ALBERT

Description: Seedlings of Avicenniagerminans (L.) Stearn., Lagunculariaracemosa (L.) Gaertn., and Rhizophoramangle (L.) were subjected to flooding, signified by soil redox potentials around −92 mV, and salinity in the range of 342 mol•m−3. Leaf conductance and net carbon assimilation rates per unit area of leaf did not change significantly under flooding or salinity treatments compared with control plants. There was no significant interaction of flooding and salinity with leaf conductance and net carbon assimilation; however, significant reduction in total leaf area per plant in response to flooding (minus salinity) was found in L. racemosa and A. germinans compared with control plants, which would result in a substantial reduction of net carbon assimilation per plant. In R. mangle, total leaf area per plant did not change significantly in response to various treatments. Generally, salinity alone or combined with flooding enhanced dry weights, whereas flooding (minus salinity) resulted in reduced dry weights. The mean values of leaf conductance and net carbon assimilation differed significantly among the study species, with greatest values recorded in A. germinans. The differences in conductance in combination with changes in net carbon assimilation rates resulted in substantial differences in water-use efficiency among these species. Water-use efficiency was greatest in L. racemosa. The overall results showed that these species were tolerant of a wide range of salinity and waterlogging conditions, with differences in physiological responses being evident in changes in biomass partitioning.

Description: Trees and their environment were studied in floodplain forests in the glaciated region of northern Missouri. Ordination of tree vegetation samples by detrended correspondence analysis indicated a primary vegetation continuum of decreasing Acersaccharinum L. and increasing Caryalaciniosa (Michx. f.) Loud with several other species associated with secondary vegetation gradients, TWINSPAN classification of tree vegetation identified three groups of plots that were dominated by A. saccharinum in varying degrees of association with other species, most importantly Populusdeltoides Bartr. and Ulmusamericana L.; two groups dominated by species of Carya and Ulmus; and two groups where dominance was more broadly distributed among lowland Quercus spp., U. americana, Aesculusglabra Willd., Fraxinuspennsylvanica Marsh., Platanusoccidentalis L., and Betulanigra L. Importance of A. saccharinum was greatest in plots where the leading dominants were young, while Quercus and Carya spp. were more common in plots with older leading dominants, suggesting that the predominant environmental influence on vegetation composition was frequency and severity of disturbance associated with flooding. Plots with younger dominant trees had lower species richness and diversity than plots with older dominants. Higher soil pH and slough location were also positively correlated with A. saccharinum importance, and Acernegundo L. was more frequent in sloughs. Analysis of overstory and understory relationships indicated that A. saccharinum is likely to remain important in the immediate future in many forests currently dominated by this species because of its abundance in subcanopy positions. Potential canopy trees of F. pennsylvanica, C. laciniosa, Caryacordiformis (Wangenh.) K. Koch, and Celtisoccidentalis L. may, in the absence of major flooding disturbance, cause long-term shifts in composition in some of these forests. While Ulmus spp. are abundant in the understory, they are unlikely to become important canopy species because of disease.

Description: The effects of climate on the growth of sugar maple (Acersaccharum Marsh.) were studied at five sites along an 800-km acidic deposition gradient from Cook County, Minnesota, to Oceana County, Michigan. Fifty increment cores were taken from 25 dominant and codominant individuals at each site in the spring of 1990. Annual ring widths (1940–1989) were measured to 0.01 mm, standardized by taking the first logarithmic differences, and averaged into chronologies using the biweight mean. The five resulting chronologies were then related to climate using least squares regression techniques. The analyses indicate that temperature is associated with sugar maple growth to a greater degree than precipitation, though there were differences in the relationships among the five study sites. Growth was also found to be significantly affected by prior growing season conditions. No evidence of an overall decline or increase in sugar maple growth rates was observed over the 50-year study period.

Description: Calamagrostiscanadensis (Michx.) Beauv. is a widely distributed rhizomatous grass that can seriously inhibit growth of white spruce (Piceaglauca (Moench) Voss) seedlings in the boreal forests of North America. We review the dynamics of this grass during four successional stages: the colonization of disturbed sites; dominance of the site by the grass a few years after disturbance; gradual loss of dominance with overstory development; and maintenance of the grass at low levels in the understory of the mature forest. We also describe C. canadensis in relation to recruitment from clonal growth and seed, environmental conditions for growth, the effects of grass litter buildup on conifer seedling microclimate, and overall competitive abilities. Control strategies for C. canadensis are as follows. If the grass is found in nearly every square metre in the understory prior to logging, there will be rapid spread when the stand is clear-cut unless clones are killed using herbicides or a deep burn. Large spruce seedlings, planted on large soil scalps or mounds, coupled with release by way of herbicides or sheep grazing, may be necessary for plantation establishment under conditions of encroachment by C. canadensis. Alternatively, the shade provided by a partial canopy may inhibit the grass sufficiently to allow spruce seedlings to establish. If grass is not abundant in the understory, we recommend (i) minimizing forest floor disturbance to reduce sites for grass seedling colonization or (ii) a slash burn with the hope of encouraging colonization by herbaceous species that have less impact on conifer seedlings.

Description: In an open-pollinated progeny trial of Pinusradiata D. Don, stem diameter assessments were cross-referenced for 410 families for ages 5, 10, and 17 years from planting. Also cross-referenced were Cyclaneusma needle cast (CYCLA) and wood density (PILO) measured by Pilodyn needle penetration. Estimated narrow-sense heritability for stem diameter declined mildly from 0.34 at age 5 to 0.25 at age 17. Estimated heritability of family means, however, only declined from 0.59 to 0.55. CYCLA and PILO gave, respectively, narrow-sense heritability estimates of 0.32 and 0.40, with repeatabilities of family means of 0.57 and 0.67. The genetic age-age correlations for stem diameter were all positive and somewhat higher than phenotypic (family-mean) age–age correlations. Such correlations indicated comparable or slightly slower rank changes among progeny families than had been reported previously for diameter, basal area, or stem volume in P. radiata and Pinustaeda L., but faster rank changes than the literature reports for tree height. A considerable contribution of CYCLA to rank changes in stem diameter was evident from path coefficients and partial correlations. PILO made no evident contribution to rank changes. Predicted gains for stem diameter at age 17 were almost maximal using year-10 data, while using CYCLA as an auxiliary selection criterion enhanced expected gain, particularly with selection at year 5. Predicted gains for stem diameter, with age–age correlations extrapolated according to the Lambeth relationship, indicated maximal gains per annum with selection at 7–8 years for rotations of 25–30 years.

Description: Crown recession rates were estimated by branch mortality dating on 357 sectioned Douglas-fir (Pseudotsugamenziesii (Mirb.) Franco) stems from temporary plots. Numerous nonlinear, logarithmic, and gamma-theory generalized linear models were developed for predicting 5-year crown recession across a range in tree, stand, and site conditions. Residual analyses and indices of fit demonstrated that a multiplicative model with lognormal errors was the most appropriate model form. The recommended logarithmic model predicts crown recession from current crown ratio, total height, breast height age, height growth, and crown competition factor. Data from southwestern Oregon indicate that within a given stand, trees with midsized crown ratios experience the most rapid crown recession.

Description: Bulk density of forest soils from nine locations in New England was closely and inversely related to the organic fraction of the soil. Measured data over the whole range of organic fractions followed the theoretical relationship Db = DbmDbo/[FoDbm + (1 − Fo)Dbo] where Db is the bulk density (Mg/m3), Fo is the organic fraction (kgo/kg), Dbo is the bulk density when Fo = 1, and Dbm is the bulk density when Fo = 0. The relation arises from assuming that (i) Dbo, the bulk density of "pure" organic matter, and Dbm, the bulk density of "pure" mineral matter, are constant and (ii) in a mixture, the volumes occupied by the organic mass and the mineral mass are additive. For forest soils on coarse-textured till in New England, Dbo = 0.11 Mg/m3 and Dbm varied from 1.45 Mg/m3 for sandy loams to 2.19 Mg/m3 for silt loams. When these parameters are known, Db can be estimated from Fo, which is more easily measured. When Fo is greater than 0.1 kgo/kg, the organic mass per unit soil volume (FoDb), or organic density, is approximately constant at 0.1 Mgo/m3. For many nutrients, separate evaluation of the organic density and the amount of nutrient per unit organic mass may facilitate intersite comparisons for studies of nutrient availability and leaching.

Description: Boles of quaking aspen (Populustremuloides Michx.), white spruce (Piceaglauca (Moench) Voss), red pine (Pinusresinosa Ait.), and jack pine (Pinusbanksiana Lamb.) were sampled after decomposing for 11–17 years. Mass loss and changes in chemical composition were determined. Density decreased by 40–73%, and the decomposition constant (k) decreased in the order aspen 〉 spruce 〉 red pine 〉 jack pine. Although the decomposition rate was strongly species dependent, it did not differ between the two sites. The decomposition constant was weakly related to lignin and phosphorus concentration in the original stems (r2 = 0.44 and 0.49, respectively) and not significantly related to original nitrogen concentrations. Nutrient concentration increased during decomposition, particularly for nitrogen and phosphorus. Nitrogen content increased during decomposition, and potassium content decreased. Calcium and magnesium content changed little during decomposition, whereas phosphorus content increased or decreased, depending on species. In general, species with the lowest initial nutrient contents had the greatest nutrient increases during decomposition. Nutrient content of boles of all species became similar during decomposition.

Description: The internal transport of carbon dioxide by water flow has until now been omitted when interpreting measurements of photosynthesis and respiration. Theoretical and empirical analyses of the behaviour of carbon dioxide within a tree show that the consideration of dissolved carbon transportation may be important when estimating the respiration rate, but not so important when considering photosynthetic production.

Description: Changes in tree form and taper over time, as affected by changes in tree, stand, and site factors for interior lodgepole pine (Pinuscontorta var. latifolia Engelm.) were investigated using detailed stem analysis data from interior British Columbia. It was found that tree shape and taper change along the stem at one time and over time with changes in tree and stand factors, particularly the diameter at breast height to total tree height ratio, crown length, and crown ratio, and with predicted quadratic mean diameter at age 50 years, a stand density measure. At young ages, the trees were parabolic in shape from ground to top. However, as they increased in size over time, different portions of the stem took different shapes because of unequal growth in diameter along the stem. Changes in tree shape and taper over time were closely related to the crown size, which is related to stand density.

Description: Estimates of individual-tree narrow-sense heritability and additive genetic coefficient of variation of seven traits of forest trees were compiled from 67 published papers. Distributions of the values for each trait were characterized and compared by calculating medians and running Kolmogorov–Smirnov and Wilcoxon signed-rank tests. Generalizations are possible about at least some of the traits examined. Heritability of wood specific gravity was almost always above 0.3 (median 0.48). Heritabilities for other traits tended to be low: medians ranged from 0.185 to 0.26, and individual values generally ranged from 0.1 to 0.4. Evidence that heritabilities of form traits tend to be higher than those of growth traits was weak. The analysis of additive genetic coefficients of variation suggested that specific gravity tends to have lower values than other traits (median 5.1%), while height and diameter (medians 8.5 and 8.6%, respectively) had lower values than straightness (median 11.65%). Individual-tree volume showed the highest levels of additive genetic coefficient of variation (median 20.3%). The levels of additive genetic variation and heritabilities suggest that reasonable levels of genetic gain can be achieved by screening relatively low numbers of trees.

Description: Tree-ring analysis was used to study historical patterns of basal area increment (BAI) by healthy (0–5% dieback) and declined (greater than 30% dieback) overstory sugar maples (Acersaccharum Marsh.) in four stands in Pennsylvania. The objectives were to establish if and when BAI decreased in healthy and declined trees and to identify causal factors associated with decreased BAI. Reduction in BAI of declined compared with healthy trees was first evident following a series of defoliations and summer droughts in the mid-1960s and early 1970s. These stresses were followed by repeated damage by pear thrips (Taeniothripsinconsequens Uzel) in the 1980s and a drought in 1988, during which large reductions in BAI occurred for both declined and healthy trees in all stands. Foliar nutrient analysis of two stands suggested deficiency of Mg and Ca. Other factors hypothesized to be associated with this sugar maple decline as either predisposing or inciting factors include unusually warm winter temperatures in the 1980s and air pollution.

Description: On the rocky shores and islands of Lake Duparquet, in the southwestern Quebec boreal forest, Thujaoccidentalis L. reaches ages in excess of 800 years. Annual ring widths from 38 trees were used to develop an 802-year chronology (1186–1987) standardized by polynomial regressions. Excellent cross dating, correlation with a shorter chronology located 14 km inland, and 33.6% common variance in a chronology subsample all point to the existence of a climatic signal. After autoregressive modeling to obtain a serially random residual chronology, correlation and response functions were used to identify the growth–climate relationship. The resulting model reduced 19.2% of the chronology variance. Precipitation in June as well as low temperature in June or July seemed to have a positive influence on growth. Likewise, a drought index was closely related to growth, indicating that the chronology could be used to estimate past drought conditions. Moisture deficits are thus inferred for the 13th century as well as during the Little Ice Age (17th century to late 19th century). Since the end of the latter period, precipitation seems to have followed an upward trend.

Description: Three southern Appalachian stands with sparse and unproductive pine–hardwood overstories and dense Kalmialatifolia L. understories were treated to restore productivity and diversity on steep slopes. An adaptation of the fell and burn practice was applied in summer and fall 1990. About one-half of the woody fuels were consumed at each site. A range of fire intensities was observed. Flame temperatures approached 800 °C, but the heat pulse into the forest floor only reached 60 °C at 5 cm. Humus and charred leaf litter remained on most of the surface after burning. Evidence of soil erosion was spotty and related to points of local soil disturbance. No soil left the sites. At the end of the first growing season, 23% of the burned surfaces were covered by growing plants and 62% by residual forest floor and woody debris. Felling and burning reduced evapotranspiration so that soil in the treated areas remained moister than under adjacent uncut stands. Opening the sites increased soil temperatures 2 to 5 °C at 10 cm during the first 16 months after treatment.

Description: Radial growth following a shelterwood seed cut in a 174-year-old stand of white spruce (Piceaglauca (Moench) Voss) in interior Alaska was compared with growth in an adjacent undisturbed stand of the same age. After a 2-year lag, radial growth of residual trees accelerated an average of 27% in 5 of the next 6 years. Net mean increase in growth after 8 years was 164%. Basal area growth of individual shelterwood trees increased 26.8% over the 14-year posttreatment period, while control trees increased 16.5%.

Description: In view of the possible applications of ectomycorrhizae to forestry, this paper discusses the important functions of ectomycorrhizae, the conditions affecting their formation, and methods for the production and application of inoculum. A rationale for selecting appropriate ectomycorrhizal fungi and considerations in selecting sites where ectomycorrhizal seedlings should be planted are presented. Suggestions are also made on encouraging the use of ectomycorrhizal technology. A cost–benefit analysis of inoculation is done.

Description: Three paired watersheds treated with a fell and burn prescription were studied to determine the effects on soil, soil water, and stream water. Soil nitrification and mineralization were measured by in situ closed-core incubation. Soil water was collected with porous cup lysimeters placed at 30 and 60 cm depths, and water samples were collected from streams draining control and burned areas on one of the three sites. All data were collected for 6 months prior to and 12 months after treatment. Soil ammonium (NH4+) content increased significantly in all three sites after burning, but the magnitude differed greatly among sites. However, there was no change in soil nitrate (NO3−) content. In situ measurements of net mineralization showed increased rates with increasing burn severity. Net nitrification displayed no treatment response. Slight and nonsignificant increases in soil water NO3− concentration occurred after burning in two of the three sites. Stream water NO3− concentrations increased in the one stream sampled. Thus, while prescribed burning increased available soil N, there was little change in N transformation rates or movement of dissolved inorganic N off site during the first year after burning.

Description: By measuring incident precipitation, throughfall, and stemflow chemistry, the roles of coniferous- and deciduous-dominated forest canopies as a source of and sink for ions in precipitation were examined. A regression technique for distinguishing between external (dry deposition) and internal (canopy leaching) sources of ions in the throughfall flux was evaluated. The effect of seasonal changes in the forest canopy on throughfall and stemflow chemistry was also examined. Throughfall comprised 74 and 84%, respectively, of the hydrologic flux at the coniferous and deciduous sites. Sulphate fluxes were highest at the coniferous site during both growing and dormant seasons, suggesting either a higher scavenging efficiency of the needles for atmospheric SO42−, or higher SO42− leaching from the foliage. The deciduous site neutralized acidic inputs, as demonstrated by its net negative H+ flux year round. The buffering capacity of the coniferous forest was exceeded by the higher amount of acid interception by the canopy. Nitrate behaved conservatively and base ions were exported from the canopy. Stemflow contributions of ions, although low, were generally higher than the contribution of stemflow to the hydrologic flux (2–3%). Independent dry deposition measurements for the growing season, when compared with net SO42− flux, overestimated dry deposition collected by the deciduous canopy, but were comparable to the flux at the coniferous site. These data suggest that dry SO2−SO42− deposition may be responsible for all SO42− enrichment seen in throughfall at these sites. A regression technique for separating internal and external ion sources in throughfall yielded inconsistent results, and attributed virtually all ion enrichment to internal sources. Problems with false assumptions and spurious correlations are discussed. We conclude that this method is not satisfactory for separating ion sources. Seasonal patterns in throughfall chemistry are present. During the growing seasons bases exchange for H+ and are exported similarly with SO42−. Hydrogen retention mirrors SO42− export. Base cations (particularly K+) are leached from the canopy primarily during senescence, but from the stem of the tree primarily during the dormant period. This was most evident at the deciduous site. Chloride behaved in a similar manner, while NH4+ and H+ were retained during the senescent period.

Description: Change in the health of sugar maple (Acersaccharum Marsh.) and associated northern hardwoods was evaluated for 3 years (1988–1990) in seven states and four provinces. Generally, levels of crown dieback and crown transparency (a measure of foliage density) in 165 stands decreased during this period. In 1990, less than 7% of all dominant–codominant sugar maples (n = 7317) exhibited crown dieback ≥ 20%. Significantly (p = 0.05) fewer of these maples were classified as having high crown transparency (≥ 30%) in 1990 compared with 1988. Crowns of maples that received moderate (31–60%) or heavy (〉 60%) pear thrips (Taeniothripsinconsequens (Uzel)) damage for 1 year recovered the following year. Crowns of maples exposed to severe drought in 1988 (Wisconsin) continued to show the effects (high transparency) of this stress in 1990. A majority (69–71%) of the dominant–codominant sugar maples with high (≥ 20%) crown dieback had bole and (or) root damage. Of those maples with crown dieback ≥ 50%, 86% had bole and (or) root damage. The condition of sugar maple in operating sugar bushes and undisturbed stands was similar. The condition of sugar maple crowns was similar in locations presumably exposed to low, medium, and high levels of sulfate deposition.

Description: A series of experiments on field-grown seed-derived trees between 2 and 17 years old demonstrated that the growth regulator paclobutrazol could be used to reduce vegetative growth and enhance flower-bud production in Eucalyptusglobulus Labill. and Eucalyptusnitens (Dean & Maid.) ex Maid. Responses to high levels of trunk injection and collar drenching persisted for up to six growing seasons, yielding both increases in frequency of flowering and heaviness of bud crop. Growth responses were expressed in the immediate growing season, but flowering responses were not evident for another year. Foliar spray treatments reduced vegetative growth in young trees of both species for one growing season, but only the E. globulus showed an associated flowering response. Assessment of seed yield per capsule and subsequent germination tests showed no deleterious effects on seed development or quality. Choice of application method will vary with objective and size of tree. Collar drenching shows the most promise for treating large numbers of seed orchard trees because application time is substantially independent of tree size and weather conditions.

Description: This paper describes a decision support system that forest managers can use to help evaluate short-term, site-specific silvicultural operating plans in terms of their potential impact on long-term, forest-level strategic objectives. The system is based upon strategic and tactical forest-level silvicultural planning models that are linked with each other and with a geographical information system. Managers can first use the strategic mathematical programming model to develop broad silvicultural strategies based on aggregate timber strata. These strategies help them to subjectively delineate specific candidate sites that might be treated during the first 10 years of a much longer planning horizon using a geographical information system and to describe potential silvicultural prescriptions for each candidate site. The tactical model identifies an annual silvicultural schedule for these candidate sites in the first 10 years, and a harvesting and regeneration schedule by 10-year periods for aggregate timber strata for the remainder of the planning horizon, that will maximize the sustainable yield of one or more timber species in the whole forest, given the candidate sites and treatments specified by the managers. The system is demonstrated on a 90 000 - ha area in northeastern Ontario.

Description: Biomass allocation to roots was studied in holm oak (Quercusilex L.), a dominant evergreen tree in broad-leaved sclerophyllous Mediterranean forests. The root systems of 32 single-stemmed holm oaks growing in shallow soils on largely unfissured bedrock were excavated in a mesic site and a xeric site in the Montseny Mountains (northeast Spain). Individual root:shoot biomass ratios (roots with diameter

Description: A simple system for the estimation of stem volume is presented based on the compatible stem profile and volume equations. This system can directly predict the stem volume above breast height from measurements of stem diameter at breast height and at an another point along the upper stem, and does not require any sample data for determining a parameter of volume equation. In comparison with the prediction accuracy of existing volume equations from the literature, using data from Cryptomeriajaponica D. Don, Chamaecyparisobtsusa Endl., and Pseudotsugamenziesii (Mirb.) Franco, this system has the advantage of reducing prediction error.

Description: The responses of CO2 assimilation rate (A), transpiration rate (E), and leaf conductance (g) to increasing leaf to air water vapor concentration difference (ΔW) were investigated (i) using excised shoots from mature trees of Abiesalba, Abiescephalonica, Abiesmarocana, and Abiesnordmanniana and (ii) in situ on a mature tree of Abiesbornmulleriana. Gas-exchange responses to increasing soil drought were also studied in plants of A. bornmulleriana, A. cephalonica, and Cedrusatlantica. Stable carbon isotope composition measurements were carried out on annual growth rings of A. bornmulleriana to estimate the time-integrated values of the ratio of intercellular leaf (Ci) to ambient (Ca) CO2 concentration. Increasing ΔW around the shoots reduced A and g in such a way that either Ci remained constant or its decrease was not pronounced enough for the changes in A to be accounted for by changes in g only. This suggests a direct effect of ΔW on photosynthesis. The different Abies species showed clear differences in water-use efficiency. Abiescephalonica and A. marocana had lower water costs of CO2 assimilation (E/A) than A. nordmanniana and A. alba. It has also been shown that A. cephalonica and A. marocana are characterized by an optimal stomatal control of leaf gas exchange. Stomata closed very rapidly in A. bornmulleriana in response to water supply being withheld, even prior to there being any important decrease in leaf predawn water potential. The stomatal response in C. atlantica was more gradual. In A. bornmulleriana, drought adaptation appears to be linked to the ability to avoid internal water stress, whereas drought adaptation in C. atlantica involves the ability to tolerate internal water stress. The high stomatal sensitivity mA. bornmulleriana is also supported by the isotopic carbon composition data, as shown by the substantial interannual variations in the estimates of Ci/Ca, ranging from 0.48 for the dryest years to 0.61 for the rainy years.

Description: Repeated measures data occur in a wide variety of experimental situations and are often analyzed without full consideration of the statistical issues involved. In this paper, a discussion of model construction, univariate versus multivariate solutions, and statistical assumptions is motivated by examples from a tree physiology experiment. In addition, several examples from the forestry literature are reviewed. It is hoped that this discussion will help scientists with little statistical training to become aware of the different analyses available and perhaps to recognize the associated models in their own research. The examples range from a simple repeated measures design with one within-subject factor and no between-subjects factors to a more complex design involving multiple within-subject and between-subjects factors. The modelling approach used here permits a straightforward comparison between the univariate and multivariate solutions. Although no single approach is consistently best, the multivariate approach is always appropriate and provides the same interpretations as the univariate approach. However, when appropriate assumptions such as sphericity are met, power considerations tend to favor the more traditional univariate analysis.

Description: In the assessment of S cycling in forest ecosystems, solutions passing through the forests are normally analyzed for inorganic SO4; other forms of S are rarely considered. In this study, organic S (estimated as the difference between total S and SO4-S) was measured in canopy and soil solutions from eight forest stands spanning a broad range of overstory and soil types. Organic-S concentrations varied among the different types of solutions and among the forests, with values ranging from 0 to 50 μmol S•L−1. Organic S was ≤10% of total S in precipitation, 5 to 54% in throughfall, 1 to 50% in stem flow, 16 to 46% in O-horizon solution, 11 to 21% in A- or E-horizon solutions, and 0 to 29% in B-horizon solutions. Organic S was positively correlated with organic C and organic N in Douglas-fir (Pseudotsugamenziesii (Mirb.) Franco) and red alder (Alnusrubra Bong.) soil solutions and in Douglas-fir stem flow (r2 = 0.68 to 0.96, p 

Description: A probabilistic model predicts means and variances of the total number and volume of large woody debris pieces falling into a stream reach per unit time. The estimates of debris input are based on the density (trees/area), tree size distribution, and tree-fall probability of the riparian stand adjacent to the reach. Distributions of volume, length, and orientation of delivered debris pieces are also predicted. The model is applied to an old-growth coniferous stand in Oregon's Cascade Mountains. Observed debris inputs from the riparian stand exceeded the inputs predicted from tree mortality rates typical of similar nonriparian stands. Debris pieces observed in the stream were generally shorter, with less volume per piece, than those predicted by the model, probably because of bole breakage during tree fall. As a second application, predicted debris inputs from riparian management zones of various widths are compared with the input expected from an unharvested stand.

Description: A simple conceptual model is proposed concerning how leaf area efficiency (stemwood growth per unit leaf area) changes with leaf area for trees within a stand. Greater leaf area is generally associated with (i) improved light environment due to greater height and (ii) a lower ratio of photosynthetic to nonphotosynthetic tissue. Greater height and improved light environment result in higher photosynthetic production, which should increase leaf area efficiency. A lower ratio of photosynthetic to nonphotosynthetic tissue suggests that the ratio of respiration to photosynthesis increases, which should decrease leaf area efficiency. In relatively small trees, the influence of increased height (associated with greater leaf area) should more than offset the influence of the increased respiration:photosynthesis ratio; as a result, leaf area efficiency should increase with leaf area. In large trees, further increases in leaf area are associated with minimal increases in height, and leaf area efficiency should decline as the respiration:photosynthesis ratio increases. Predictions from this conceptual model were examined with data from stands of subalpine fir (Abieslasiocarpa (Hook.) Nutt.).

Description: Ten forest litters with decomposition state varying from 16.6 to 100% weight remaining were partitioned into sub-samples; each subsample was analyzed for proximate carbon fractions using one of two chemical analysis procedures (forage fiber and forest products analyses). Proximate carbon fractions from the simpler forage fiber techniques accurately estimated extractives, cellulose, lignin, and acid-hydrolyzed carbohydrates (R2 〉 0.83) determined by the more complex forest products analyses. Decomposition state accounted for most of the residual variance and significantly improved predictive equations for lignin and extractives. The relationship between proximate carbon fractions from the different techniques also varied somewhat among wood, hardwood leaves, and conifer leaves; however, variations were minor relative to the overall trend. Equations developed can be used to extend data availability for developing and validating decomposition models.

Description: We studied radial growth reduction in American beech (Fagusgrandifolia Ehrh.) in relation to the level of defect induced by beech bark disease, in second-growth and old-growth northern hardwoods stands in New Hampshire. In the second-growth stand at Moose Mountain (n = 243 trees), 1989–1990 radial growth declined significantly with increasing severity of external symptoms. The severity of external symptoms increased significantly with DBH. To examine temporal trends, internal defect induced by the disease was quantified as the percentage of growth sheath cankered in each year, by cross-sectioning a subsample of 40 trees. Internal defect first appeared in 1950, increased through 1969, then declined until a major pulse of infection in the period 1983–1987. Sectioned trees were divided into infection classes based on a cumulative measure of internal defect. Growth of severely infected trees first fell below that of uninfected trees in 1965, and was consistently lower after 1972. The growth ratio of severely infected to uninfected trees generally declined from 1960 to 1990; by 1990, growth of severely infected trees was reduced by more than 40% relative to healthy trees. This decline in the growth ratio corresponded well to the increase in cumulative internal defect in the stand, suggesting that disease stress had cumulative effects on tree vigor. The relation between beech bark disease and growth was also examined on an individual-tree basis; recent growth decline was significantly greater for trees with higher levels of internal defect. Internal defect was a better predictor of growth trends than was external defect. External defect was only moderately correlated with internal defect (r2 = 0.503). In the old-growth stand at Bartlett, N.H. (n = 40 trees) infection was quantified from external symptoms only. As in the second-growth stand, the growth of severely infected trees in the old-growth stand fell significantly below that of uninfected trees. However, significant differences in growth between uninfected and severely infected trees occurred earlier in the old-growth stand, first appearing in 1949. Delayed growth reductions in the second-growth stand may be associated with changes in shade and moisture affecting the beech scale, changes in tree physiological stress after selective logging, or changes in the density of large trees. Beech may survive long periods of infection by beech bark disease. However, our results demonstrate clearly that beech bark disease has reduced the growth of American beech in both second-growth and old-growth northern hardwoods stands for several decades.

Description: The renewal of boreal fir stands after harvesting is related to the abundance of fir advanced regeneration. The objective of this study was to compare the advanced regeneration in 45-years-old balsam fir stands of second growth to determine if ecological site conditions could explain some regeneration problems noted in the balsam fir – white birch ecoclimatic domain. Seven ecological phases were studied; these are among the most common in the Laurentians north of Québec. On the basis of fir seedling densities (2 years old and more), three groups could be distinguished using a cluster analysis method for grouping means. Dry balsam fir – herb-and-moss type on well drained tills formed a first group characterized with very high seedling densities (〉 60 000 seedlings/ha). A second group, characterized with high seedling densities (25 000 – 40 000 seedlings/ha), was constituted of three phases with an important moss cover:the mesic balsam fir – moss-and-herb type on moderately well drained tills, the humid balsam fir – moss-and-herb type on imperfectly drained tills with seepage, and the humid balsam fir – moss type on imperfectly drained tills. A third group, with low seedling densities (

Description: Stratum-based timber harvest schedules must be disaggregated into operational plans prior to implementation. In most cases this is an expensive and time-consuming manual task that does not ensure consistency between the long-term harvest schedule and short-term operational activities. This paper presents the results of applying the CRYSTAL algorithm, which automates the disaggregation and allocation of a stratum-based harvest schedule into harvest blocks, to a small forest in New Brunswick. The results indicate that it is possible to use a set of allocation guidelines to quickly delineate harvest blocks in a consistent, reproducible manner. We also discuss how the algorithm is used in conjunction with a Monte Carlo integer programming model to estimate the potential losses in timber harvest volumes attributable to deviations from the stratum-based schedule and the addition of adjacency constraints.

Description: Limits are frequently encountered in the range of values of independent variables included in data sets used to develop individual tree mortality models. If the resulting model is to be utilized, its ability to extrapolate to conditions outside these limits must be evaluated. This paper describes the development and evaluation of six assumptions required to extend the range of applicability of an individual tree mortality model previously described. The assumptions deal with mortality in very dense stands, mortality for very small trees, mortality on habitat types and regions poorly represented in the data, and mortality for species poorly represented in the data.

Description: Thirty years of cone production records for subalpine fir (Abieslasiocarpa (Hook.) Nutt.) and mountain hemlock (Tsugamertensiana (Bong.) Carr.) (two sites each) in the Cascade Mountains of Washington and Oregon were compared with basal area increment and weather records to determine relationships among weather, radial growth, and cone crop. Results show that the size of subalpine fir cone crops was negatively related to large crops and positively related to radial growth in the previous 2 years. Mountain hemlock cone crops were negatively related o a large cone crop and positively related to July or August temperature in the previous year. Radial growth in heavy cone years was inhibited more for subalpine fir than for mountain hemlock. Results are explained by differences in the location of cone production between species. It is concluded that global climate warming could result in fewer and more irregular cone crops for these species.

Description: We describe a visual survey technique for evaluating acorn production. In contrast with previously proposed methods, our technique yields ratio-level data on annual productivity that are analyzable with standard statistics and, by sampling the same trees each year, data on the reproductive patterns of individual trees. We compared this technique with two independent sets of acorn-trap data acquired on oaks of three species at Hastings Reservation in central coastal California. Correlations between acorns counted by the visual surveys and collected from acorn traps under the same trees were significant for all three species. Most scatter in the data appeared to be attributable to three causes: (1) sampling error, especially among trees with very small crops, (2) finite counting speed, leading to a lack of discrimination among trees with very large crops by the visual surveys, and (3) arboreal acorn removal by animals. This latter factor can be particularly large, rendering visual surveys more reliable than the use of traps. Furthermore, only the high efficiency of visual surveys allows for the practical assessment of samples large enough to accommodate high within-population variation and detect widespread geographic variation in acorn production. Visual surveys offer a method of assessing the fruit or cone crops of many hardwood and conifer species that is not only more efficient but also more accurate than the use of traps.

Description: Geostatistics provides tools to model, estimate, map, and eventually predict spatial patterns of tree size and growth. Variogram models and kriged maps were used to study spatial dependence of stem diameter (DBH), basal area (BA), and 10-year periodic basal area increment (BAI) in an old-growth forest stand. Temporal variation of spatial patterns was evaluated by fitting spatial stochastic models at 10-year intervals, from 1920 to 1990. The study area was a naturally seeded stand of southwestern ponderosa pine (Pinusponderosa Dougl. ex Laws. var. scopulorum) where total BA and tree density have steadily increased over the last decades. Our objective was to determine if increased stand density simply reduced individual growth rates or if it also altered spatial interactions among trees. Despite increased crowding, stem size maintained the same type of spatial dependence from 1920 to 1990. An isotropic Gaussian variogram was the model of choice to represent spatial dependence at all times. Stem size was spatially autocorrelated over distances no greater than 30 m, a measure of average patch diameter in this forest ecosystem. Because patch diameter remained constant through time, tree density increased by increasing the number of pine groups, not their horizontal dimension. Spatial dependence of stem size (DBH and BA) was always much greater and decreased less through time than that of stem increment (BAI). Spatial dependence of BAI was close to zero in the most recent decade, indicating that growth rates in 1980–1990 varied regardless of mutual tree position. Increased tree crowding corresponded not only to lower average and variance of individual growth rates, but also to reduced spatial dependence of BAI. Because growth variation was less affected by intertree distance with greater local crowding, prediction of individual growth rates benefits from information on horizontal stand structure only if tree density does not exceed threshold values. Simulation models and area estimates of tree performance in old-growth forests may be improved by including geostatistical components to summarize ecological spatial dependence.

Description: Application of base-cation fertilizers has been shown to increase tree growth and vigour in declining sugar maple (Acersaccharum Marsh.) stands in southern Quebec but little is known about the effects of such fertilizers on litter quality or decomposition. Sugar maple foliage litters from fertilized and unfertilized plots on a base-poor site and from a naturally base-rich site were incubated in litterbags of 1- and 3-mm mesh sizes on fertilized and unfertilized plots at the base-poor site. Mass loss of unfertilized litter was slower in fertilized than in unfertilized plots, suggesting a negative effect of fertilization on the decomposer community. Faster mass loss of fertilized than unfertilized litter incubated in the same plot indicated that changes in litter quality brought about by fertilization enhanced decomposition. Mass loss of fertilized litter on fertilized plots did not differ from that of unfertilized litter on control plots, indicating that although decomposition processes are affected by fertilization the overall effect on decomposition is negligible. Mass loss was significantly, but only slightly, higher in large mesh than in small mesh bags indicating that larger soil fauna play a limited role in litter decomposition in this forest.

Description: Stand tables from forest inventories representing more than 22 × 106 ha of forests in tropical Asia were used to estimate aboveground biomass (point and 99% confidence interval). The mean inventory-based biomass for moist forests (225 Mg/ha) was lower than that reported by direct measurements for mature forests in the same region (350 Mg/ha), whereas the mean inventory-based biomass for dry forests (82 Mg/ha) was higher than estimates based on direct measurements (55 Mg/ha). Our analyses demonstrated that human use of the forests in tropical Asia is intense, leading to degradation. Between two national forest inventories of Peninsular Malaysia in 1972 and 1981, the total area and biomass of forests declined by 18 and 28%, respectively. Modeling land-use changes and carbon dynamics of tropical Asian forests must take into consideration human impact on vegetation because such use of forests reduces their biomass and may stimulate forest growth and carbon uptake.

Description: Measuring the mass of snow on cut branch tips soon after snowfalls during two winters provided comparisons of catch by Engelmann spruce (Piceaengelmannii Parry), subalpine fir (Abieslasiocarpa (Hook.) Nutt.), and lodgepole pine (Pinuscontorta var. latifolia Engelm.). Analysis of these and other reported measurements confirmed (i) snow bridging by cohesion, (ii) bouncing of snow crystals by elastic rebound, and (iii) branch bending as mechanisms that determine the sigmoidal growth curves characterizing snow interception relative to snowfall. The fraction of snowfall intercepted by the branches was largest when storm accumulations were 3–4 mm water equivalent, with low specific gravity (0.04–0.07). Percent catch in snowfalls with 10 mm water and low specific gravity was near 50% for Engelmann spruce and about 45% for subalpine fir and lodgepole pine, but values decreased to near 30% in 20-mm storms. Catch was inversely proportional to the density of snow accumulations in the specific gravity range 0.04–0.13. Average branch catch was only about 10% of a storm with 10 mm water equivalent at 0.13 specific gravity. Meteorological conditions were more important than branch growth form in determining snow interception amounts on the conifers tested. The results suggest, as a hypothesis, a computational function for the fraction of snowfall caught on conifer crowns.

Description: Identifying the factors that influence foresters to adopt management innovations is essential for developing models that accurately forecast the rate at which new technology will be adopted. Models were developed to evaluate the impact of sociodemographic characteristics, attitudes, and structural–external variables on New Hampshire foresters' decisions to adopt three distinct soil information sources. The soil information sources studied were County Soil Surveys, Important Forest Soils Groups, and Leak's Habitat Classification System. Discriminant analytic models were developed and evaluated for each information source. Information on use and other variables were collected by a mail survey with a response rate of approximately 60%. The results reveal that structural variables exerted the greatest influence on adoption decisions, while sociodemographic characteristics were the least important. College and noncollege sources of information about the innovations were the most important variables in the discriminant models. Other significant structural variables included employer, prior use of soils information, and participation in continuing education. All three models correctly classified more than 75% of the respondents, with one model classifying more than 90%. The difference in accuracy may be attributed to differences in information access among the three soils information sources.

Description: Radial growth patterns, canopy recruitment characteristics, and disturbance histories were examined in a shade-tolerant species, Nyssasylvatica Marsh., and a shade-intolerant species, Liriodendrontulipifera L., to determine the influence of canopy gaps in species with contrasting life histories. Tree cores of these co-occurring species were taken from three mixed-Quercus forests in northern Virginia. Most N. sylvatica individuals became established prior to 1850 and experienced multiple release and suppression periods coinciding with logging during the late 1800s and early 1900s. Many L. tulipifera became established during the early 1900s following logging, and only a few individuals experienced prolonged suppression periods. Regardless of site, L. tulipifera grew faster than N. sylvatica (average radial growth 〉1.70 mm/year for L. tulipifera vs. 2 mm (15–37 years for L. tulipifera vs.

Description: Western hemlock (Tsugaheterophylla (Raf.) Sarg.) seedlings from four dormancy induction treatments (DIT) (i.e., long-day dry, long-day wet, short-day dry, short-day wet) were planted on a coastal reforestation site in British Columbia and monitored for physiological response and morphological development over the first growing season. Seedlings in all DIT showed a similar net photosynthetic, needle conductance, and shoot water potential response to seasonal low temperature, non droughty, and late summer drought conditions. Pressure–volume analysis at the beginning of the growing season (late February) showed short-day wet seedlings to have lower saturated and turgor loss point osmotic potentials than other DIT, while at the end of the growing season (October) there was no osmotic potential difference between DIT. At the end of the growing season, short-day wet seedlings had a much lower maximum modulus of elasticity than other DIT, indicating their shoots were still elongating. Morphological assessment 1 month after field planting showed short-day DIT seedlings had less needle damage and greater root development in response to continuous low temperature exposure. Four months after planting, long-day wet seedlings had the greatest shoot (i.e., height, root collar diameter) and root development in response to mild temperatures and high soil moisture conditions. Eight months after planting, survival was between 95 and 97% for all DIT. Long-day DIT seedlings had the greatest height and all DIT had similar root collar diameter and root development. Short-day DIT seedlings had the best shoot/soil roots ratio. Short-day wet seedlings had the least number of stem units per centimetre of new shoot development throughout the growing season. Results are discussed in reference to stock quality assessment presented in the first paper of this series.

Description: The relevance of organosulfur formation and sulfate adsorption as S retention mechanisms in forest soils based upon the sulfur status of samples collected by horizon was investigated. Several forests of varying elevation, vegetation, location, and soil type were considered. Organic S was found to constitute over 78% of total S in the uppermost mineral (0–20 cm; A,E) horizons. This trend was also observed for both intermediate (20–40 cm; primarily A/B) and deeper (40+ cm; B,C) horizons, where organic S exceeded 65% of total S in all but one site examined. Adsorbed sulfate generally constituted only a minor component of the S pool in the uppermost mineral horizons (

Description: This paper reports results of a study designed to examine the control that soil temperature exerts on soil processes associated with nutrient flux, and in turn, on tree nutrition in interior Alaska black spruce ecosystems. Approximately 50 m2 of forest floor in a 140-year-old black spruce ecosystem, which had developed on permafrost, was heated to 8–10 °C above ambient temperature. This perturbation amounted to approximately a 1589 degree-day seasonal heat sum (above 0 °C), 1026 degree-days above the control total of 563 degree-days. The forest floor, surface 5 cm of mineral soil, and soil solution were compared with those of an adjacent control plot to evaluate the change in nutrient content and decomposition rate of the forest floor. The nutritional response to soil heating of current black spruce foliage also was evaluated. Soil heating significantly increased decomposition of the forest floor, principally because of an increase in biomass loss of the O21 layer. The increased decomposition resulted in greater extractable N and P concentrations in the forest floor, higher N concentrations in the soil solution, and elevated spruce needle N, P, and K concentrations for the experimental period. These results are discussed in light of the importance of soil temperature and other state factors that mediate ecosystem function.

Description: We test one of the fundamental assumptions of most dendroclimatological research, that the radial growth response of trees to climate does not vary with age once the biological growth trend has been removed. Piceaglauca (Moench) Voss from three sites in the western Northwest Territories, Canada, are disaggregated into age-classes, and their response to climate examined through response function and linear regression analyses. These data are then used in multiple regression analyses to estimate June-July temperatures at Norman Wells, N.W.T., from 1909 to 1989 using both age-dependent and standard (age-independent) models. The response function and regression analyses suggest that the response of Piceaglauca radial growth to climate differs between trees greater than 200 years old and less than 200 years old. These results suggest that the assumption of an age-invariate climate–growth function is therefore invalid at these sites. These apparent age-dependent responses are site specific and may reflect physiological changes related to aging. One possible causal factor of age dependence is that the trees are becoming increasingly moisture stressed with age owing to a reduction in the efficiency of water and nutrient translocation mechanisms. The estimation of June–July temperatures based on an age-dependent model produced improved calibration and verification statistics as compared with a reconstruction based on standard dendroclimatic modelling. If present, age-dependent climate–growth relationships may result in less accurate reconstructions of past climate, particularly during the early portions. However, age-dependent responses could also be used to increase the number of tree-ring based predictor variables for dendroclimatic reconstructions.

Description: The reactions of sessile oak (Quercuspetraea (Mattuschka) Liebl.) to drought were studied under natural conditions in a 32-year-old stand near Nancy (northeastern France) during the summers of 1989 (strongly rain deficient) and 1990. A plot of five trees was subjected to imposed water shortage, while a group of irrigated trees was used as a control. Measurements of xylem sap flows and water potential enabled the computation of plot transpiration, canopy conductance, and specific hydraulic conductance in the soil–tree continuum. Stomatal conductance was measured directly with a porometer. Specific hydraulic conductance of our oaks was of the same order of magnitude as that reported for other species. It decreased significantly during spring because of a time lag between cambial growth and leaf area expansion. Measured transpiration was close to potential evapotranspiration, except during days with high vapor pressure deficits, which promoted stomatal closure in the absence of soil water deficits. Imposed drought caused predawn leaf water potentials to reach values as low as −2.0 MPa and a progressive decline in hydraulic conductance, which was probably attributable to modifications in hydraulic properties at the soil–root interface. This gradual decline in conductance was attributed to their deep rooting (1.40 m). This study revealed that Q. petraea may be considered as drought tolerant because of adaptations like deep rooting, efficient and safe xylem sap transport, maintenance of significant stomatal conductance, and significant transpiration, even during strong drought stress.

Description: Coarse woody debris from streamside forests plays important biological and physical roles in stream ecosystems. The distance from stream bank to rooting site was determined for at least 30 fallen trees at each study site on 39 streams in the Cascade and Coast ranges of Oregon and Washington. The study sites varied in channel size (first- through third-order), side-slope steepness (3 to 40°), and age of surrounding forest (mature or old-growth stands). The distribution of distances from rooting site to bank was similar among streams, with 11% of the total number of debris pieces originating within 1 m of the channel and over 70% originating within 20 m. Stands with taller trees (old-growth conifers) contributed coarse woody debris to streams from greater distances than did stands with shorter (mature) trees.

Description: This review presents information on the occurrence, persistence and nontarget effects of Bacillusthuringiensis Berliner (B.t.) in soil, with particular emphasis on forest ecosystems. Both field and laboratory studies have confirmed that B.t. is able to survive for several years after spraying, but studies on long-term persistence and possible accumulation of spores in soils are inadequate. The ecological role of B.t. in the soil ecosystem is poorly understood; we do not even know where and under what conditions B.t. multiplies in nature. Information on the effects of B.t. on soil microflora is contradictory, with studies variously suggesting that B.t. caused an increase, a decrease, or did not affect indigenous bacterial populations. Toxicity of B.t. to species of several invertebrate taxa (Acarina, Nematoda, Collembola, Annelida, Hymenoptera) inhabiting the soil has been demonstrated, but only rarely is it possible to relate dosage information to field situations, and in many cases the B.t. subspecies tested are not currently used for pest control in North America. There is an urgent need for further research to elucidate the relationships between B.t. and the natural soil microflora and fauna.

Description: Seedlings of Douglas-fir (Pseudotsugamenziesii (Mirb.) Franco), lodgepole pine (Pinuscontorta Dougl.), and white spruce (Piceaglauca (Moench) Voss) were grown in a container nursery from February to July 1988 and then exposed to three temperatures and three levels of drought stress applied factorially during mid-July to October 1988. Seedlings were retained in a shelter house until January 1989, when they were cold-stored until early May. Measurements of stomatal conductance (gs), transpiration (E), and specific leaf area (SLA) were made at the end of the treatment period in September 1988 and again after growth the following year at the end of June. Root growth capacity (RGC) was tested in early May 1989. Results were considered in conjunction with performance of other samples of the same plants that had been planted in sand beds in April 1989, where irrigation was regulated to provide three levels of moisture stress. Low temperature (13 °C) generally reduced gs and E, which were adjusted for xylem pressure potential, and SLA in all species by the time nursery treatment was completed at the end of September. No effect of nursery temperature treatment on gs and E could be detected when new needles were measured in June and July (after 9 to 12 weeks of growth), but SLA of lodgepole pine increased with nursery temperature treatment, and SLA of white spruce decreased with nursery temperature treatment. RGC was higher for the 13 °C treatment than for the 16 and 20 °C treatments. Survival of outplanted seedlings was mainly inversely related to nursery temperature. Low nursery temperature reduced gs, E, and SLA and increased RGC. SLA of planted lodgepole pine increased with level of nursery drought treatment, and severe nursery drought increased gs under stress, when measured in June. No other effects of drought were detected, although drought treatment was effective in increasing survival of planted seedlings.

Description: Development of humus layer soil microbial biomass C (Cmic) and N (Nmic), fungal biomass (as soil ergosterol content), microbial respiration activity, and the soil organic C (Corg) and N (Ntot) were determined in coniferous forest soils that had received a single prescribed fire treatment at different times over a period of 45 years. The ratio of soil respiration rate to microbial biomass C (qCO2) and the Cmic/Corg and Nmic/Ntot percentages were derived from the measurements taken. All the measured biomass indicators reacted identically to show recovery from prescribed burning within 12 years. A raised metabolic quotient (qCO2) was detected in soils over the first 2 years following the fire treatment, but after the third year it had decreased to a stable level. These observations suggest that during the first few years after fire the soil microflora can be characterized on the basis of simple substrate–decomposer relationships. The first 12 years were characterized by increasing Cmic/Corg and Nmic/Ntot percentages, which then stabilized at mean values of 1.3 and 5.5%, respectively. The observed rise in the Cmic within a large pool of Corg suggested increasing availability of energy-rich C sources. These C sources are probably derived from the organic C input resulting from postfire plant succession.

Description: During a 3-year study, soil microbial biomass C and N, length of the fungal hyphae, soil respiration, and the percent mass loss of needle litter were recorded in coniferous forest soil humus layers following a prescribed burning (PB) treatment or a forest fire simulation (FF) treatment (five plots per treatment). Unburned humus from adjacent plots served as controls (PC and FC, respectively). Prescribed burning was more intensive than the forest fire, and this was reflected in all the measurements taken. The amounts of microbial biomass C and N, length of fungal hyphae, and soil respiration in the PB area did not recover to their controls levels, whereas unchanged microbial biomass N and recovery of the length of the fungal hyphae to control levels were observed in the FF area. The mean microbial C/N ratio was approximately 7 in all the areas, which reflected the C/N ratio of the soil microbial community. Deviation from this mean value, as observed during the first three samplings from the PB area (3, 18, and 35 days after fire treatment), suggested a change in the composition of the microbial community. Of the two treated areas, the decrease in soil respiration (laboratory measurements) was much more pronounced in the PB area. However, when the humus samples from both areas were adjusted to 60% water holding capacity, no differences in respiration capacity were observed. The drier humus, due to higher soil temperatures, of the PB area is a likely explanation for the low soil respiration. Lower soil respiration was not reflected in lower litter decomposition rates of the PB area, since there was a significantly higher needle litter mass loss during the first year in the PB area followed by a decline to the control level during the second year. Consistently higher mass losses were recorded in the FC area than in the FF area.

Description: The reliability of a leaf-disk assay to assess resistance of Populusdeltoides Bartr. to Melampsoramedusae Thuëm. f.sp. deltoidae, the causal agent of poplar leaf rust, was evaluated. Leaf disks of eight host genotypes were inoculated in a spore settling tower with 11 isolates of the pathogen in all possible combinations, and the latent period, infection probability, sporulation, and progeny/parent ratio were recorded. Correlations were established between these measures and measures derived from field epidemics, namely the relative area under the disease progress curve, the apparent rate of infection, the final disease severity, and the number of days before defoliation. Four genotypes were highly resistant to all 11 isolates tested with the leaf-disk assay and to the local inoculum in field tests. Three genotypes were highly susceptible in inoculation assays and were also susceptible in the field. One genotype was highly resistant in inoculation assays but had intermediate resistance in the field. Significant correlations (p 〈 0.05) were found between all leaf-disk and all field parameters.

Description: Red spruce (Picearubens Sarg.), black spruce (Piceamariana (Mill.) B.S.P.), and balsam fir (Abiesbalsamea (L.) Mill.) seedlings were root inoculated with tomato mosaic virus, potted, and maintained in a cold frame. The virus was detected by enzyme-linked immunosorbent assay in root extracts of several inoculated seedlings of each species 6–12 months postinoculation, and in root extracts of approximately 35–40% of both inoculated and noninoculated seedlings of each species 12–18 months postinoculation. Virus spread apparently occurred from the roots of infected to noninfected seedlings within the cold frame. The presence of virus in root extracts was confirmed by immunoelectron microscopy. The virus was not detected in the needles of any seedling at any time by either enzyme-linked immunosorbent assay or immunoelectron microscopy. A transitory needle chlorosis was observed in approximately 14% of the inoculated black spruce and 30% of the inoculated red spruce seedlings, but tomato mosaic virus was not detected in all plants with symptoms.

Description: Rising atmospheric carbon dioxide concentrations may have important consequences for forest ecosystems. We studied above- and below-ground growth and leaf gas exchange responses of Populusgrandidentata Michx. to elevated CO2 under natural forest conditions over the course of a growing season. Recently emerged P. grandidentata seedlings were grown in native, nutrient-poor soils at ambient and twice ambient (707 μbar (1 bar = 100 kPa)) CO2 partial pressure for 70 days in open-top chambers in northern lower Michigan. Total leaf area and shoot and root dry weight all increased in high CO2 grown plants. Photosynthetic light and CO2 response characteristics were measured 28, 45, and 68 days after exposure to elevated CO2. In ambient grown plants, light saturated assimilation rates increased from day 28 to day 45 and then declined at day 68 (15 September). This late-season decline, typical of senescing Populus leaves, was due both to a decrease in the initial slope of the net CO2 assimilation versus intercellular CO2 partial pressure relationship and to decreased CO2 saturated assimilation rates. Specific leaf nitrogen (mg N•(cm2 leaf area)−1) did not change during this period, although leaf carbon content and leaf weight (mg•cm−2) both increased. In ambient grown plants stomatal conductance also declined at day 68. In contrast, plants grown at elevated CO2 showed no late-season decline in photosynthetic capacity or changes in leaf weight, suggesting a delay in senescence with long-term exposure to high CO2. High CO2 grown plants also maintained photosynthetic sensitivity to increasing Ci throughout the exposure period, while ambient CO2 grown plants were insensitive to Ci above 400 μbar on day 68. These results indicate the potential for direct CO2 fertilization of P. grandidentata in the field and provide evidence for a new mechanism by which elevated atmospheric CO2 could influence seasonal carbon gain.

Description: Nitrogen (N) mineralization and nitrification were compared among ecosystems representing a primary successional sequence on the Tanana River floodplain of interior Alaska. These processes displayed marked seasonality, were closely related to substrate chemistry, and reflected the impact of vegetation clearing. The highest rates of N mineralization were encountered in the June to July incubation periods, and rates generally declined during the remainder of the summer. The early season period (June to July) was the interval of most favorable litter and mineral soil temperature and most available energy supply for microbial mineralization of detrital materials. Minimal rates were encountered during the winter. Litter layer N mineralization rates were highest in the early-successional poplar–alder (Populusbalsamifera–Alnustenuifolia (Nutt.) stage and declined with advancing succession in poplar (Populusbalsamifera) and mature white spruce (Piceaglauca) (Moench) Voss) stands. The poplar–alder stage displayed the highest rate of nitrification. Nitrate constituted 98% of the mineralized N in early-successional poplar–alder forest floors but fell to 4 and 0% in poplar and white spruce forest floors, respectively. Nitrogen mineralization was closely related to significant increases in the lignin/N ratio across the sequence of vegetation types. The rate of surface mineral soil net N mineralization increased with succession in response to higher soil organic matter content. The range of average total seasonal net N mineralization (260–1600 mg N•m−2) for litter layer plus mineral soil among successional stages in this study was generally lower than the 1200–8400 mg N•m−2 reported by investigators for other studies in temperate latitudes. Vegetation clearing increased the magnitude of temporal fluxes as well as total annual mineral N production. The most consistent increases were encountered in the poplar–alder vegetation type. The average seasonal total net N mineralization for forest floor plus mineral soil in this vegetation type increased from 1500 to 3264 mg N•m−2 as a result of clearing. Soil temperature declined with advancing succession and generally increased as a consequence of clearing. However, these changes were not as closely correlated with N mineralization as were the changes in substrate chemistry encountered across this successional sequence.

Description: A combination of antiaggregation and aggregation pheromones was tested for protecting stands of Douglas-fir (Pseudotsugamenziesii (Mirb.) Franco) at high risk for infestation by the Douglas-fir beetle (Dendroctonuspseudotsugae Hopkins). The antiaggregation pheromone, 3-methylcyclohex-2-en-1-one (MCH), was applied in a bubble capsule formulation to the perimeter of 1-ha circular plots at a rate of 60 g/plot. Treated plots also had three or four clusters of four Lindgren funnel traps baited with frontalin, seudenol, 1-methylcyclohex-2-en-1-ol, and ethanol located outside of the plot but within 160 m of the boundary. Mean (±SE) accumulated catches in all traps per plot were 73 658 ± 19 721 Douglas-fir beetles and 12 892 ± 2 513 Thanasimusundatulus (Say), a predator of the Douglas-fir beetle. The mean percentage of Douglas-fir trees ≥20 cm DBH that were mass attacked was reduced by 80% within the treated plots compared with the untreated plots. However, there was an eightfold increase in the percentage of trees mass attacked in the area outside the treated plots in the vicinity of the funnel traps. The net effect of the treatment was to concentrate mass-attacked trees within a limited area outside of the protected stand. Our results indicate that Douglas-fir beetle antiaggregation and aggregation pheromones can be used effectively to reduce the probability of infestation in small, high-value stands.

Description: Variation in diameter, height, and stem volume of 57-year-old Douglas-fir (Pseudotsugamenziesii var. menziesii (Mirb.) Franco) was related to distance of these trees from a 27 m wide strip in the same Douglas-fir plantation that had been interplanted with red alder (Alnusrubra Bong.). Within the interplanted strip and despite its greater total stand density, bole volume of dominant and codominant Douglas-fir averaged 1.27 m3 compared with 0.55, 0.45, 0.46, or 0.49 m3 in trees 15, 30,45, or 60 m, respectively, from the edge of the mixed stand. Some positive influence of nitrogen-fixing red alder apparently extended about 15 m beyond the edge of the mixed stand at this poor quality site in southwest Washington. We infer that similar ribbonlike distributions of naturally regenerated red alder could be retained to improve growth of nearby conifers on nitrogen-deficient sites.

Description: Subalpine fir (Abieslasiocarpa (Hook.) Nutt.) regeneration following fire was studied at two locations that burned in 1902 northeast of Mount Rainier, Washington. Tree establishment dates were compared with local climatic records using multiple and logistic regression to identify potential relationships between seasonal climate and annual tree establishment. The influence of microsite features on forest regeneration was also explored. Little regeneration occurred in the first 30 years after the fires, and most trees established in the 1950s, 1977, 1983, and 1989. The dominance of trees

Description: Attraction of scolytids and other bark- and wood-dwelling beetles to volatile constituents of Norway spruce (Piceaabies (L.) Karst.) was studied in field experiments in central Sweden. The volatiles were released from chips of newly cut spruce stems as well as from stems stored over winter. These host materials were kept in cylinders covered on both ends with fine nylon mesh. Attracted insects were caught in barrier traps. The composition of volatile constituents of wood samples was determined using gas chromatography. Hylurgopspalliatus (Gyll.), Hylastescunicularius Er., Ipstypographus (L.) (Scolytidae), and Glischrochilusquadripunctatus (L.) (Nitidulidae) were particularly attracted to stored spruce wood. Trypodendronlineatum (Oliv.) (Scolytidae) and Pityophagusferrugineus (L.) (Nitidulidae) were only attracted to stored wood. There were indications that Tomicuspiniperda (L.) and Hylastesbrunneus Er. (Scolytidae) preferred fresh wood. Chemical analyses revealed that during storage ethanol and acetaldehyde increased considerably in two out of the five examined tree stems.

Description: Rooted cuttings of three clones of Sitka spruce (Piceasitchensis (Bong.) Carr.) were grown in 2 m tall transparent acrylic tubes of peat in insulated boxes out of doors. Roots of many of the trees formed mycorrhizas with Thelephoraterrestris Ehrh.:Fr.; the extra-matrical mycelium and associated strands were visible in the tubes, together with the roots. Some of the tubes were waterlogged to submerge the lower part of the root and mycelial systems. Waterlogging was carried out in October, when roots were growing slowly, or in November, when growth had stopped. The fungal mycelium was growing on both occasions. The tubes were drained in the following March, and survival of main roots and fungus was measured to the point of regeneration. There were large differences between the two waterlogging treatments in root survival. October waterlogging caused substantial dieback, and roots survived to a mean depth of only 122 mm below the water table. November waterlogging resulted in little dieback, and survival depth was 308 mm. Differences between clones in root survival were statistically significant but small. The extra-matrical hyphae of Tterrestris died in both treatments, but all of the strand systems survived and regenerated in the following spring. Factors affecting survival are discussed, including growth and oxygen transport in roots and fungal strands.

Description: The net annual flux of carbon from south and southeast Asia as a result of changes in the area of forests was calculated for the period 1850 to 1985. The total net flux ranged from 14.4 to 24.0 Pg of carbon, depending on the estimates of biomass used in the calculations. High estimates of biomass, based on direct measurement of a few stands, and low estimates of biomass, based on volumes of merchantable wood surveyed over large areas, differ by a factor of almost 2. These and previous estimates of the release of carbon from terrestrial ecosystems to the atmosphere have been based on changes in the area of forests, or rates of deforestation. Recent studies have shown, however, that the loss of carbon from forests in tropical Asia is greater than would be expected on the basis of deforestation alone. This loss of carbon from within forests (degradation) also releases carbon to the atmosphere when the products removed from the forest burn or decay. Thus, degradation should be included in analyses of the net flux of carbon from terrestrial ecosystems. Degradation may also explain some of the difference between estimates of tropical forest biomass if the higher estimates are based on undisturbed forests and the lower estimates are more representative of the region. The implication of degradation for estimates of the release of carbon from terrestrial ecosystems is explored. When degradation was included in the analyses, the net flux of carbon between 1850 and 1985 was 30.2 Pg of carbon, about 25% above that calculated on the basis of deforestation alone (with high estimates of biomass), and about 110% above that calculated with low estimates of biomass. Thus, lower estimates of biomass for contemporary tropical forests do not necessarily result in lower estimates of flux.

Description: One-year-old Cornusflorida L. seedlings were randomly assigned to four treatments of simulated rain (pH 2.5, 3.5, 4.5, and 5.5) in two greenhouse–laboratory studies in 1989 and 1990. After 10 applications over a 42-day period, the seedlings were moved to a temperature-controlled laboratory, placed in plastic bags, humidified, and sprayed with a spore suspension of five Disculadestructiva isolates. About 30 days later, the seedlings were examined for percentage of leaves exhibiting anthracnose symptoms and disease severity on affected leaves. Both trials showed that as the acidity of the simulated rain increased, the incidence and severity of anthracnose leaf symptoms increased. The 1989 study included a soil lime treatment that showed the same trend but the overall occurrence and severity of symptoms was higher.

Description: This study presents an individual tree height prediction model for white spruce (Piceaglauca (Moench) Voss) and trembling aspen (Populustremuloides Michx.) grown in boreal mixed-species stands in Alberta. The model is based on a three-parameter Chapman–Richards function fitted to data from 164 permanent sample plots using the parameter prediction method. It is age independent and expresses tree height as a function of tree diameter, tree basal area, stand density, species composition, site productivity, and stand average diameter. This height-prediction model was fitted by weighted nonlinear regression for spruce and unweighted nonlinear regression for aspen. Almost all estimates of parameters were significant at α = 0.05 and model R2-values were high (0.9192 for white spruce and 0.9087 for aspen). No consistent underestimate or overestimate of tree heights was evident in plots of studentized residuals against predicted heights. The model was also tested on an independent data set representing the population on which the model was to be used. Results showed that the average prediction biases were not significant at α = 0.05 for either species, indicating that the model appropriately described the data and performed well when predictions were made.

Description: Beech bark disease occurs when either Nectriagalligena Bres., or Nectriacoccinea var. faginata Lohman, Watson, and Ayers kills bark that is or has been infested and altered by the beech scale, Cryptococcusfagisuga (Lind.). Introduced to Nova Scotia around 1890, this insect now occurs as far southwest as Ohio, West Virginia, Virginia, and North Carolina. I determined the relative occurrence of the two pathogens in forests affected for varying times and the temporal changes in their relative occurrence in recently affected stands of the Monongahela National Forest, West Virginia (MNF). Stands in the Canadian Maritimes, New England, New York, and Pennsylvania were sampled in 1985–1986; and in the Catskill Park, New York in 1988 and 1991. Stands on the MNF were sampled in 1982, 1983, 1984, 1986, 1988, and 1991. The following trends were indicated: N. galligena dominated recently affected stands in Pennsylvania and West Virginia, and N. coccinea var. faginata dominated long-affected stands in Canada, New England, and New York. On the MNF, only N. galligena was isolated in 1982; by 1988, N. coccinea var. faginata was isolated from 8 of 16 stands and from all 16 stands in 1991. Stands where N. galligena persisted had higher proportions of tree species highly susceptible to N. galligena than stands dominated quickly by N. coccinea var. faginata. Once present, however, N. coccinea var. faginata eventually will replace N. galligena as the dominant pathogen.

Description: The prevalence of individual-tree growth decline was determined for red spruce (Picearubens Sarg.) populations at three locations in the southern Appalachians: Mount Rogers National Recreation Area, the Black Mountains, and Great Smoky Mountain National Park. An index of annual stemwood volume increment (AVI) was computed from dendrochronological data and a site-specific DBH–height regression equation. Individual-tree AVI time series were analyzed to identify changes in 20-year periodic mean AVI and AVI trend. The proportion of red spruce that exhibited decreasing mean AVI or negative AVI trend was determined for the most recent 20-year period, and this was compared with the estimated historical prevalence of these indications of growth decline. Also, the prevalence of growth decline was compared among subpopulations that differed with regard to various tree, stand, and site characteristics. Of 263 red spruce sampled, 25% exhibited a decrease in mean AVI during the period 1967–1986, 8% exhibited a negative AVI trend without a reduction in mean AVI, and 17% exhibited a reduction in the slope of the AVI curve. The proportion of trees that exhibited decreasing or slowed growth after 1967 was substantially greater among trees growing at 1980 m than in populations at lower elevations; no relationship was found between elevation and growth decline below 1980 m. No difference was found in prevalence of growth decline between subpopulations that differed with regard to age, DBH, competitive status, stand density, slope aspect, or site exposure. The prevalence of individual-tree growth decline for the most recent 20-year period did not exceed estimated levels for historical periods of decline in the Great Smoky Mountains population.

Description: Phenotypic variation in defoliation of ponderosa pine, Pinusponderosa Laws., by the pine sawfly, Neodiprionfulviceps, is reported. Two field and one laboratory bioassays of paired phenotypically resistant and susceptible trees consistently indicated that sawfly egg, larval, and total survival were 5–14% lower on resistant trees. A suite of host-plant traits were assayed on resistant and susceptible trees including anatomical characteristics of the needles and concentration of terpene and nitrogen. Resistant trees had thicker outer cell layers, more fiber cells, higher toughness, and lower soluble nitrogen than susceptible trees. Resistant and susceptible trees were not distinguished by differences in terpenes. Generational survival of N. fulviceps was inversely related to foliage toughness and number of fiber cells.

Description: We classified red spruce (Picearubens Sarg.) sites from northern Maine by radial growth release history. Two major releases were apparent for a majority of the sites. The first was a reduction and subsequent increase in radial increment in 1920. The second was an increase in radial increment from 1935 to 1955. Red spruce radial growth reduction in the 1960s is apparent only for sites released from 1935 to 1955 (approximately 54% of the sites in this study). These sites are now approaching the radial growth rates of the unreleased stands. Birch dieback is suggested as a probable contributor to the 1935–1955 red spruce growth increase and subsequent 1960s growth reduction.

Description: In a routine determination of jack pine (Pinusbanksiana Lamb.) tree ages, several rings were overlooked or were difficult to count. Sections of the wood, when examined at higher magnification, revealed an unusually high proportion of "light rings." Light rings are characterized by having a small number of latewood cells whose walls are not as thick as those of latewood cells found in normal rings. Under low magnification, bands of these light rings may be interpreted as a single annual ring. Light rings may thus be a source of error in determining tree ages for forest productivity studies, particularly in older stands that have been affected by defoliators and root disease.

Description: A base-age invariant site index equation for jack pine based on the Chapman–Richards function was produced that satisfied nine criteria of preferred behavior for site index equations. A difference form of the Chapman–Richards equation produced the best behavior; height equalled site index at base age, and the shape of the curves reflected the data. The data structure used to fit the difference equation was all possible differences rather than the conventional nonoverlapping sequential intervals because this improved the behavior of the model. Height-prediction equations typically use height at base age (site index) as a predictor variable. As site index is measured with error, the equation will be biased. This bias will be evident in the predicted height at base age and in the shape of the curves. Base-age invariant equations predict height and site index with the same equation and thus diminish the effect of stochastic predictor variables. The equation performed comparably to a previously published equation with a specific base age of 50 years.

Description: Measurements of the spatial mean values of global irradiance, photosynthetic photon flux density, and the downward longwave irradiance under a 26-year-old, second-growth Douglas-fir (Pseudotsugamenziesii (Mirb.) Franco) forest canopy on a 26° south-facing slope were made with a tramway system, which scanned a 20-m transect in 12 min. The diffuse solar irradiance under the canopy was measured with a stationary pyranometer equipped with a shadow band. The extinction coefficients for the direct, global, and photon components were derived as functions of the solar incident angle over the range of 15° to 85°. The extinction coefficient for the diffuse radiation was found to correlate well with the ratio of the direct to diffuse irradiance above the canopy. Complete diurnal cycles of the downward longwave irradiance were simulated with a simple model based on the air temperature inside the stand. Analyses of the measurements of all the shortwave and longwave components were made using an effective leaf area index, which was derived from the measurements of the direct irradiance above and below the stand. It was found that the distribution of the leaf inclination angle of a Douglas-fir canopy has strong planophile characteristics, and that in the case of a forest stand on a slope, it is critical to obtain the characteristics of the light transmission through the canopy over the entire incident angle range before effective leaf area index is calculated. Warren Wilson's 57.5° approximation did not hold for the Douglas-fir canopy, which had distinct foliage clumping features.

Description: The problem of fitting height–diameter curves for repeated measurements on growth plots is addressed. The context of the problem is fitting historical data with varying sampling protocols and varying measurement accuracy. A key consideration is obtaining good estimates of top height and top-height increment. A particular model and objective function for fitting are presented. The model has two parameters for each measurement and one common parameter; limited crossovers in the height–diameter curves for the various measurements are allowed. The objective function minimizes errors in predicted height and in predicted change in height. The programming is described, and the availability of code is announced. Examples show both the strengths and weaknesses of this approach.

Description: In a Rocky Mountain aspen forest, the detailed pattern of mass loss from decomposing leaf litter of trembling aspen (Populustremuloides Michx.) during the first 6 months of decay was compared with that from aspen leaves modified to produce a more recalcitrant litter type by removal of leachable material (31.7% of original mass). Leaching litter removed substantial quantities of N (24%) and P (54%), but did not change the litter's C/N ratio (77:1); and leached leaves still contained 33% labile (benzene alcohol soluble) material. Decomposition of intact aspen litter was best described by a double exponential model (k1 = −7.91/year, k2 = −0.21/year), except during the first 2 weeks, when an extremely rapid mass loss (14.2%) apparently resulted from leaching. Microbial metabolism was probably responsible for most of the subsequent decay (35% total in 6 months). In contrast, decomposition of leached aspen showed no exponential trend and was best described by a simple linear regression with a slope of −19.7%/year. Additional data from a 2nd year (12–15 months decay) reduced the regression estimates of decay rates but did not alter the best fit models. Fits were improved slightly if temperature sum replaced time in the regressions, especially if 2nd-year data were included.

Description: Reciprocal cross pollinations between western larch (Larixoccidentalis Nutt.) and alpine larch (L. lyallii Parl.) were done in Spring, 1991. The cross to alpine larch females was highly successful; 63% of the seeds developed mature embryos and 79% germinated. However, the cross to western larch females resulted in only 4% filled seed of which 68% germinated. Open-pollinated western larch and alpine larch averaged 26 and 32% filled seed with 48 and 44% germination, respectively. Less than 1% of the alpine larch hypocotyls were reddish in color; most of them were green. About 15% of the western larch and about 17% of the hybrids from alpine larch females had reddish hypocotyls, whereas hybrids from western larch females were intermediate between western larch and alpine larch. Hybrid seed began germinating before the parental types and initial height growth exceeded that of the parental types. However, 4 months following germination, western larch seedlings were ca. four times taller than the hybrids and 10 times taller than alpine larch. Stems of hybrids were significantly thicker than those of either western larch or alpine larch. Hybrid seedlings are robust, stocky, and may be useful in revegetating cold, moist sites between the elevational ranges of alpine and western larch.

Description: Height and diameter growth, stem volume production, leaf phenology and leaf number, and number of branches of Populustrichocarpa Torr. & Gray, Populusdeltoides Bartr., and their F1 hybrids (P. trichocarpa × P. deltoides) were studied for 4 years in a research plantation in western Washington, United States. Twelve clones (three of each species and six of the hybrids) grew under a short-rotation silviculture regime in monoclonal plots at spacings of 1 × 1 m (10 000 stems/ha). Clones represented a north-south gradient within the geographic distribution of both the two North American poplar species and the parentage of the hybrid material. The results support earlier work by contributing additional evidence for the superiority of the hybrids. However, the relative hybrid superiority in these monoclonal plots was less pronounced than that found earlier in field trials with single-tree plots because of heightened intraclonal competition. After 4 years, mean estimated stem volume of the hybrids was 1.5 times that of P. trichocarpa and 2.3 times that of P. deltoides. Total tree height of the hybrids was 1.1 times that off. trichocarpa and 1.3 times that off. deltoides. Clonal variation was the dominant theme in height and diameter growth, stem volume productivity, time of bud break and bud set, tree mortality, and number of branches. Populustrichocarpa had the highest number of sylleptic branches, P. deltoides had the lowest, and hybrids were intermediate. Significant clone by replicate interactions were observed in height, diameter, and volume growth. Phenological traits, such as the dates of bud break and bud set, and the length of growing period only partly explained the observed differences in growth between the P. trichocarpa × P. deltoides hybrids and the parental species.

Description: Nitrogen transformations were examined in two pairs of adjacent, 55-year-old forests dominated by conifers (primarily Douglas-fir (Pseudotsugamenziesii (Mirb.) Franco)) and by conifers and nitrogen-fixing red alder (Alnusrubra Bong.). Nitrogen availability was examined in aerobic and anaerobic incubations in the laboratory, and with resin-core and buried-bag incubations in the field. Rates of nitrogen mineralization and immobilization were examined in the field incubations using [15N]ammonium. Net nitrogen mineralization in a year-long series of resin-core incubations of forest floor plus 0–0.15 m depth soil was about 9 kmol•ha−1•year−1 for the two alder–conifer stands, but almost nil in the conifer stand at the low-productivity Wind River site, and 2.1 kmol•ha−1•year−1 in the conifer stand at the high-productivity Cascade Head site. The 15N pool dilution experiment showed that buried-bag incubations demonstrated more differences among stands than did resin cores; resin cores typically gave greater rates than buried bags. Previous estimates of nitrogen budgets were coupled with net mineralization estimates to examine how well the estimated fluxes balanced at an ecosystem scale. This tabulation of the complete nitrogen cycles showed substantial discrepancies, prompting caution in interpretation of some of the differences among the stands.

Description: Nitrification and denitrification were measured in a 120-year-old Piceaglauca (Moench) Voss forest in southwestern Alberta. Nitrifying activity could not be detected using short-term incubations of F–H and upper mineral soil horizons. Long-term incubations for nitrifying potential indicated that nitrifying organisms were present. The results suggest that the population of nitrifying organisms must have been small, their activity was limited, or the microhabitat necessary for their activity was not adequately simulated by the soil slurries in the short-term incubations. Low rates of denitrification were detected but probably did not represent a substantial loss of N from the soil–plant system. Low rates of denitrification may have been due to a lack of substrate (NO3−), reflecting the low rates of nitrification.

Description: The nutritional requirements of Eucalyptusgrandis Hill ex Maiden seedlings were studied in glasshouse pot experiments. Nitrogen (N), phosphorus (P), and potassium (K) demand was assessed by bioassays, in which the uptake of tracers (15N, 32P, or 86Rb) by roots excised from the seedlings grown at different levels of nutrient supply was measured. A pilot study showed that there was an inverse relationship between nutrient supply and influx. The measured influx was affected by the length of time that the roots were stored before the bioassay was applied. In a 33 N, P, and K factorial nutrient experiment, growth was primarily influenced by N. Maximum growth occurred at the highest level of supply of each of the three nutrients. N and K influx in the root bioassay was inversely related to foliar N and K concentration, but P uptake was not related to foliar P concentration. These results indicate that the root bioassays are more sensitive determinants of nutrient limitation than. foliar analysis. Interactions between N and P and between N and K alter the stem:leaf ratio, which may be of importance in optimizing wood production.

Description: The effects of DiPel® 176, a commercially available Bacillusthuringiensis Berliner subsp. kurstaki (B.t.k.) formulation, on microbially mediated carbon and nitrogen mineralization processes, and the persistence of B.t.k. following application of DiPel® 176 to an acidic, coniferous forest soil were evaluated in the laboratory using simple microcosms. Litter (L) and fermentation–humus (FH) material were exposed to DiPel® at the recommended field application rate (FA), DiPel® at 1000× the field application rate (1000× FA), or left untreated. Respiration, substrate induced respiration (SIR), microbial biomass C, metabolic quotients (qCO2), NH4-N, NO3-N, cellulose decay, and B.t.k. viability were monitored regularly over 8 weeks. The FA treatment had no significant impact on soil processes in either the L or FH. The 1000× FA treatment increased SIR and biomass C and decreased qCO2 consistently in both the L and FH. No other effects of the 1000× FA treatment were evident in the L, while in the FH this treatment stimulated respiration initially, then reduced it below control levels; it enhanced cellulose decay; and it inhibited ammonification and nitrification after 8 weeks incubation. In both the L and FH there was no significant loss in viability of B.t.k. in either of the DiPel® treatments over 8 weeks. The microcosms used in this study were simple, inexpensive, and effective, with respiration, SIR, biomass C, and qCO2 being the least variable measurements and the most sensitive to perturbation. This approach is recommended for ecotoxicological and fate testing as outlined in the GuidelinesforRegistrationofNaturallyOccurringMicrobialPestControlAgents.

Description: Microbial isolates from conifer foliage and strawberry were evaluated for biocontrol of Botrytiscinerea Pers.:Fr. in container-grown seedlings of black spruce (Piceamariana (Mill.) B.S.P.). In growth room assays, seedlings were environmentally predisposed to the pathogen and coinoculated with the isolates and B. cinerea. Sporulation incidence of the pathogen was later estimated on needle segments that were cut from the seedlings and incubated on paraquat–chloramphenicol agar medium. The isolates suppressed sporulation by 0–100%. Those of Alternaria, Cladosporium, Epicoccum, yeasts, and bacteria were of low to moderate effectiveness, but those of Gliocladium, Myrothecium, Trichoderma, and Trichothecium were moderately to highly suppressive. In two greenhouse tests, spore suspensions of Gliocladium roseum Link:Bainier and of Myrotheciumverrucaria (Alb. & Schw.) Ditm. ex Stendel. (applied three times) at 2- to 3-week intervals suppressed incidence of B. cinerea by 50–69% and 42–60%, respectively, and were at least as effective as recommended fungicide treatments. Two or three additional applications of the antagonists did not further suppress the pathogen. Isolates of Fusarium sp. and Penicillium sp. were generally ineffective in the greenhouse tests. The four fungi ranked similarly in biocontrol effectiveness in the growth room and greenhouse. It is concluded that the growth room assay provided a strong indication of biocontrol performance in the greenhouse and that G. roseum and M. verrucaria have potential for managing B. cinerea in black spruce in greenhouses.

Description: Forest floors and mineral soils from early (open willow), middle (poplar–alder), and late (white spruce) floodplain primary successional stages were examined for nitrogen fixation and denitrification. The acetylene-reduction and acetylene-inhibition techniques were used separately and in combination to measure nitrogenase and denitrification activities, both in laboratory and field studies. In situ N2O production was undetectable at all sites and during all sampling periods. Denitrifying activity measured in the field with acetylene amendments was low to undetectable, except after a brief flood in the open willow stand when N2O production ranged from undetectable to 34 ng N•cm−2•h−1 within the newly deposited alluvium–old mineral soil interface. Intact core assays also had low to undetectable denitrification activities; the highest activities (259 ng N•g−1 h−1) were measured in the poplar–alder forest floor in the fall. Laboratory studies showed that potential denitrification enzyme activity (DEA) was also greatest in the poplar–alder forest floor (4332 ng N•g−1•h−1), once again occurring in the fall. In early and midsuccessional stages, the interactive effects of temperature, carbon, and NO3− limited denitrification, yet even with the addition of the limiting amendments, low to undetectable DEA was observed in mineral soils. The later white spruce successional stage also had low to undetectable DEA, increasing only with the addition of the full DEA media and independent of temperature changes. Nonsymbiotic nitrogenase activities were highly variable, ranging from undetectable to 30 ng N•cm−2•h−1. Highest activities were seen in the open willow, newly deposited alluvium–old mineral soil interface immediately after a flood and approximately 1 month after the flood on the newly deposited silt surface. Only the white spruce forest floor had measurable nonsymbiotic nitrogenase activity at all sampling times. Alder root nodule nitrogenase activity showed no significant differences between sampling periods. The estimated annual nitrogen fixation rate of 164 kg N•ha−1 for alder root nodules is a substantial N contribution to the alder stand and to the floodplain ecosystem in general.

Description: The variable-form taper function was tested on a data set of 5074 trees, consisting of jack pine (Pinusbanksiana Lamb.), lodgepole pine {Pinuscontorta Dougl.), white spruce (Piceaglauca (Moench) Voss), and trembling aspen (Populustremuloides Michx.), from Alberta. When compared with the results of tests performed on the same data using the variable-exponent function and the segmented polynomial model, the variable-form model was found to be superior (based on residual mean squares) for estimating both upper stem diameter ratios and total stem volumes. The two pine species could be combined and a single taper function used without significant loss of accuracy. Stem form in two of the volume sampling regions was significantly different from that in other regions of the province, but this may have been because the samples were not representative of the two regions. The variable-form function gave accurate and unbiased estimates of both merchantable height and merchantable volume.

Description: Eighteen natural second-growth stands of Douglas-fir (Pseudotsugamenziesii (Mirb.) Franco) from a wide range of stand densities and stocking levels were measured for stand leaf area index (LAI) from allometric relationships (direct LAI) and the LI-COR LAI-2000 plant canopy analyzer (indirect LAI). LAI was based on one-half total leaf area (m2) per square metre of ground surface area, normal to any slope. The LAI-2000 underestimated direct LAI by 62%. The underestimate of LAI was due to the confinement of needles within branches (74%) and nonrandom branch distribution (26%). In effect, the branches behaved like big leaves. The LAI-2000 can be used to estimate LAI across a variety of stand density and stocking levels by using a single stand clumping factor of 2.63.

Description: Boreal forests contain large quantities of soil carbon, prompting concern that climatic warming may stimulate decomposition and accentuate increasing atmospheric CO2 concentrations. While soil warming increases decomposition rates, the accompanying increase in nutrient mineralization may promote tree growth in these nutrient-poor soils and thereby compensate for the increased carbon loss during decomposition. We used a model of production and decomposition to test this hypothesis. In black spruce (Piceamariana (Mill.) B.S.P.), white spruce (Piceaglauca (Moench) Voss), and paper birch (Betulapapyrifera Marsh.) forests, decomposition increased with the soil warming caused by a 5 °C increase in air temperature. However, increased nitrogen mineralization promoted tree growth, offsetting the increased carbon loss during decomposition. In the black spruce forest, increased tree production was maintained for the 25 years of simulation. Whether this can be maintained indefinitely is unknown. In the birch forest, tree production decreased to prewarming levels after about 10 years. Our analyses examined only the consequences of belowground feedbacks that affect ecosystem carbon uptake with climatic warming. These analyses highlight the importance of interactions among net primary production, decomposition, and nitrogen mineralization in determining the response of forest ecosystems to climatic change.

Description: The effect of convection column air temperature and live needle moisture content on ignitability of tree branches was verified and quantified by exposing branches of three conifer species to a hot-air convection column, at temperatures between 400 and 640 °C, and measuring time to ignition. The three species were ponderosa pine (Pinusponderosa Laws.), Douglas-fir (Pseudotsugamenziesii var glauca (Beissn.) Franco), and lodgepole pine (Pinuscontorta Dougl.). The experiment was repeated monthly over the course of a year, taking advantage of the natural fluctuation of live needle moisture content. Three multiple regression equations for the prediction of time to ignition with air temperature and needle moisture as the independent variables were developed.

Description: Four species of boreal forest conifers share a similar southern limit of natural distribution in the three Prairie Provinces of western Canada. The southern boundaries of boreal forest and aspen parkland were compared with geographic patterns of several climate variables to provide a preliminary assessment of how global climate change could affect forest distribution in the future. Forest zonation corresponded most closely with climatic moisture regimes (annual precipitation minus potential evaporation). In contrast, thermal characteristics of climate (mean July and annual temperature, growing degree-days) showed an inconsistent relationship with forest zonation. It is postulated that moisture limitations prevent conifer regeneration south of the present limit of natural distribution. Alternatively, the more arid climates south of the boreal forest may have promoted higher fire frequencies historically, thus preventing conifers from achieving sufficient longevity to regenerate. The driest areas of boreal forest in the region occur at low elevations in west-central Manitoba, throughout Saskatchewan and Alberta, and the southwestern Mackenzie District, Northwest Territories; these areas may be most vulnerable to increased climatic dryness. Climatically induced losses of forest cover from these low-elevation areas could eventually lead to the fragmentation of the boreal forest in western Canada.

Description: Parent and preflight-adult Douglas-fir beetles (Dendroctonuspseudotsugae Hopk., Coleoptera: Scolytidae) were collected from felled Douglas-fir (Pseudotsugamenziesii (Mirb.) Franco) trees. Fungi isolated from the beetles included blue-stain fungi, such as Ophiostomapseudotsugae (Rumbold) von Arx, Leptographiumterebrantis Barras & Perry, and Leptographiumabietinum (Peck) Wingfield; other ascomycetes, such as Graphium spp. and Leptographium spp.; yeasts; and unidentified basidiomycetes. All fungal cultures derived from parent beetles, preflight adults, beetle eggs, larvae, frass, and bark from galleries had yeasts present. Ophiostomapseudotsugae was isolated from 67% of the parent females, 19% of the parent males, and 100% of the preflight adults of both sexes collected from their pupal cells. Rinsing the beetles with 70% ethanol prior to fungus isolations reduced the frequency of all fungi, except yeasts. Parent and preflight adult beetles (of both sexes) were examined with a scanning electron microscope, where spores of O. pseudotsugae were visible in shallow pits on the elytra of both male and female beetles. Spores of other fungi were observed in shallow pits on elytra and in much smaller but deeper pits on the scutellum of both sexes. The results indicate a close association between Douglas-fir beetles and fungi, accompanied by anatomical differentiation on the beetles that allows the dissemination of blue-stain fungi to new Douglas-fir hosts and substrates.

Description: Differences in N, P, and K status and N efficiency of some North American willow species and their clones were observed and evaluated. Salixeriocephala Michx., S. lucida Mühl., and S. exigua Nutt., represented by five clones each, were raised from cuttings in potted loamy sand for 113 days. Plants were subjected to three levels of fertilization (equivalent to 125, 250, or 500 kg N/ha) applied at exponential rates of addition (0.06, 0.071, and 0.082, respectively), using complete nutrient solutions and were compared with control plants (no fertilizer added). The experiment showed that nutrient concentration and nutrient uptake in the plants depended not only on treatments but also on species and clones. Thus, species and clones that removed smaller amounts of nutrients per unit of biomass could be selected. Species had a larger effect than clones on nutrient concentration and content. Total nutrient content, at the same treatment level, depended primarily on biomass production. Significant variation existed among species and clones in the ratio of biomass/total N. Salixeriocephala (the best species) produced 35.1% more than the other species, and the best experimental clones within species produced 27.8–41.6% more stem per unit of N than the poorest clones.

Description: Concepts and procedures are presented for the analysis of progeny trials that incorporate clonal replication as a means to resolve variance arising from nonadditive gene effects. Components of variance from the linear model may be expressed in terms of expected covariances among relatives, and these, in turn, may be used to derive approximations of additive, dominance, and epistatic components of genetic variance. In addition to the usual assumptions applied to conventional progeny trials, the use of this expanded genetic model in the analysis of tests with clonal replicates assumes that the greatest portion of the total epistasis is due to interactions involving groups of more than two or three loci. If this assumption is not satisfied, estimates of additive and dominance variance, including those from trials without clonal replicates, will be contaminated by a large fraction of epistasis, and total epistasis will be underestimated by a corresponding amount. Heritability and gain formulae for alternative selection and deployment schemes are developed and illustrate the use of genetic parameters in the comparison of seedling and clonal reforestation strategies.

Description: Identical pair crosses, including reciprocals, in Pinussylvestris L. (Scots pine) were made on ramets of the same clones in three clonal archives (seed orchards) in Sweden: Sävar (64°N), Röskär (59.5°N), and Degeberga (56°N). The offspring were used to test the hypothesis that the parental environment could affect the performance of the progeny (aftereffects). Growth and freezing tests were performed in the controlled conditions of the Stockholm Phytotron. Parental environment affected seed weight: the heaviest seeds came from Röskär and the lightest seeds, from Degeberga. Height development was affected in the two growth periods tested: seeds from Sävar produced the shortest plants and seeds from Röskär, the tallest plants. There was an effect on the autumn frost hardiness in the first growth period that disappeared after the second growth period. The most hardy progenies came from Sävar. The aftereffects of the parental environment were less than the maternal effects on seed weight and also less than the effects of full-sib families on growth and autumn frost hardiness. Small but mostly significant reciprocal effects were found for height and height increment during the second growth period. There was a significant reciprocal effect for seed weight. Seed weight differences could explain only a small part of the effects on growth and none of the effects on hardiness.

Description: The frost hardiness of 15- to 25-year-old Scots pine (Pinussylvestris L.) and Norway spruce (Piceaabies (L.) Karst.) growing under central Finnish conditions was followed during 1985–1987. Shoots were subjected to artificial frost in the laboratory. Frost hardiness was assessed by the impedance method and by visual scoring. Frost hardiness varied during the years from −3.5 °C to lower than −40 °C. The rate of dehardening increased after about mid-April in both tree species when the daily mean temperature increased by several degrees above 0 °C. The maximum rate of dehardening varied slightly from year to year. In both species the frost hardiness of the previous year's shoot decreased during shoot elongation. This phenomenon was more prominent in pine than in spruce. Shoots were most susceptible to frost damage at the time when shoot elongation was ceasing. The onset and development of hardiness in autumn varied from year to year, especially in spruce. Some difference in hardening was found between the current and the previous year's shoots. The rate of hardening increased typically around mid-September in both species, when the mean daily temperatures decreased to within the range of 5–10 °C.

Description: The biochemical bases of water-stress tolerance in a pedigree consisting of black cottonwood (Populustrichocarpa Torr. & Gray female) and eastern cottonwood (Populusdeltoides Bartr. male) parental clones and four hybrid progeny were investigated. Trees were grown outdoors in pots; well-watered trees (soil water potential greater than −0.03 MPa) were kept moist in trays, and stressed trees (soil water potential less than −2.0 MPa) were subjected to repeated cyclical stress of 1 or 2 days duration over the 14-week study. Analysis of the major metabolites and ions in fully expanded leaves demonstrated that the greatest degree of osmotic adjustment was displayed by male hybrid 242, the P. deltoides male parent, and male hybrid 239 to a lesser extent. Osmotic adjustment in leaves of both hybrid 242 and the P. deltoides male parent was primarily constituted by malic acid, K, sucrose, and glucose, with the same metabolites also increasing in fine roots of hybrid 242, the only clone to display osmotic adjustment in roots. Female clone 240 and P. deltoides displayed organic solute-based adjustments to water stress that were offset by declines in inorganic ions, particularly Na and Mg. Given that the P. trichocarpa female parent did not display osmotic adjustment in either tissue, the hybrids' capacity for adjustment was likely conferred by the P. deltoides male parent.

Description: Sampling of 24-, 34-, and 44-year-old patch cuts (324–2400 m2) in the Bartlett Experimental Forest, New Hampshire, was undertaken to assess the effect of gap size and location within a gap on tree species abundance (relative basal area and relative density). Shade-tolerant species, especially eastern hemlock (Tsugacanadensis (L.) Carr.) and American beech (Fagusgrandifolia Ehrh.), were relatively more abundant in small gaps and gap edges and generally decreased with increasing gap size. Shade-intolerant species, including paper birch (Betulapapyrifera Marsh.) and pin cherry (Prunuspensylvanica L.f.), were relatively more abundant in large gaps and gap centers and increased with increasing gap size. Intermediately shade-tolerant species, especially yellow birch (Betulaalleghaniensis Britt.) and red maple (Acerrubrum L.), were relatively more abundant in gap centers. Striped maple (Acerpensylvanicum L.) was relatively more abundant in gap edges. Many of these relationships were complex due to interactions with gap age and slope. Sugar maple (Acersaccharum Marsh.) relative abundance was not associated with gap size or location within a gap. Analyses isolating irradiance as a factor influencing species composition were inconclusive. Instead, other effects of gap disturbance and characteristics associated with different locations in the gap, such as soil conditions and root competition, may play an important role in the gap dynamics of this northern hardwoods forest. Gap age had a strong effect on species relative abundances and these patterns reflected typical successional sequences in northern hardwood forests. The gap disturbances increased species richness and diversity in this forest. Gaps contained species not present in the old-growth forest, and the species compositional variations among different gap sizes suggest that a forest with a range of gap sizes will have high diversity. Competitive exclusion appeared to be prevented by the gap disturbances, a likely consequence of the release of previously unavailable resources.

Description: The temporarily increased fire hazard that is believed to result from the process of thinning is included in a single-stand model for assessing the economic benefits of juvenile spacing. Formulas for the expected net present value and the land expectation value are given along with methods for determining the age of financial maturity and the optimal rotation age. A numerical example is given to illustrate the degree of loss due to the increased fire risk. The problem of commercial thinning when the risk of fire is present is addressed using continuous-time models. It is shown how, when the fire hazard is exogenous to the thinning activity, the problem reduces to one of deterministic optimal control with the discount rate adjusted upward by an amount equal to the fire hazard rate. In the case when the fire hazard increases whenever thinning is taking place, it is shown that in general the optimal thinning policy is qualitatively different from that which is optimal in the no-risk case and involves periods of thinning at the maximum rate interspersed with periods of no thinning activity.

Description: Some physical, thermal, and chemical properties of bark of 16 tree species native to the central hardwood region were measured to determine their potential to protect the vascular cambium from damage by fire. The relationship between DBH and bark thickness for each of 16 species was determined. For purposes of monitoring seasonal trends, two species (Quercusmacrocarpa Michx. and Acersaccharinum L.) were sampled periodically during one growing season. Temperature response to bark surface heating of 11 species was monitored at the cambial layer during simulated fires conducted in the field. Bark samples were analyzed for moisture content, specific gravity, dry weight, volatile matter content, and time until ignition. Overall, during simulated fires, temperature gradients were decreased and maximum cambial temperatures were reduced as bark thickness increased. Thick-barked species had lower maximum cambial temperatures, longer times to reach peak temperatures, slower rates of heat loss, and shorter time until surface ignition. Populusdeltoides Marsh, was the most heat resistant among species tested, while Acersaccharinum was the least. Higher specific gravities were associated with higher rates at which cambial temperatures rose as well as with increased time required for surface ignition.

Description: Biomass and nutrient dynamics were examined in a subalpine Piceaengelmannii Parry–Abieslasiocarpa (Hook.) Nutt. forest within Loch Vale watershed in north central Colorado by quantifying annual above- and below-ground production, biomass, nutrient pools, and internal nutrient transfers. Subalpine forest covers only 6% of the watershed, and the values reported here refer only to the forested area. Total ecosystem biomass was 42 kg•m−2, of which 30% was soil organic matter, 33% was detrital biomass (including deadwood and forest floor), and 36% was living biomass. Total forest biomass (not including soil organic matter) was 28.9 kg•m−2, of which root biomass was 11%. Net primary production was 520 g•m−2•year−1, of which fine root production was about 27% and foliar production was 30%. Much more N was recycled via fine root turnover than via aboveground litter fall (1.6 versus 0.9 g•m−2•year−1), whereas four times more Ca was returned via litter fall than via fine roots. Compared with other temperate coniferous forests, this subalpine forest had low production. Nutrient resorption contributed between 35 and 38% of the annual requirements of N, P, and K, but only 9% of Ca and 12% of Mg. Although a higher percentage of annual N requirement was met by resorption, this forest used N less efficiently than a similar forest in southwestern Alberta.

Description: We tested the hypothesis that naturally occurring nitrogen (N) isotope ratios in foliage (from plants that do not symbiotically fix atmospheric N2) are an indicator of soil N dynamics in forests. Replicate plots were established at eight locations ranging in elevation from 615 to 1670 m in Great Smoky Mountains National Park in eastern Tennessee, U.S.A. The locations selected ranged from N-poor (low-elevation) to N-rich (high-elevation) forest stands. Soils were sampled in June 1992; plants, forest floors, and upper mineral soils were sampled in August 1992. Net N mineralization and net nitrification potentials for surface mineral soils and organic matter layers at each site were determined by aerobic laboratory incubations. Soils and organic layers from high-elevation sites had greater net N mineralization and nitrification potentials than soils from low-elevation sites. There were significant (P ≤ 0.05) differences between study sites in soil 15N abundance. Therefore, we examined correlations between measures of soil N availability and both mean foliar δ15N values and mean enrichment factors (εp−s = δ15Nleaf − δ15Nsoil). In evergreens, maples, and ferns, mean foliar δ15N values and mean enrichment factors were positively correlated with net N mineralization and net nitrification potentials in soil. The observed relationships between natural 15N abundance in plant leaves and soil N availability were explained by a simple model of soil N dynamics. The model predicts how the isotopic composition of plant N is affected by the following factors: (i) varying uptake of soil NH4-N and NO3-N, (ii) the isotopic composition of different soil N pools, and (iii) relative rates of soil N transformations.

Description: A 1992 study of serotiny in lodgepole pine (Pinuscontorta Dougl. ex Loud. var. latifolia Engelm.) in Yellowstone National Park asked four questions: (i) are there morphological characteristics that can be used to estimate pre-fire proportion of serotinous trees in forests that burned in 1988?; (ii) at what spatial scale does percent serotinous trees vary across the landscape?; (iii) which environmental factors are correlated with serotiny?; and (iv) what is the relationship between prefire serotiny and postfire lodgepole pine seedling density? We first sampled cone characteristics in serotinous and nonserotinous trees along four 2950-m transects in unburned forests, and examined burned trees nearby. Results indicated that asymmetrical cones and an acute angle of cone attachment to the branch were reliable indicators of serotiny even in burned trees. We then sampled nine patches of lodgepole pine forest that had burned in 1988, and varied in size from 1–3600 ha. We sampled serotiny at varying intervals along two perpendicular transects that crossed in the center of each patch. At each sample point, the 12 nearest canopy lodgepole pines were classified as serotinous or nonserotinous. We concluded that the percentage of serotinous trees is most variable at intermediate scales of 1–10 km, and is relatively homogeneous at both fine scales (