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.     

Regeneration niches of Norway spruce (<Emphasis Type="Italic">Picea abies</Emphasis> [L.] Karst.) saplings in small canopy gaps in mixed mountain forests of the Bavarian Limestone Alps

Natural regeneration measurements are the main silvicultural objective in overaged protective forests of the Bavarian Limestone Alps. While manifold problems with these stands, especially the impact of browsing, are widely recognised, the regeneration niches of Norway spruce (Picea abies [L.] Karst.) are insufficiently known. The purpose of this study was to determine favourable combinations of site factors for the development of spruce in small, unfenced canopy gaps, located on Aposerido-Fagetum caricetosum albae forest sites. We recorded the occurrence of spruce saplings (as dependent variable) and of six site factors (as independent variables) on 480 0.5 m²-subplots. In addition, we estimated the coverage of six acid adapted plant species to determine correlations with the humus depth. A binary logistic regression analysis was used to predict the probability of the occurrence of a spruce sapling in dependency of the different site factors. Supported by other studies, we assumed that the supply of solar radiation was adequate for the sufficient regeneration of spruce within the canopy gaps. Other site factors significantly determined the regeneration niches of spruce saplings. More spruce saplings were found near hindrances and on rough surfaces than would be expected from a random occurrence of saplings. These microsite types may have characteristics, especially protection against snow gliding that promotes spruce establishment. A calculated “hindrance index”, which accounted for the number, and the distance of surrounding hindrances might be a good specific value to describe the influence of hindrances on steep slopes. The sapling establishment decreased on thin humus layers. Our assumption for the sites was that thick organic layers might represent a good seedbed for spruce. Decayed dead wood was scarce, but was exceedingly favoured by spruce saplings. Results obtained suggest that the natural regeneration establishment of spruce on steep slopes can be successfully influenced by site factors which inhibit the influence of snow gliding. According to a “positive microsite” concept, we recommend for artificial regeneration measurements with spruce, microsites close to hindrances (e.g. stumps, downed trees) and Vaccinium myrtillus as a predictor for thick, acid humus layers.     

Influence of artificial gaps in tropical forest on survival, growth, and Phytolyma lata attack on Milicia excelsa

Milicia excelsa and M. regia are important timber species in moist tropical areas of Africa. They have not been successfully grown largely because of attacks by gall-forming psyllids in the genus Phytolyma. Our objectives were to evaluate the growth of planted Milicia seedlings and incidence of psyllid attack in small (4.2 m²), medium (18.5 m²) and large (>500 m²) artificial gaps in the Bobiri Forest Reserve in the Moist Semi-Deciduous Forest of Ghana. After 13 months, height and diameter growth of Milicia seedlings were significantly greater in the medium and large gaps than in the small gaps. Insect attacks occurred first and most severely in the large gaps, but spread to gaps of all sizes between the 11th and 13th months after planting. While gap size significantly affected the susceptibility of seedlings to psyllid attack, it is not the only factor important in determining susceptibility of Milicia excelsa under field conditions. We conclude that gap sizes in the range of 10–50 m², where irradiances are from 30–60% of full sunlight, in forests similar to those at the study site, seem to be most suitable for regeneration of Milicia.     

The effects of clearcutting on snowshoe hare (Lepus americanus) relative abundance in central Labrador

To assess the effects of clearcutting on snowshoe hare (Lepus americanus) relative abundance, we surveyed pellets in 1 m² circular plots and, vegetation and browse surveys in 4.5 m² circular plots among four different aged clearcut (30, 20, 10, 5 years post-harvest) and mature forests (>150 years old) in central Labrador, Canada. Data were modelled at three grain sizes: transect (4400 m²), plot (314 m²) and subplot scales (4.5 m²). Betula papyrifera, distance from mature forest edge, tree and herb cover as well as remotely sensed forest inventory data were used as predictors for hare pellets. We found pellet abundance was 5 and 37 times greater (new and old pellets, respectively) in clearcut stands 30 years old than the next highest in 20 year old cuts. There were few hare pellets in the remaining stand ages. B. papyrifera was the most proportionately used browse species and most important of our fine-detailed vegetation in predicting hare pellets. The coarse-detailed, forest inventory and topographic data better predicted hare pellets than the fine-detailed vegetation data.     

Management of advanced regeneration in secondary forests of the Brazilian Amazon

Gap formation is a critical process for plant establishment that may be absent or infrequent in second-growth forests, negatively influencing their ability to become diverse mature forest. I simulated treefall gaps within a 10-year-old secondary forest established on a clear-cut area, because cut areas may have a greater potential for economic exploitation than areas used for pastures and shifting cultivation. I explored the effects of varying degrees of canopy opening on the growth of plants already established as advanced regeneration. Five intensities of canopy opening were created: total removal (100%), 80–90%, 50–60%, 20–30% and a 0% control. Only trees >5 cm DBH were cut. The established plant community was diverse, and comprised 37 families and 126 species and morpho-species, of which 72% had some use reported in the literature. I found that the canopy opening treatments resulted in very little disturbance to the established seedling and sapling community, with no significant increase in mortality, but had a strong positive effect on their growth. Light levels reaching the understory were increased from 1% of incident photosynthetic active radiation in control plots to 35% in 100% canopy opening plots. During the first year, differences in plant growth were not significant, but by the second year, differential plant growth associated with canopy opening became evident. Most seedlings (51%) grew approximately 25 cm in height, and approximately one quarter grew more than 25 cm. Only 2.6% of the seedlings decreased in height, and 12% did not grow at all. Fifty percent canopy removal resulted in an almost threefold increase in plant height compared to control plots (0.52 ± 0.4 m and 0.20 ± 0.2 m, respectively), and was not significantly different from 80 or 100% removal. This indicates that intermediate levels of canopy removal had positive effects and total removal was unnecessary to stimulate the best growth responses. I conclude that regeneration of diverse mature forest can be accelerated under some conditions by partial removal of secondary forest canopy.     

Species composition and dynamics in two hardwood stands in Vermont: a disturbance history

Stand composition and structure utilizing stem analysis was studied in two hardwood stands in Vermont. In a mixed hardwood stand with some white pine and hemlock, a major entry of new trees in the main canopy seems associated with harvesting coincident with land exchange. More recent partial cuttings have promoted establishment of new seedlings or development of suppressed advanced-growth shade-tolerant beech, (Fagus grandifolia, Ehrh.), hophornbeam (Ostrya virginiana, (Mill) K. Koch), and striped maple (Acer pensylvanicum, L.). Very few sugar maple and red oak seedlings and saplings are present.

In a northern hardwood stand some red spruce (Picea rubens, Sarg.), that were 240 to 306 years old, became established before any known harvest, and exhibited release following harvests of the mid-1800s. This major harvest, coupled with the differential growth between spruce and hardwoods, and seed/seedling availability, resulted in a major change in stand composition. Trees now in the main canopy of sampled stands appear to have either been released or newly established following various harvests. Harvests have been of such frequency that natural disturbances seem insignificant. Many of the competitive understory species have become abundant following harvests of the 1960s and 1980s and may have been present as advanced growth and responded to the release. Following the harvest of 1981–1982, abundant yellow birch became established on skid trails. Elsewhere in the stand, yellow birch seedlings and saplings are only in great abundance in areas that were possibly sizable gaps following earlier harvesting.

The dynamics of tree entry and growth in gaps of small or large size probably occur in a similar way in many other stands of the region. Though the sampling of this study is limited, there is no suggestion of continuous tree establishment at any particular location, the new age classes seem associated with either a gap or stand replacing disturbance attributed to harvesting.     

The effects of selective logging on stand structure and the regeneration of subboreal forests in Hokkaido, northern Japan

We investigated the effects of selective logging on stand structure and regeneration in selectively logged subboreal forests in Taisetsuzan National Park in Hokkaido in northern Japan. The basal area decreased and the size structure of trees altered in the stands studied due to repeated, intense selective logging, in which larger trees were cut down as a priority. Sapling density in the stands was much lower than that in primary forests. In the simple and multiple regression analyses that were used to estimate the effects of selective logging on sapling density, sapling density had a significant positive correlation with tree density and had little correlation with the density of logged stumps or the height ofSasa (dwarf bamboo) growing on the forest floor. These results suggest that the establishment sites around canopy trees influenced the establishment of saplings, rather than the gaps caused by selective logging. However, both the coefficient of determination and the standardized partial regression coefficient of multiple regression analysis were higher for the stand with a dense cover ofSasa than for the stand with a sparse cover ofSasa. Thus, the success of regenerating forests with selective logging depends on both the site of advanced regeneration and the light conditions that regulate growth.     

The ecology of tree regeneration in mature and old forests: combined knowledge for sustainable forest management

We focused our attention on quantifying the factor complex of forest regeneration in 423 mature and old stands with contrasting environmental conditions. We recorded the microhabitat selection of tree recruits, the frequency of tree seedlings, and evaluated the drivers of sapling abundance and diversity. The majority of forest regeneration was established on undisturbed forest floor. Dead wood was a frequent substrate in spruce-(co)dominated forests. Seedling frequency within a stand was related to the site-type specific productivity gradient of stands—pine seedlings were common in low-productivity and spruce in high-productivity boreal forests. Seedlings of temperate broad-leaved trees dominated in productive boreonemoral forests, except for oak, which showed a uniform distribution of abundance in all forest site-types. Sapling abundance was dictated by forest site-type, and facilitated by stand diversity, variability in stand closure, lying dead wood, abundant moss, and a thick organic layer. Only in boreal forests was sapling abundance suppressed by the abundant spruce and younger trees. Upon considering the relationship between sapling abundance and species richness, sapling diversity was dependent on forest site-type, suppressed by stand density and dead wood (old gap) abundance, and facilitated by stand diversity. In addition, boreonemoral stands, competition from herbs, and facilitation by mosses occurred. The observed pattern of tree recruitment points to the importance of top-down effects of the overstory, competing or facilitating interactions with forest floor vegetation, and availability of regeneration microhabitats, which in complex make their ecology comparable with forest herbs. Natural forest regeneration can be enhanced if silvicultural methods support mixed stands and enhance field layer diversity. Oak can provide the universal tree species to improve stand structure over a wide range of habitats.     

Regeneration patterns of Eastern white pine (Pinus strobus L.) in hardwood-dominated forests in Virginia, USA

Eastern white pine (Pinus strobus L.) is a moderately shade-tolerant species that co-occurs with hardwood tree species in many forests of the eastern United States, as well as in pure stands. The species is valued for its timber, as well as for wildlife and recreation. Regeneration of this species is somewhat unpredictable and often occurs in patches of similarly-aged cohorts. We described the regeneration patterns of this species and examined their relation to environmental variables within hardwood forests of southwestern Virginia, USA. An average of 5.3 white pine patches per ha were observed in this study. The majority of patches consisted of saplings (85%), with 9% of patches in pole size classes, and 6% in seedling size classes. The average density of patches was 43.5 stems with an average age of 20 years. The size of patches averaged 80.6 m². The total density of seedlings and the number of regeneration patches of all sizes of regeneration (seedlings, saplings, and poles) in plots was related to the surrounding density of large white pine trees (potential seed trees). The density of seedlings or patches was not significantly related to current vegetation cover or soil surface cover variables, but more than half of regeneration patches were located in or adjacent to old canopy gaps, most of which were old logging gaps. While seedling regeneration may occur within the understory of these forests near seed trees, advancement to the sapling and pole stage appears to be associated with canopy gap formation.     

Productivity and economics of mixed two-storied spruce and birch stands in Southern Finland simulated with empirical models

The profitability of growing a naturally emerged birch (Betula pendula Roth or Betula pubescens Ehrh.) overstory in a young Norway spruce (Picea abies (L.) Karst.) plantation was examined with empirical stand structure, growth and yield, logging cost, and logging damage models. In the projected alternatives, an overstory of either birch species was thinned to 200–1000 stems per ha at the age of 15 years and retained for another 15 years. Development of the remaining spruce stand was simulated up to rotation age (70–85 years). Alternative treatments included removing the overstory completely at 15 years, and managing a pure spruce stand that was kept free of birch throughout.

Growing a birch overstory of 200–1000 stems per ha up to age 30 years resulted in a 61–93 m³ ha⁻¹ or 9.1–16.8% yield loss for the spruce stand due to growth retardation, and a mortality of 382–498 out of 1900 stems per ha through logging damage. This was compensated for or exceeded by the additional yield of the birch (54–173 m³ ha⁻¹) except for the lowest stocking (200–400 stems per ha) alternatives with B. pubescens. Treatment regimes with a birch overstory were clearly the most profitable alternatives, yielding up to 151% (B. pendula) and 113% (B. pubescens) of the net present value of the pure spruce alternative, at 4% interest rate. Removing the birch overstory already at 15 years was the least profitable alternative with 79 and 83% net present values, respectively. The most profitable treatment with current technology, price, and cost structure appears to be to grow 500–800 birch per ha up to about age 40 years for B. pendula and 45–50 years for B. pubescens.     

The impact of hemlock looper (Lambdina fiscellaria fiscellaria (Guen.)) on balsam fir and spruce in New Brunswick, Canada

In 1989, the first recorded outbreak of hemlock looper (Lambdina fiscellaria fiscellaria (Guen.)) occurred in New Brunswick, Canada. Data were collected from ten plots established in an area infested from 1992–1994, to assess impacts of hemlock looper. Ocular and branch sample assessments of current defoliation and ocular assessments of total defoliation (all age classes of foliage) were conducted for balsam fir (Abies balsamea [L.] Mill.), white spruce (Picea glauca [Moench] Voss), and black spruce (Picea mariana (Mill.) B.S.P.). Stand response was assessed and related to cumulative defoliation. Ocular assessments were found to accurately estimate defoliation, which was significantly related to tree mortality. Ninety-two percent of balsam fir trees that had cumulative defoliation >90% died. Mortality of balsam fir was significantly (p < 0.05) related to tree size, in both lightly and severely defoliated plots; trees with DBH <11 cm sustained 22–48% higher mortality than larger trees. Mortality of balsam fir, in terms of both percent stems/ha and m³/ha merchantable volume, increased exponentially in relation to three estimates of cumulative (summed) plot mean defoliation. The strongest relationships (r² = 0.75–0.79) were between mortality and the ocular defoliation assessment for 1990–1993 foliage. Tree mortality caused by the looper outbreak ranged from 4–14% stems/ha in lightly defoliated and from 32–100% in severely defoliated plots; merchantable volume killed was 3–14 m³/ha and 51–119 m³/ha, respectively. Relationships between mortality and defoliation were similar when defoliation was assessed for 1987–1993 and 1990–1993 foliage age classes.     

Pits and rocky microsites in a subalpine forest stand facilitate regeneration of spruce saplings by suppressing dwarf bamboo growth inside a deer-proof fence

We determined patterns of microsite suppression in dwarf bamboo Sasa nipponica when grazing deer were absent. This bamboo species is able to outcompete Hondo spruce (Picea jezoensis var. hondoensis) saplings under many environmental circumstances. We set up two 10 × 100 m plots inside a deer-proof fence within a subalpine forest on Mt. Ohdaigahara, central Japan, and two similarly sized plots outside the fence. Within the plots, we surveyed microsites where spruce saplings grew. We measured height and shoot elongation of all spruce saplings, and culm height and cover ratios of dwarf bamboo growing around each spruce sapling. Spruce sapling density and average height were higher inside the deer-proof fence than outside, as were bamboo height and cover. Thus, there was a negative effect of deer browsing on vegetation parameters outside the fence and a suppression of the negative effect of bamboo on spruce sapling growth inside the fence. Spruce sapling height was higher in tree-fall pits than in other microsites inside the fence, whereas both dwarf bamboo height and cover were lower in pits and rocky sites than elsewhere. In soil and collar microsites, spruce sapling shoot growth was lower and bamboo height and cover were higher than in pits and rocky sites. Inside the fence, dwarf bamboo cover was high, but pits and rocks suppressed its growth, allowing spruce saplings to flourish. To restore heavily damaged spruce forests with advanced saplings, it will be necessary to construct deer-proof fences and create and maintain microsites with pits and rocks.     

Variable-density thinning promotes differential recruitment and development of shade tolerant conifer species after 17 years

Promoting patchy recruitment of shade tolerant tree species into the midstory is an important step in developing structural diversity in second-growth stands. Variable-density thinning (VDT) has been proposed as a strategy for accelerating structural diversity, as its combination of within-stand treatments (harvest gaps, thinning, and non-harvested skips) should create variable overstory and understory conditions. Here we report on western hemlock (Tsuga heterophylla (Raf.) Sarg.) seedling and sapling densities in five mixed-conifer stands and Sitka spruce (Picea sitchensis (Bong.) Carr.) seedling and sapling densities in two stands in western Washington at 3,7, 10, and 16–17 years after VDT. Additionally, we report on western hemlock advance regeneration growth and survival in two stands over 14 years. Western hemlock seedling density was highest in the thinned treatment but only significantly so in Year 10. In contrast, the gaps contained significantly more western hemlock saplings in Years 7 and 10 and significantly greater growth of western hemlock advance regeneration through Year 10. Skips embedded within the VDT did not differ significantly from unharvested reserves in terms of seedling or sapling densities of either species. Sitka spruce seedling density was highest in the gap and thinned treatments, but saplings were uncommon in all treatments. Collectively, these results indicate that our variant of VDT promoted patchy, midstory recruitment of western hemlock but failed to recruit Sitka spruce saplings in either stand where it established. Consequently, more intensive variants of VDT may be required to promote midstory recruitment of species less tolerant of shade than western hemlock.

     

Oak (<Emphasis Type="Italic">Quercus robur</Emphasis> L.) regeneration as a response to natural dynamics of stands in European hemiboreal zone

The oak (Quercus robur L.) regeneration intensity was assessed in the core area of the Białowieża National Park (BNP) in Poland with respect to the selected ecological factors. The emphasis was placed on the response of oak regeneration to disturbances, including the large-scale dieback of spruce stands. Defining their effect could help predicting the role of oak in naturally developing lowland forest ecosystems in the European hemiboreal zone. The results of the study challenge the opinion that the ‘lime-oak-hornbeam forest’ is a ‘climax’ community, confirming a very poor regeneration represented by only two saplings taller than 0.5 m per hectare. By contrast, in spruce-dominated communities, from 49 to 848 taller saplings per hectare were found. The occurrence of saplings was associated with discontinuous canopy of late seral stage of stands, as well as with large gaps. Most of the best quality grown-up oak saplings developed in the immediate neighbourhood of spruce logs. The results of the research indicate that ‘lime-oak-hornbeam forest’ (Tilio-Carpinetum) should be rather perceived as a transient community, evolved from relict, culturally modified, oak woodlands. Because spruce had become a dominating species only after abandoning in mid-1800s the historical regime of anthropogenic disturbances (involving frequent forest fires), the observed phenomena related to the disintegration of spruce stands had probably no precedent over the last 500 years. To confirm whether the massive decline of spruce stands will finally result in the successful establishment of the new canopy oak generation, both in Białowieża and other forests of hemiboreal zone, further research is needed.     

Tree allometry, leaf size and adult tree size in old-growth forests of western Oregon

Relationships between tree height and crown dimensions and trunk diameter were determined for shade-tolerant species of old-growth forests of western Oregon. The study included both understory and overstory species, deciduous and evergreen angiosperms and evergreen conifers. A comparison of adult understory species with sapling overstory species of similar height showed greater crown width and trunk diameter in the former, whether the comparison is made among conifers or deciduous trees. Conifer saplings had wider crowns than deciduous saplings, but the crown widths of the two groups converged with increase in tree height. Conifer saplings had thicker trunks than deciduous saplings of similar crown width, possibly because of selection for resistance to stem bending under snow loads. The results suggest that understory species have morphologies that increase light interception and persistence in the understory, whereas overstory species allocate their biomass for efficient height growth, thereby attaining the high-light environment of the canopy. The greater crown widths and the additional strength requirements imposed by snow loads on conifer saplings result in less height growth per biomass increment in conifer saplings than in deciduous saplings. However, the convergence in crown width of the two groups at heights greater than 20 m, and the proportionately smaller effect of snow loads on large trees, may result in older conifers equalling or surpassing deciduous trees in biomass allocation to height growth.     

Understory bamboo flowering provides a very narrow light window of opportunity for canopy-tree recruitment in a neotropical forest of Misiones, Argentina

Chusquea ramosissima is a native monocarpic bamboo species growing in subtropical forests of northeastern Argentina, which can dominate gaps and open forests in the region, particularly after human disturbance. This bamboo species started to flower in different areas of northeastern Argentina in 2001, with the flowering peak during 2002 and 2003 and small isolated flowering events still occurring until 2010. We studied the effects of C. ramosissima flowering and die-back on microclimate, litter decomposition, nutrient availability, sapling growth, abundance and regeneration of tree canopy species. We wanted to know how environmental conditions and ecosystem processes change through time after bamboo flowering and if bamboo die-back would favor regeneration of canopy trees. Twenty 50 × 50 m plots of flowering and non-flowering bamboo were permanently marked and vegetation dynamics as well as nutrient cycling and microclimate studies were performed. C. ramosissima die-back enhanced growth and reduced mortality rate of tree saplings during the first year after flowering. Only growth of tree saplings previously established was enhanced by the flowering event and tree-species richness and saplings abundance of canopy trees did not change as expected due to bamboo flowering. The short-term effect of tree saplings growth was likely due to incident solar radiation at the forest floor which doubled in the first year after the bamboo flowering event. Increased light availability at the forest floor simultaneously promoted the growth of other understory plants such as ferns, lianas and Piper spp. that rapidly colonized gaps and intercepted a percentage of the incident solar radiation after the first year, which together with an increased litter layer due to the senescence of the bamboo, may have inhibited establishment of new tree individuals and affected tree growth. Contrary to predictions, soil water, litter decomposition and soil nutrients were not significantly affected by bamboo flowering. Thus, successful tree regeneration in gaps following bamboo flowering appears to be restricted to a very narrow window of increased light availability (i.e., 1 year) before growth of other understory plants and rapid re-colonization of bamboo. Changes in resource availability, and the opportunity for overstory regeneration after bamboo flowering events appears to depend on climatic and community characteristics of the ecosystem where the flowering event occurs and also, on the flowering patterns and their synchronicity.     

Canopy tree development and undergrowth bamboo dynamics in old-growth Abies–Betula forests in southwestern China: a 12-year study

Interactions between forest canopy characteristics and plants in the forest understory are important determinants of forest community structure and dynamics. In the highlands of southwestern, China the dwarf bamboo Bashania fangiana Yi is an understory dominant beneath a mixed canopy of the evergreen Abies faxoniana (Rheder & Wilson) and the deciduous Betula utilis (D. Don). The goal of this study was to better understand the role of bamboo dominance, canopy characteristics, and periodic bamboo dieback on forest development. To achieve this goal, we measured tree seedling, tree saplings, and trees, forest canopy characteristics, and bamboo cover in permanent forest (n = 4) and gap plots (n = 31) in a mixed A. faxoniana and B. utilis forest in Sichuan, China. Dwarf bamboos died off in 1983 in the gap plots, and in three of the four forest plots. Forest development was assessed for the period 1984–1996. The seedling bank in forest and gap plots increased after bamboo die-off. A. faxoniana seedlings increased more than B. utilis in forest plots; the opposite pattern characterized gap plots. The proportion of seedlings on raised micro-sites on the forest floor also changed and new seedling were more abundant on the forest floor. By 1996, bamboo seedling cover and biomass had recovered to ca. 45% or their pre-flowering values. Rates of bamboo seedling recovery were faster beneath canopy gaps and deciduous trees than beneath forest or evergreen trees. Tree mortality exceeded recruitment in plots with dense bamboo; the opposite pattern was found in the plot with little bamboo. The mortality rate for B. utilis trees (2.4% year⁻¹) was higher than that for A. faxoniana (0.8% year⁻¹) and forests with dense bamboos became more open over the census period. Tree mortality was size-dependent and intermediate sized trees had the lowest rates of mortality. Stand basal area increased mainly due to greater basal area gain than loss for A. faxoniana. Interactions between tree species life history, canopy type, and bamboo life-cycles create heterogeneous conditions that influence tree and bamboo regeneration and contribute to the coexistence of A. faxoniana and B. utilis in old-growth forests in southwestern China.     

Stand Structure in Pine,Spruce and Deciduous Forests: A Comparison Between Finnish and Russian Karelia

This study investigated the stand structure in pine, spruce and deciduous forests in the border district of Finland and Russia. A total of 46 mature forest stands was selected as pairs, the members of each pair being as similar as possible with respect to their forest site type, age, moisture and topography. The stands were then compared between the two countries by means of basal areas and number of stems. The proportions of dominating tree species were 2-12% lower, and correspondingly the proportions of secondary tree species higher, in Russian forests. The density of the forest stock was also higher in each forest type in Russia. The forests in the two countries differed most radically in terms of the abundance of dead trees. The amount was two to four times higher in Russian deciduous and spruce forests, and in pine forests the difference was 10-fold. The stand structures indicated that Russian coniferous stands, in particular, were more heterogeneous than intensively managed pine and spruce stands in Finland.     

Spatial point-pattern analysis for detecting density-dependent competition in a boreal chronosequence of Alberta

In the boreal forest of Alberta, fire and wind often open gaps in the canopy where late-successional species can establish and over time cause a shift in the species distribution from deciduous (e.g., trembling aspen) dominated to mixedwood, to shade-tolerant conifer (e.g., white spruce) dominated stands. This study attempted to understand the change of density-dependent competition in a boreal chronosequence and the role of tree competition in affecting stand structure and mortality. Four 1-ha stem-mapped plots were established to represent a chronosequence comprised of aspen dominated, mixedwood, and spruce dominated stands in Alberta. Second order spatial point-pattern analysis using Ripley's K(t) function showed that intraspecific competition is a prevailing force causing conspecific tree mortality and thus shaping the stand structure. The results of bivariate K(t) function analysis did not reveal sufficient evidence of interspecific competition. This suggested that competitive interaction among heterospecific trees was not strong enough to cause significant tree mortality, but the analysis of marked correlation function revealed that interspecific competition could have a negative impact on tree growth. This study highlights the importance of density-dependent competition in understanding stand dynamics of boreal forests over succession.     

Tree regeneration in gap-understory mosaics in a subtropical Shorea robusta(Sal) forest

Spatial variation in tree-regeneration density is attributed to the specialization of tree species to light availability for germination and growth. Light availability,in turn, varies across the gap-understorey mosaic. Canopy gaps provide an important habitat for the regeneration of tree species that would otherwise be suppressed in the understory. In subtropical forests, there is still a knowledge-gap relating to how canopy disturbances influence tree-regeneration patterns at local scale, and if they disproportionately favor regeneration of certain species. We aim to analyze whether canopy gaps promote tree regeneration, and tree species are specialized to gaps or understory for germination and growth. We sampled vegetation in 128 plots(0.01 ha), equally distributed in gaps and below canopy, in two subtropical Shorea robusta Gaertn.(Sal) forests in Nepal, recording the number of tree seedlings and saplings in each plot. We compared the regeneration density of seedlings and saplings separately between gaps and the understorey. The mean densities of seedlings and saplings were higher in the gaps at both sites;although there was no difference in the seedling density of the majority of the species between the habitats. No species were confined to either gap or understorey at the seedling stage. We conclude that gaps are not critical for the germination of tree species in Sal forests but these are an important habitat for enabling seedlings to survive into saplings. The classification of trees into regeneration guilds mainly based on germination does not apply to the majority of tree species in subtropical Sal forests. Our results reaffirm that gap creation promotes tree regeneration by favouring seedling survival and growth and can influence forest management for conservation, as well as for plantations.