Growth–climate responses of high-elevation conifers in the central Hengduan Mountains,southwestern China

Improved understanding of tree-growth responses to climate is needed to model and predict forest ecosystem responses to current and future climatic variability. We applied dendroclimatological techniques to assess the effects of inter-annual climate variations on radial growth of high-elevation conifers in the central Hengduan Mountains, southwestern China. Eight tree-ring width chronologies of the major tree genera Abies and Picea that are aligned along an elevation gradient from 3200 to 4200 m a.s.l. were developed. Correlation and principal component analyses for the eight chronologies identified three groups of sites, representing different patterns of growth–climate relationships. Correlation and redundancy analyses with regional climate data revealed that radial growth of fir growing at high-elevation sites is enhanced by normal or warm summer temperatures (June and July) during the current growing season. In addition, radial growth of trees growing from high to middle elevations is sensitive to low temperatures during winter season. At low-elevation sites, trees display low sensitivity to temperature variation. However spring moisture availability becomes crucial for radial growth regardless of tree species. High- to middle-elevation conifers in the central Hengduan Mountains may benefit from the current climate warming, especially from rising winter temperatures.     

Chronology development of Hiba arbor-vitae (<Emphasis Type="Italic">Thujopsis dolabrata</Emphasis> var. <Emphasis Type="Italic">hondae</Emphasis>) and dating of timbers from an old building

Two site chronologies of the ring widths of Hiba arbor-vitae (Thujopsis dolabrata var. hondae) were developed for the northern-most part of Honshu Island, Japan (AOTK: 1715-2001, AOSO: 1753-2000). These are highly coherent, satisfactorily indicating high values of expressed population signal and mean correlation between trees. The two series were merged into a single composite chronology, spanning a period of 287 years from 1715 to 2001, because they correlated significantly with each other. The developed chronology is the first quality-controlled tree-ring chronology for this region extending back to the eighteenth century. Using the master chronologies, we successfully crossdated three historical timbers from an old building near the sampling sites. The tree-ring dates strongly supported the documented construction date of the building. It is revealed that Hiba arbor-vitae has sufficient potential to contribute to future dendrochronological study in this region by extending the chronologies back into the past.     

Tree-ring response of Larix chinensis on regional climate and sea-surface temperature variations in alpine timberline in the Qinling Mountains

Tree-ring width chronologies of Larix chinensis were developed from the northern and southern slopes of the Qinling Mountains in Shaanxi Province,and climatic factors affecting the tree-ring widths of L.chinensis were examined.Correlation analysis showed that similar correlations between tree-ring width chronologies and climatic factors demonstrated that radial growth responded to climate change on both slopes.The radial growth of L.chinensis was mainly limited by temperature,especially the growing season.In contrast,both chronologies were negatively correlated with precipitation in May of the previous year and April of the current year.Spatial climate-correlation analyses with gridded land-surface climate data revealed that our tree-ring width chronologies contained a strong regional temperature signal over much of northcentral and eastern China.Spatial correlation with seasurface temperature fields highlights the influence of the Pacific Ocean,Indian Ocean,and North Atlantic Ocean.Wavelet coherence analysis indicated the existence of some decadal and interannual cycles in the two tree-ring width chronologies.This may suggest the influences of El Nin˜o-Southern Oscillation and solar activity on tree growth in the Qinling Mountains.These findings will help us understand the growth response of L.chinensis to climate change in the Qinling region,and they provide critical information for future climate reconstructions based on this species in semi-humid regions.     

Larix olgensis growth-climate response between lower and upper elevation limits:an intensive study along the eastern slope of the Changbai Mountains,northeastern China

Larix olgensis is a dominant tree species in the forest ecosystems of the Changbai Mountains of northeast China.To assess the growth response of this species to global climate change,we developed three tree-ring width and biomass chronologies across a range of elevations in the subalpine forests on the eastern slope of the Changbai Mountains.We used dendroclimatic analyses to study key factors limiting radial growth in L.olgensis and its variation with elevation.The statistical characteristics of chronologies suggested that elevation is a determinant of tree growth patterns in the study area.Response function analysis of chronologies with climate factors indicated that climate–growth relationships changed with increasing elevation:tree growth at high elevation was strongly limited by June temperatures of the previous year,and as elevation decreases,the importance of temperature decreased;tree radial growth at mid-elevation was mainly controlled by precipitation towards the end of the growing season of the current year.Biomass chronologies reflected a stronger climatic signal than tree-ring width chronologies.Spatial correlation with gridded climate data revealed that our chronologies contained a strong regional temperature signal for northeast China.Trees growing below timberline appeared to be more sensitive to climate,thus optimal sites for examining growth trends as a function of climate variation are considered to be just below timberline.Our study objective was to provide information for more accurate prediction of the growth response of L.olgensis to future climate change on the eastern slope of the Changbai Mountains,and to provide information for future climate reconstructions using this tree species in humid and semi humid regions.     

Canopy openings may prevent fir broom rust (<Emphasis Type="Italic">Melampsorella caryophyllacearum</Emphasis>) infections

Melampsorella caryophyllacearum is a heteroecious fungus that produces disease on silver fir resulting in galls that depreciate the value of trees. Currently, little is known about its epidemiology. The purpose of this study was to identify those site or management factors associated with fir broom rust infection. A 29-plot systematic sampling was carried out throughout all silver fir forests in the Spanish Pyrenees. In every sampling plot, we examined each tree for the presence of M. caryophyllacearum galls or witch’s brooms. We differentiated their position as either on the main stem or on the branches of trees. We identified the most susceptible stands by relating a compendium of tree and stand variables with the presence and severity of symptoms. Disease symptoms were observed in 11.8% (95% CI: 5.8–17.9%) of trees. Fir broom rust-infected trees tended to be those of larger diameters. The rust infection was observed in 55.2% (95% CI: 35.7–73.6%) of the stands. We identified a wide range of severities with 5 of 29 stands having more than 30% of trees infected. Fir broom rust appeared to be a widespread disease, although we observed important differences in terms of severity. The presence and severity of the rust in stands was associated with understories composed of more shade tolerant plant species and with longer periods without thinnings. Canopy opening should be experimentally tested as a possible control method.     

Fungal colonization of coastal Douglas‐fir following mechanical commercial thinning damage

Commercial‐aged Douglas‐fir trees mechanically damaged during commercial thinning activities were investigated to quantify fungal taxa present 14 years after thinning. A total of 235 wood cores from 18 damaged and 18 non‐damaged control trees were obtained. Fungi were isolated on malt extract agar and molecular methods were used to identify 48 taxa. Damaged Douglas‐fir trees were more than twice as likely to contain wood‐inhabiting fungi as undamaged trees (p = 0.07) and contained more than 2.5 times more fungi (p = 0.006). However, typical Douglas‐fir stem‐decay fungi were not isolated. Fungi present were mostly endophytes and no significant differences were found in frequencies of decay/non‐decay soft‐rot fungi between damaged and control trees (p = 0.67). Likewise, no significant differences were found for the trend in number of fungi isolates over radial depth into the stem between damaged and control trees (p = 0.29). Frequency of fungal isolates was non‐linearly related to depth in the tree bole. Significant differences between damaged and control trees (p = 0.03) was influenced by the data in the outermost 3 mm. With the exception of these data, there were no significant differences between damaged and control trees in the radial trend in fungal frequency (p = 0.37). Results of this study suggest that wounding in commercial‐aged Douglas‐fir may not contribute significantly to occurrence and species composition of wood inhabiting fungi over the period of a short‐rotation final harvest. However, damage mitigation is recommended supported by a related study showing vertical discolouration affecting all wounded stems.     

The pine pathogen Diplodia sapinea is associated with the death of large Douglas fir trees

Climate change poses severe pressures to European conifer forests. Using non-native tree species, such as Douglas fir (Pseudotsuga menziesii), is one proposed strategy to circumvent adverse effects for forest management. However, novel forest health risks can impair the cultivation of non-native trees. In 2022, we observed large Douglas fir trees (approximately 40–50 years old, diameter at breast height (dbh) 21–41 cm) that had recently died in spring or summer 2022 in three forest stands in Eastern Austria. Intensive resin flow, blue-staining of the sapwood and the absence of bark- and wood-boring insects indicated a fungal infection. Isolations from blue-stained sapwood of the dead trees consistently yielded cultures of the opportunistic pathogen Diplodia sapinea. In a greenhouse wound inoculation experiment, seven D. sapinea isolates obtained from Douglas fir caused phloem necrosis, blue-staining of sapwood and mortality and thus displayed pathogenicity towards seedlings of both Ps. menziesii and its common host, Scots pine (Pinus sylvestris). Diplodia sapinea produced significantly longer areas of blue-stain as well as higher and faster mortality in Ps. menziesii compared to P. sylvestris. We conclude that D. sapinea substantially contributed to the death of seven of the 13 examined large Douglas fir trees. While this fungus has been described as a pathogen of young Douglas fir trees before, this is the first report that it can potentially kill large individuals of this conifer species under drought conditions. Thus, our results indicate that D. sapinea could represent a severe threat to the cultivation of Ps. menziesii in European forestry.     

Stamm- und Wurzelfäulen in Douglasien,Pseudotsuga menziesii (Mirb.) Franco1

Stem and root rot of Douglas fir, Pseudotsuga menziesii (Mirb.) Franco. In 20 year old Douglas fir most of the butt rot was caused by Fomes annosus. Stem decay was central as well as eccentric reaching the sap wood in either case. The side roots of 20 year old Douglas fir were compared with those of a 40 year old stand with butt rot. In individual trees with decay there were less roots with Fomes annosus decay in the younger stand. Calocera viscosa was more abundant in the older stand. Soil conditions which might have favoured root rot in the young stand are discussed.     

Size-mediated climate-growth relationships in temperate forests: A multi-species analysis

In most dendrochronological studies, climate-growth relationships are established on dominant trees to minimize non-climatic signals. However, response to environmental factors may be affected by tree-size, which begs the question of the representativeness of dominant trees on the stand level. To highlight the variations in climate-growth relationships among sizes and species, under a wide range of ecological conditions (climate and soil properties), 61 pure even-aged stands were sampled across France. At each stand, two tree-ring chronologies were established from 10 big- to 10 small-diameter trees. Our objectives were, (1) to assess variations in climate sensitivity between the two size-diameter classes, and (2) to investigate the role of species and ecological conditions on these variations. The climate-growth relationships were evaluated from 122 tree-ring chronologies (1 220 trees) through extreme growth years and correlation function analyses. Sensitivity to climate of shade-intolerant and moderately shade-tolerant species (Picea abies (L.) Karst., Pinus sylvestris L. and Quercus petraea (Matt.) Liebl.) remained constant between the size-diameter classes for both temperature and hydric balance, while the shade-tolerant species Abies alba Mill. and Fagus sylvatica L. displayed significant differences, with larger trees being more sensitive to summer drought than smaller trees. This difference increased with increasing climatic xericity. Our results suggest that, for shade-tolerant species, (1) big trees could be more sensitive to climatic change especially under xeric climate, and (2) future tree ring studies should include trees stratified by size to produce unbiased estimation of sensitivity to climate.     

Relationships between tree-ring cell features of <Emphasis Type="Italic">Pinus koraiensis</Emphasis> and climate factors in the Changbai Mountains,Northeastern China

Anatomical characteristics have been proven useful for extracting climatic signals. To examine the climatic signals recorded by tree-ring cell features in the Changbai Mountains, we measured cell number and cell lumen diameter, in addition to ring widths, of Korean pine (Pinus koraiensis) tree rings at sites of varied elevation, and we developed chronologies of cell number (CN), mean lumen diameter (MLD), maximum lumen diameter (MAXLD) and tree-ring width (TRW). The chronologies were correlated with climatic factors monthly mean temperature and the sum of precipitation. As shown by our analysis, the cell parameter chronologies were suitable for dendroclimatology studies. CN and TRW shared relatively similar climatic signals which differed from MLD and MAXLD, and growth-climate relationships were elevation-dependent, as shown by the following findings: (1) at each elevation, MLD and MAXLD recorded different monthly climatic signals from those recorded by TRW for the same climatic factors; and (2) MLD and MAXLD recorded climatic factors that were absent from TRW at lower and middle elevations. Cell lumen diameter proved to be an effective archive for improving the climate reconstruction for this study area.     

Climatic responses of tree-ring widths of <Emphasis Type="Italic">Larix gmelinii</Emphasis> on contrasting north-facing and south-facing slopes in central Siberia

An analysis was performed on the climatic responses of the radial growth of Larix gmelinii (Rupr.) Rupr. on contrasting north-facing and south-facing slopes in Tura, central Siberia. We developed chronologies of tree-ring width for four plots, designated as north-upper, northlower, south-upper, and south-lower. Both residual and standard chronologies of tree-ring widths exhibited a significant positive correlation with temperature from the end of May until early June in all four plots. The chronologies of ring width did not reveal any major difference in the response to temperature among the four plots. The standard chronologies of ring widths on the north-facing slope were negatively correlated with precipitation during the winter (October–April) and in early and mid-May, whereas the residual chronologies did not reveal clear relationships with precipitation during the winter and May. The significant correlation between ring width and temperature from the end of May until early June indicates that temperatures in springtime play a significant role in the radial growth of L. gmelinii. The negative correlations between standard chronologies of tree-ring width and precipitation in the winter and in May on the north-facing slope indicate that lowfrequency fluctuations in snowfall have negative effects on the radial growth. However, these effects vary and depend on the microscale topography. Part of this report was presented at the 55th Annual Meeting of the Japan Wood Society, March 2005, Kyoto     

The relationship between Swiss needle cast symptom severity and level of Phaeocryptopus gaeumannii colonization in coastal Douglas‐fir (Pseudotsuga menziesii var. menziesii)*

This study examined 108 15‐year‐old Douglas‐fir (Pseudotsuga menziesii var. menziesii) trees to investigate whether trees exhibiting less severe Swiss needle cast (SNC) symptoms were more resistant (had less fungal colonization) or more tolerant (maintained healthy foliage under similar infection levels). Trees were sampled from six open pollinated families that were categorized into three disease severity groups (two families for each group; mild, moderate and severe disease symptoms). The amount of retained foliage and level of discoloration were visually assessed on trees in the field. Fungal colonization (as determined by proportion of stomata occluded with pseudothecia and by amount of Phaeocryptopus gaeumannii DNA in sampled needles) was measured on 1‐ and 2‐year‐old needles in the laboratory. Trees in the different disease severity groups were similar with respect to amount of fungus in their needles, yet the trees in the mild symptom group retained higher proportions of needles and maintained greener foliage. The relationship between amount of P. gaeumannii in needles and SNC symptom severity was statistically significant (p < 0.05) for amount of fungal DNA in 1‐year‐old needles and average needle retention (NR) over the last four growing seasons. Average NR decreased with increased amount of pathogen DNA in the mild disease symptom families. This relationship was reversed in the severe disease symptom group and there was no relationship in the moderate disease symptom group. Because the amount of P. gaeumannii DNA in foliage did not differ significantly among the groups, differences in symptom severity were attributed to tolerance, not resistance. Visual scoring of individual trees for average NR over the past four growing seasons could be used to effectively assess for SNC tolerance in Douglas‐fir.     

THE PLACE OF A SPECIES IN THE FOREST, WITH SPECIAL REFERENCE TO WESTERN NORTH AMERICAN SPECIES OF CONIFER USED IN BRITAIN

The nature of the differences between the climates of the QueenCharlotte Islands and of England and Wales are discussed briefly.The author then examines the site tolerance of the principalspecies within their own natural range, comparing particularlythe place of Sitka spruce (Picea sitcbensis Carr.) in the foreston Graham Island with its place around Terrace and describingthe Douglas fir forests of the drier eastern side of VancouverIsland and the place that Sitka spruce has in them. Factorsaffecting the growth of Douglas fir (Pseudotsuga taxifolia Brit.)are discussed. It is suggested that care is needed before itis assumed that a markedly dry summer period such as occursin much of the Douglas fir region entails a ‘dry’period as regards supply to the tree. Illustrations are givenfrom eastern North America of changes in type of site occupiedby a species with change in climatic or edaphic environment.Observation of the relative position of a species in the forestas climate changes makes it possible to suggest the place ofSitka spruce and Douglas fir in Great Britain.     

暖温带幼林树轮宽度气候变化响应分析--以山东蒙山黑松树轮气候研究为例

根据采自山东蒙山地区两组幼龄黑松树木年轮样本,研制出单点和区域树轮宽度年表,分析了蒙山黑松树轮宽度年表的统计参数和径向生长的气候限制因子,并尝试重建相关密切的气候因子,探讨蒙山黑松径向生长与大范围气候变化的关联。研究结果表明,（1）蒙山黑松树轮宽度年表的统计参数都比较高,说明蒙山黑松树轮宽窄变化包含有较多的气候环境变化信息。（2）蒙山黑松径向生长变化与当年4-9月降水量呈显著正相关。同时,当年5-7月异常高温也对蒙山黑松径向生长变化产生显著抑制。通过黑松树轮宽度年表对4-9月降水量的模拟重建结果表明蒙山黑松具有较大树轮气候研究潜力。（3）蒙山黑松树轮宽度生长与太平洋大范围气候波动存在显著关联。     

Polyporus schweinitzii Fr. und Sparassis crispa (Wulf. in Jacq.) ex Fr. als Fäuleerreger in einem Douglasienbestand (Pseudotsuga menziesii (Mirb.) Franco) mit hohem Stammfäuleanteil1

Polyporus (Phaeolus) schweinitzii and Sparassis crispa as frequent decay fungi in Douglas fir. Of fifty, 55 ears old trees in a Pseudotsuga menziesii stand. 90% showed decay of the heart- wood of the butt. The possibility of transmission of the most frequent decay fungi, P. schwei- nitzii and S. crispa, which also are frequent parasites in the roots and butts of pine, from the roots of the Pinus sylvestris trees of the previous crop into the roots of the Douglas fir is discussed. A first identification of the fungi was made possible by storing the stem discs under moist conditions. The mycelia of P. schweinitzii and S. crispa grew out of the decayed wood and possessed a characteristic form and colour.     

Necrophylactic periderm formation in the roots of western larch and Douglas‐fir trees infected with Armillaria ostoyae. I. The response to abiotic wounding in non‐infected roots

The sequence of events leading to necrophylactic periderm formation was studied throughout the year following the abiotic wounding of the non‐infected roots of 10‐ and 27‐year‐old western larch (Larix occidentalis) and 11‐ and 25‐year‐old Douglas‐fir (Pseudotsuga menziesii) trees that were infected with Armillaria ostoyae. The sequence was the same for both ages and species of trees. Wound repair was more rapid in the summer compared with the spring and autumn. Following cell hypertrophy, a zone of lignified impervious tissue was in the initial stages of formation within 10 days of wounding in the summer and 14 days in the spring or autumn. The new phellogen produced a layer of phellem three to four rows of cells thick after 20 days in the summer or 40 days in the spring. Modified cells abutting the inner boundary of the impervious zone frequently developed thick lignified abaxial walls and thin suberized adaxial walls. A typical exophylactic periderm in healthy root bark tissue of both western larch and Douglas‐fir consisted of stone phellem one to four rows of cells thick and a layer of thin‐walled phellem three to six rows of cells thick in western larch and two to three rows thick in Douglas‐fir, a single row of phellogen cells and one to three rows of phelloderm cells. Mature thin‐walled phellem cells had pigmented contents, red in western larch and light brown in Douglas‐fir. In response to wounding, 27‐year‐old western larch and 25‐year‐old Douglas‐fir developed necrophylactic periderms with annual bands of phellem. The bands included a layer of phellem that was six to 12 and nine to 15 rows of cells thicker than the layer of phellem observed in the respective naturally developed exophylactic periderms. Fifty days following wounding in the summer, stone phellem, one to three rows of cells thick, was observed in the necrophylactic periderm of 10‐year‐old trees. When fully developed, the necrophylactic periderm in 27‐year‐old western larch also had a layer of stone phellem three to five rows of cells thick in each band. Stone phellem development was only sporadic in 25‐year‐old Douglas‐fir. Wounds in the winter showed no signs of activity associated with repair until dormancy broke in the spring.     

The effect of climate on radial growth of horse chestnut (<Emphasis Type="Italic">Aesculus hippocastanum</Emphasis> L.) in the Świętokrzyski National Park in central Poland

The objectives of this study were to construct a site tree-ring width chronology of horse chestnut (Aesculus hippocastanum L.) growing in a single locality situated in the Świȩtokrzyski National Park, and to determine its sensitivity to the two main climatic elements of air temperature and precipitation. Fifteen horse chestnut trees, growing on a moist site and not attacked by the leaf miner (Cameraria ohridella Deschka et Dimić), were selected for the study. Relationships between monthly values of air temperature, precipitation, and width of tree rings during 1932–2003 were analyzed, in addition to using bootstrapped response and correlation functions for single and multiple time intervals (the computer program DENDROCLIM2002 was used). High air temperature of the previous winter, as well as the air temperature of August of a given growing season positively affected the radial growth of horse chestnut. The cambium activity was also favored by ample precipitation in December preceding a given growing season. Excessive precipitation in August, which raised the existing high water table, had a negative effect on tree-ring width. The tree-ring chronology, elaborated for horse chestnut during this study, may be a local growth standard of this species. Thus, the model of climate–tree radial increment relationships may depend on characteristics of the locality where the trees are growing. Therefore, it would be advisable to undertake comparative studies.     

Age structure and regeneration of subalpine fir (Abies fargesii) forests across an altitudinal range in the Qinling Mountains,China

Age structure and regeneration dynamics of subalpine fir (Abies fargesii) forest were studied across the altitudinal range in both the north and south aspects of the Qinling Mountains, China. Ages of individual fir trees were determined based on the number of rings counted from cores and the number of years to reach coring height estimated using age–height regression. Fir age structure and regeneration dynamics were similar in both the north and south aspects. A unimodal population age structure was found at the low- and mid-elevations in both aspects, indicating that environmental factors might play an important role in shaping A. fargesii age structure and regeneration at those sites. There was a recruitment pulse during the time period 1830–1890 at each altitudinal site, but no stem recruitment occurred at the low- and mid-elevations in the last century, which might be attributed to the intensive cover of understory bamboo. Fir trees were, however, persistently recruited at the upper limits during the last 150 years, and the fir tree density at the upper limits was significantly higher than that at the lower limits in both aspects. The fir population at the upper limits showed a significant increase in recruitment and stem density relative to the fir population at the low- and mid-elevations in the last century. We propose that the differences in recruitment might promote variations in stand structure and regeneration dynamics of the subalpine fir forests along the altitudinal gradient in the Qinling Mountains, China.     

Lesion formation and host response to infection by Armillaria ostoyae in the roots of western larch and Douglas‐fir

The process of lesion formation and host response to natural infection by Armillaria ostoyae were studied in the roots of western larch (Larix occidentalis) and Douglas‐fir (Pseudotsuga menziesii ssp. glauca) trees in the three age classes, 6–8, 18–19 and 85–95 years. The characteristics of lesions on infected roots were recorded and bark samples were dissected from infection points and lesion margins in the field and stored in liquid nitrogen for macroscopic study in the laboratory. Infection in the roots of 6‐ to 8‐year‐old trees advanced freely, overcoming any host resistance, quickly girdling the root collar and killing the trees. In 18‐ and 19‐year‐old trees, however, 43% of infections on western larch and 27% of the infections on Douglas‐fir roots were confined to lesions bounded by necrophylactic periderms with multiple bands of phellem. Host response was similar in 85‐ to 95‐year‐old trees, but the percentage of confined lesions was higher than in younger trees. The results suggest that larch shows resistance to A. ostoyae at a younger age and with greater frequency than Douglas‐fir.     

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

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