁻¹ spacing experiment on Douglas-fir that was replicated 3 times across the region. The material was used to predict (1) the number of branches whorl⁻¹, (2) branch angle, (3) status (living/dead) of the branches within the living crown, (4) maximum branch diameter whorl⁻¹, and (5) relative diameter of branches within a whorl. For each of these models (except branch status), both a linear and nonlinear, generalised hierarchical mixed effects equation was developed. The comparison of the linear and nonlinear approaches showed that both had a relatively similar level of bias, but the nonlinear equations generally performed better (reduction in mean absolute error of 1.1–69.5%). Overall, individual branch and tree properties were sufficient to give logical and precise predictions of the branch characteristics for the models across the range of sampled stand densities. In addition, the models showed a similar behaviour compared to models on Douglas-fir crown structure from the Pacific Northwest, USA. This suggests that the allometric relationship between tree size and branch characteristics for a given species may be relatively consistent across regions, even ones with highly contrasting growing conditions like in this study. The models performed well across a range of stand conditions and now will be further integrated into an individual tree growth and yield simulations system.