Influence of local stand basal area on density and growth of regeneration in uneven‐aged Picea abies stands

In single‐tree selection, trees removed by harvest or lost through mortality are replaced by ingrowth from the seedling/sapling bank. Because the level of ingrowth is governed not only by the recruitment rate of new seedlings, but also by mortality and growth rates within the seedling/sapling stratum, knowledge of how these processes are related to the tree stratum is important for successful application of single‐tree selection. Therefore, Norway spruce (Picea abies (L.) Karst.) regeneration (0.1 m ≤ height ≤ 2.0 m) was measured on seventy 100 m² circular subplots at each of two sites in central and northern Sweden. Both sites had previously been selectively logged, but the time elapsed since the last harvest was at least 30 yrs. Basal area of trees of at least 2 m height within three different radii from the subplot centres was measured. Measurements on regeneration included height and leader length. Influence of local stand basal area on density and growth of regeneration was analysed by ordinary least‐squares linear regression for each of four height intervals. Regeneration was significantly aggregated at both sites, but number of Norway spruce seedlings and saplings per circular subplot was not correlated to local stand basal area. Out of 36 regressions, only eight showed a significantly (p < 0.05) negative correlation between seedling and sapling height growth and local stand basal area. In five of these cases, removal of one point (subplot) made p > 0.05. The results suggest that density and growth of Norway spruce regeneration in selectively logged uneven‐aged Norway spruce stands is affected more by ground conditions than by local stand basal area.     

Relationship between seedbed properties and the emergence of spruce germinants in recently cut Norway spruce selection stands in Southern Finland

The purpose of this study was to examine the effects of type and amount of ground vegetation and substrate on the germination and early survival of Norway spruce (Picea abies (L.) Karst) seedlings in uneven-aged stands on mineral soils in Southern Finland. The study was conducted on two permanent plots representing average site quality for spruce stands in the area. A gradient in selection cut intensity and residual stand density was imposed across each of the two permanent plots. Regeneration and vegetation surveys were repeated annually on a grid of unmanipulated microplots and on a set of soil treatment microplots. The results indicated that the number of spruce germinants per unit area was positively correlated with herb cover but negatively correlated with total plant cover. Germinant density was also related to vegetation-free seedbed conditions, high overstory competition index and presence of larger spruce seedlings. Regeneration microsites dominated by moss or with plants with a relatively high light demand were associated with low germinant numbers. Bare humus layer and decayed stumps seemed to promote germinant emergence. Exposure of mineral soil resulted in germination rates that were 4.3–6.9 times higher than in undisturbed conditions during the first growing season.     

Microsite patterns of conifer seedling establishment and growth in a mixed stand in the southern Alps

In many forests of the Alps, permanent forest cover and, therefore, its continuous renewal is the main silvicultural goal. Regenerating these forests must be based on a sound understanding of the ecology of the tree species in question. The regeneration of silver fir (Abies alba Mill.) and Norway spruce (Picea abies (L.) Karst.) in the upper montane zone of the southern central Alps has so far received little attention. The aim of this study was, therefore, to characterize the ecological niche for the establishment and growth of silver fir and Norway spruce seedlings in this zone. The study was conducted as a case study on a plot of 1.5 ha in the upper montane zone of southern Switzerland, at an altitude of 1380 m. The basal area of the mixed Norway spruce–silver fir stand was 43.7 m² ha⁻¹. Seedlings were censused on 375 plots of 1.0 m² area. Seedling density was 1.30 ± 4.25 m⁻² (mean ± standard deviation) for Norway spruce and 0.87 ± 1.50 m⁻² for silver fir. Logistic regression models were used to test the effect of microsite characteristics on seedling occurrence, and general linear models for effects on seedling height growth and biomass increment. Most seedlings received less than 10% light (photosynthetic photon flux density) as compared to values in the open. Silver fir occurrence was positively related to microsites at the edge of canopy trees, but unrelated to ground cover type, light and micro-relief. Norway spruce occurrence was only, and positively, related to the presence of mosses. Height growth and biomass increment of seedlings of both species were only loosely correlated with microsite conditions. Seedlings without canopy cover grew faster than those under canopy cover, probably as a result of light and moisture limitations under the canopy of adult trees. Diffuse radiation was positively correlated with average annual biomass increment of silver fir, but not of Norway spruce seedlings. In general, the results suggest that silver fir seedlings have less specific microsite requirements than Norway spruce seedlings in terms of ground cover. They are also more shade tolerant, and therefore, grow faster than Norway spruce in low-light environments of the upper montane zone of the southern Alps. While the microsite concept can be helpful in designing silvicultural operations, it has limitations when only patch characteristics that are easy to assess are used, and others neglected. Finally, our study suggests that counting the number of green shoots is a promising method to quickly and non-destructively estimate the biomass of a great number of small seedlings.     

Dynamics of seedling establishment and survival in uneven-aged boreal forests

The aim of this study was to describe the dynamics of seedling establishment and development in spruce-dominated uneven-aged boreal forests. The study was based on empirical data from 15 stands with permanent plots, which had been intensively monitored for 10 years in southern Finland. All trees (height > 1.3 m) were measured every fifth year. Regeneration was measured on 64 permanent sample plots (4 m²) in each stand. The establishment of first-year seedlings was analysed on a yearly basis. The survival and development of older Norway spruce (Picea abies (L.) H. Karst.) seedlings were analysed based on observations made every five years. The establishment of spruce seedlings was closely correlated with the abundance of seed crops. Seedling cohorts originating from abundant seed crops were clearly detectable in the development of seedling height distributions over time. It took about 15 years for spruce seedlings to reach a height level of 15-30 cm. Local basal area had hardly any effect on the emergence or survival of small spruce seedlings, while the number of higher spruce seedlings decreased with increasing local basal area.     

From planting to 26 years of age – actual growth and estimated volume scenarios for spruces and pines

The objective of this study was to compare the survival and volume of conifer stands at 26 years of age with their status at planting. Survival, growth and damage were studied in eight clear felled stands regenerated in 1972. Five of the areas were planted with Norway spruce (Picea abies (L.) Karst.) and three with Scots pine (Pinus sylvestris L.). The plantings were examined in 1972 and 1974. In 1974, the number of living undamaged planted seedlings was low (10–15%). However, the number of undamaged seedlings was supplemented by naturally regenerated conifer and birch seedlings. The total number of undamaged seedling in 1974 was equivalent to 20–30% of the number of seedlings planted. In 1998, the main species in three stands had changed from Norway spruce to Scots pine, and in one stand from Norway spruce to birches. Actual volume in 1998 for the stands was compared to stand volume generate according to five scenarios based on recommended and actual seedling number in 1972 and 1974. The actual volume was 64% of that expected if the recommended number of trees had been planted. Naturally regenerated Scots pine and Norway spruce increased stand density in 1998. The actual volume was 37% higher than the average volume in the surrounding county. On average, 36% of the trees were damaged. More than 50% of the total damage was caused by moose (Alces alces L.). For Scots pine, moose or other browsing animals damaged 30% of the trees. The results of this study indicate that the 1998 volume was higher than expected, considering the low number of undamaged seedlings in 1974. This was mainly due to the large amount of naturally regenerated plants. In addition, the results indicate that the volume could have been higher if the initial conditions had been better. Despite the low number of undamaged seedlings in 1974, seven of the eight studied stands produced a higher volume than the average stand for the region. In practise, high numbers of seedlings should be planted on scarified areas. In most cases there will be a supply of naturally regenerated seedlings.     

Interference of bracken (Pteridium aquilinum L. Kuhn) with Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies L. Karst.) seedling establishment

The nature of interference of bracken with Scots pine and Norway spruce seedling establishment was considered in three field experiments. In a seeding experiment, it was found that Scots pine germination was highest on exposed mineral soil and lowest when intact bracken litter and humus were present, suggesting adverse effects of litter and humus on pine regeneration probably due to phytotoxicity. In a second experiment, smothering by bracken caused high mortality of Scots pine seedlings while Norway spruce seedlings were relatively unaffected. Mortality for both Scots pine and Norway spruce seedlings was low when planted in a adjacent Scots pine-bilberry stand with no bracken. Annual shoot growth of Norway spruce was higher in bracken than in Scots pine-bilberry vegetation while no differences in shoot growth between these two vegetation types occurred for Scots pine. In a third experiment, activated carbon was added to the ground under Norway spruce seedlings planted in bracken to adsorb possible phytotoxic compounds released by bracken. The addition of carbon had no effect on seedling mortality or growth rate, indicating that the seedlings were not susceptible to allelochemicals released by bracken. Since large Norway spruce seedlings were relatively unaffected by bracken interference in this study, artificial regeneration with containerized Norway spruce seedlings is suggested to achieve an acceptable conifer tree establishment on clear-cuts invaded by bracken.     

Relationships between insect diversity and habitat characteristics in plantation forests

A methodological approach to the identification of biodiversity indicators in commercial forest stands is illustrated by analysis of the relationships between syrphid (hoverflies) and carabid (ground beetles) community composition and diversity, and stand structure and field layer vegetation. Data were collected from 12 commercial forest sites encompassing a range of climatic conditions and different crop types (Scots pine, Sitka spruce, Norway spruce and Corsican pine) across the UK. Comparisons were also made between unmanaged semi-natural Scots pine woods and Scots pine plantations. For both syrphids and carabids, no differences in species richness and diversity were recorded between semi-natural stands and plantations; one rare syrphid considered to be restricted to semi-natural pine stands was also found in spruce. Syrphid species diversity and richness was higher in southern spruce sites than in the northern sites. Northern sites had distinctive carabid communities, as did sites in the New Forest, a large ancient woodland in southern England. Of the measured habitat variables, vertical stand structure showed the best correlation with species richness and diversity of both carabids and syrphids. Richness and diversity were less in stands with high vertical cover values for canopy layers. Stands with higher field layer cover supported greater syrphid diversity, but lower carabid diversity. Measures of stand structure could be used as potential indicators of syrphid and carabid diversity, but additional habitat parameters also need to be tested.     

Germination and seedling establishment of Norway spruce (<Emphasis Type="Italic">Picea abies</Emphasis>) after clear-cutting is affected by timing of soil scarification

Natural regeneration of Norway spruce (Picea abies (L.) Karst.) is a relatively common practice in Norway on medium to low site indices. However, seedling establishment is often hampered by rapid regrowth of competing vegetation in scarified patches. The aim of our study was to examine the effect of coordinating scarification towards an expected seed-fall, by studying germination and seedling establishment in scarified patches of different age (fresh, one- and two-year-old). The experiment was conducted in two stands in southeast Norway that were clear-cut in 2007. Scarification was applied to subplots in autumn 2008–2010. To simulate seed-fall, seeds were sown in fresh scarification patches in spring 2009–2011, in one-year-old patches in 2010 and 2011, and in two-year-old patches in 2011. Both germination and seedling survival were negatively affected by the age of the scarified patches. Germination was higher, and mortality lower, at the small fern woodland site, compared with the bilberry woodland site. Sowing in fresh patches also resulted in increased height and root collar diameter of the seedlings compared with sowing in older patches. It is likely that the competing vegetation both on the site and in the scarification patches affected the growth of the seedlings. In conclusion, the age of the scarified patches affected both germination and mortality, as well as early growth of the seedlings.     

Effects of site preparation on emergence and early establishment of Larix olgensis in montane regions of northeastern China

Larix olgensis is one of the important commercial timber tree species in northeastern China, but it does not naturally regenerate in L. olgensis forests even after cutting. To identify the causes of the regeneration failure, the authors tested the effects of site preparation in a thinned stand and in a clearcut on the emergence and early establishment of L. olgensis seedlings. The results indicated that there was no correlation between cumulative emergence percentage (CEP) of L. olgensis seedlings and canopy openness, but the CEP was significantly affected by site preparation (P < 0.001). The average seedling survival rates after treatments consisting of removing both litter cover and understory vegetation cover were significantly higher in the clearcut than in the thinned stand (P < 0.001). Without site preparation, all seedlings in both the thinned stand and the clearcut disappeared within the first growing season, meaning that ground cover (understory vegetation cover and litter cover) was not beneficial to seedling survival. The average growth of seedlings exhibited significant differences between the thinned stand and the clearcut, and among the site preparations (P < 0.001). Intensely thinned stands without ground cover improved seedling growth. These results imply that light and ground cover might be the main factors limiting the early establishment of L. olgensis seedlings in managed plantation forests     

The effect of scalping on seedling establishment after seed tree cutting of Scots pine stands in drained peatlands in northern Finland

We studied the establishment of natural seedlings after seed tree cutting and scalping in two drained Scots pine peatland stands in northern Finland (Simo and Sievi). Approximately, 50?ha^?1 of Scots pine stems were retained on both sites. During the six subsequent years, five seedling surveys were conducted. The effect of site preparation was analysed using generalized linear mixed models (GLMM). The mean density of pine seedlings increased more quickly in the scalped plots and was ca. 1.1?m^?2 after six years compared to 0.87?m^?2 in the non-scalped plots, on average. Scalping increased the number of pine seedlings 2-fold compared to that in non-treated plots in Simo, however, in Sievi non-treated plots had 30% higher pine seedling density after six years. Downy birch seedling density was 9–10-fold compared to that of non-treated plots in both sites. In terms of seedling density, seed tree cutting led to satisfactory pine regeneration after just two growing seasons in drained pine peatlands, even without scalping. Scalping significantly increased the number of pine seedlings, but it also enhanced the germination of birch seedlings strongly.     

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

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

Natural regeneration after gap cutting in Scots pine stands in northern Finland

The objective of the study was to examine the success of regeneration in gaps of variable size in pine-dominated stands in terms of seedling density. It was based on an experiment in central Lapland containing circular clear-cut gaps of 20, 40 and 80 m in diameter on typical sub-xeric and xeric pine sites with site preparation (patch scarification). The observation period covered the first five years after cutting. The average number of seedlings that had emerged after gap cutting was about 22 000?ha^?1 for pine and 7 000?ha^?1 for birch. The proportion of regeneration sample plots without any pine seedlings was less than 10%. The seedling density diminished constantly with greater distance from the edge stand but indicating sufficient density up to the largest gap size in the study (diameter 80 m, area ca. 0.5?ha). Site preparation promoted regeneration remarkably, and we conclude that successful regeneration would be achieved in most cases with soil scarification exposing just 10–20% of the soil surface.     

The response of ground vegetation to structural change during forest conversion in the southern Black Forest

The aim of this study is to investigate the effects of forest conversion on forest floor vegetation. ‘Ecological’ forest conversion, as defined by an interdisciplinary southern Black Forest project group, describes the transformation of even-aged spruce (Picea abies L. H. Karst.) stands to structured continuous cover forests consisting of spruce (Picea abies), silver fir (Abies alba Mill.) and beech (Fagus sylvatica L.). In order to analyse the conversion process, four conversion stages were defined in a conceptual forest development model. Four forest districts deemed to be representative of the southern Black Forest region were selected for the study. The ground vegetation was initially classified independently from the stand structures. Subsequently, the relationship between stand structures, as determined by development stage, and ground vegetation was analysed. It was revealed that forest conversion modified the ground vegetation. The main factors influencing ground vegetation were the influence of broadleaves, predominately beech (F. sylvatica), on the canopy composition and litter coverage on the one hand; and the canopy coverage of spruce, the proportion of needle litter and the associated light penetration on the other. The prevalence of moss and vascular plant species preferring acidic sites found in spruce forests decreased during the transition process, whereas, species requiring a moderate base supply increased in abundance. The continuous cover forest representing the final stage of conversion increasingly contained a mixture of ground vegetation species normally associated exclusively with either conifer or deciduous forest. Due to the fact that there is an associated ground vegetation specific to the different stages of forest conversion in stands dominated by Norway spruce or European beech and a mixture in the latest conversion stages, large-scale forest conversion should be avoided in favour of management promoting a diversity of silvicultural goals and treatments in neighbouring stands. Only a variety of treatments ensures the maintenance of floristic diversity in the long-term.     

Modelling above and below ground carbon dynamics in a mixed beech and spruce stand influenced by climate

Tree growth and carbon dynamics are important issues especially in the context of climate change. However, we essentially lack knowledge about the effects on carbon dynamics especially in mixed stands. Thus, the objective of this study was to test the effects of climatic changes on the above and below ground carbon dynamics of a mixed stand of Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica L.) by means of scenario simulations. To account for the typical tree interactions in a mixed-species stand a spatial explicit tree growth model based on eco-physiological processes was applied. Three different climate scenarios considering altered precipitation, temperature, and radiation were calculated for an unthinned and a thinned stand. The results showed significant changes of above and belowground biomass over time, especially when temperature and radiation were increased additionally to decreased precipitation. The reduction in biomass increments of Norway spruce were more attenuated above than below ground. In contrast, the results for beech were the opposite: The belowground increments were reduced more. These results suggest a shift in the species contribution to above and belowground biomass under dryer and warmer conditions. Distinct effects were also found when thinned and unthinned stands were compared. A reduced stand density changed the proportions of above and below ground carbon allocation. As a main reason for the changed growth reactions the water balance of trees was identified which lead to changed biomass allocation pattern. This article belongs to the special issue “Growth and defence of Norway spruce and European beech in pure and mixed stands”.     

Frequencies and patterns of browsing by large herbivores on conifer seedlings

Browsing by large herbivores on planted and naturally regenerated conifer seedlings (Picea abies and Pinus sylvestris) was recorded in 104 clear‐cuts in east‐central Sweden during 1990 and in 47 clear‐cuts in 1991. The number of seedlings browsed and the browsing patterns were analysed in relation to seedling type. Browsing frequencies were also compared between forest stands with different site productivities and subjected to different management practices. The variation in the number of seedlings browsed in 1990 was explained mainly by seedling category. Among both planted and naturally regenerated seedlings, pine was browsed more than spruce. Two‐year‐old containerized seedlings of spruce was browsed more than 4‐yr‐old bare‐rooted spruce. In 1991, browsing was more equal among species and seedling types. Number of seedlings with their leader browsed and the amount of biomass left after browsing differed significantly between seedling types. Seedlings that had been browsed in 1990 experienced significantly higher browsing frequencies in 1991 when compared with unbrowsed seedlings. The effects of stand characteristics were not found to be significant.     

Pre-commercial thinning,birch admixture and sprout management in planted Norway spruce stands in South Sweden

Early management of the regenerated seedlings shapes the future stand properties. To address these issues, pre-commercial thinning (PCT) and control treatments were applied to planted Norway spruce (Picea abies L. Karst) and naturally regenerated birch (Betula pendula Roth., Betula pubescens Ehrh.) stands in forest experiments in southern Sweden (lat. 56–57?N) containing 1.1–5.5?m tall saplings. The treatments were retention of 1000 or 2000?stems?ha^?1 of Norway spruce, with no birch or birch at 1000?stems?ha^?1. Treatments were replicated with and without annual removal of birch sprouts from stumps. The periodic annual increment (PAI) over five years was calculated for total stand volume and individual trees. The mean PAI of dominant trees was significantly higher both following all PCT treatments than controls, and following low rather than high-density PCT. Birch retention did not affect growth of the dominant trees but PAI was lower in plots with uncontrolled sprouting. The PAI of birch was significantly higher in low-density Norway spruce plots than in control plots and the high-density plots. The treatment response was significant even in stands with initial heights of only 1–2?m.     

Stand density and growth of Norway spruce (<Emphasis Type="Italic">Picea abies</Emphasis> (L.) Karst.) and European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.): evidence from long-term experimental plots

On the basis of nine Norway spruce (Picea abies (L.) Karst.) and ten European beech (Fagus sylvatica L.) thinning experiments in Germany, for which both residual and removed stock had been registered first during 1870, I scrutinize how moderate and heavy thinning from below (B-, C-grade) affects the production of merchantable volume compared with light thinning (A-grade). In relation to A-grade, cumulative merchantable volume (CV) of B- and C-grade amounts in average to 103–107% in juvenile and to 97–102% in mature Norway spruce stands. The corresponding findings for European beech are 101–106% and 94–102%. CV of individual stands varies between 89% and 130% for Norway spruce and 73% and 155% for European beech (CV of A-grade = 100%). These findings are substantiated by the relation between stand density (SDI) and periodic annual increment (PAI). On the B- and C-grade plots of spruce and beech, respectively, SDI was reduced down to 41–91% and 31–83% of the A-grade. When SDI is reduced in young stands, PAI follows a unimodal curve. Norway spruce’s PAI culminates in 109% if SDI is reduced to 59%; European beech’s PAI culminates in 123% when density is reduced to 50%. Whereas Norway spruce’s growth reacts most positively on thinning under poor site conditions and with increment reduction on favourable sites, European beech behaves oppositely. With stand development the culmination point of the unimodal relation moves towards maximum density, so that in older stands PAI follows the increasing pattern, which is the left portion of a unimodal curve. A model is presented which apparently unifies contradictory patterns of stand density–growth reactions by integrating relative stand density, average tree size and site fertility effects, and makes the findings operable for forest management.     

Stand and landscape level effects of a major outbreak of spruce beetles on forest vegetation in the Copper River Basin,Alaska

From 1989 to 2003, a widespread outbreak of spruce beetles (Dendroctonus rufipennis) in the Copper River Basin, Alaska, infested over 275,000 ha of forests in the region. During 1997 and 1998, we measured forest vegetation structure and composition on one hundred and thirty-six 20-m × 20-m plots to assess both the immediate stand and landscape level effects of the spruce beetle infestation. A photo-interpreted vegetation and infestation map was produced using color-infrared aerial photography at a scale of 1:40,000. We used linear regression to quantify the effects of the outbreak on forest structure and composition. White spruce (Picea glauca) canopy cover and basal area of medium-to-large trees [≥15 cm diameter-at-breast height (1.3 m, dbh)] were reduced linearly as the number of trees attacked by spruce beetles increased. Black spruce (Picea mariana) and small diameter white spruce (<15 cm dbh) were infrequently attacked and killed by spruce beetles. This selective attack of mature white spruce reduced structural complexity of stands to earlier stages of succession and caused mixed tree species stands to lose their white spruce and become more homogeneous in overstory composition. Using the resulting regressions, we developed a transition matrix to describe changes in vegetation types under varying levels of spruce beetle infestations, and applied the model to the vegetation map. Prior to the outbreak, our study area was composed primarily of stands of mixed white and black spruce (29% of area) and pure white spruce (25%). However, the selective attack on white spruce caused many of these stands to transition to black spruce dominated stands (73% increase in area) or shrublands (26% increase in area). The post-infestation landscape was thereby composed of more even distributions of shrubland and white, black, and mixed spruce communities (17–22% of study area). Changes in the cover and composition of understory vegetation were less evident in this study. However, stands with the highest mortality due to spruce beetles had the lowest densities of white spruce seedlings suggesting a longer forest regeneration time without an increase in seedling germination, growth, or survival.     

White spruce recruitment distances relative to lodgepole pine in southern Yukon,Canada

Eleven seral, postfire forest stands in southern Yukon (Canada) were sampled to determine where western white spruce (Picea albertiana ssp. albertiana) seedlings occurred with respect to distance to the nearest lodgepole pine (Pinus contorta var. latifolia) tree. Seedling-to-nearest tree distances were assessed at 10-cm increments up to 220?cm. On average, seedlings occurred 54?cm from the nearest pine (n?=?490), but peak frequencies were 20–50?cm away, compared to a potential separation distance of 103?cm. Greatest average seedling density occurred 10–20?cm from pine (0.81?m^?2), with values between 10 and 120?cm decreasing logarithmatically with increasing distance from pine (r?=?0.994, p?n?=?11). Spruce seedling densities were <0.02?m^?2 beyond 120?cm. The differences in frequency, which represented moderately strong aggregation (Clark-Evans Index 0.34–0.52), and density suggest greater spruce recruitment near lodgepole pine was facilitated by more favorable ecological conditions than further away, for example, greater nutrient availability. The bias in seedling-to-nearest tree distances occurred regardless of stand age (57–165 years), pine density (1599–5935 stems?ha^?1), or understory vegetation type, although the bias may be weakened by the abundant presence of feathermosses (Hylocomium splendens) on the forest floor.