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.     

Regeneration growth in different light environments of mixed species, multiaged, mountainous forests of Romania

Growth of regenerating trees in different light environments was studied for the mountainous, mixed-species forests in the Carpathian Mountains of Romania. The primary species in these mixtures were silver fir (Abies alba Mill.), European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst). Seedlings/saplings of these species were selected and measured in different stands from two different geographical locations. Regenerating trees were measured for height and diameter growth during the summer of 2002. For each seedling/sapling, percentage of above canopy light (PACL) and stand basal area (BA) were used to assess available and occupied growing space respectively. Regeneration growth was compared against these two variables and regression relationships were developed. Using these models, we predicted the dynamics of regeneration as both growth and species composition. Our results showed that in low-light environments (PACL<20–35%; BA>30 m²/ha), shade tolerant fir and beech clearly outcompeted the spruce. Therefore, in dense stands, spruce could be eliminated by the shade tolerant species. For intermediate levels of cover (PACL=35–70%; BA=15–35 m²/ha) the spruce grew at comparable rates as the beech and fir. All three species showed similar growth rates in open conditions (PACL>80–90%; BA<15–20 m²/ha) with the spruce having a tendency to outgrow the others. However, in terms of establishment, such conditions favor spruce and inhibit fir and beech.     

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.     

Tree Regeneration and Sapling Damage of Pedunculate Oak Quercus robur in a Grazed Forest in Galicia, NW Spain: A Comparison of Continuous and Rotational Grazing Systems

Forest grazing has been recognised as being a useful tool in fire-risk reduction, in addition to having the potential to preserve or enhance forest biodiversity if managed correctly. Concern for natural regeneration of forest trees in Europe has also prompted interest in the effects of large herbivores on seedling and sapling growth and mortality. An investigation was carried out into sapling damage and density of natural regeneration of oak (Quercus robur) in a mature, pony-grazed, Pinus radiata forest in Galicia, NW Spain under two different grazing regimes (continuous and rotational). In all treatments significantly more oak seedlings and saplings were recorded in areas of grass sward than areas dominated by taller ground flora species. Damage to oak saplings was assessed from the form (height and canopy) relative to stem diameter. The height and average canopy diameter of similar-aged saplings were found to be significantly greater in ungrazed (control) than both continuous and rotationally grazed treatments. Height and canopy diameters of similar-aged oak were not significantly different between the two grazing treatments. Significant differences were observed in tree form, with unbrowsed saplings having the greatest height to canopy width ratio and those in the continuously browsed plots having the smallest. An obvious decrease in the goodness of fit (R²) of regression analyses were found in continuously grazed areas compared to rotational and control plots for both height and canopy data. The differences in damage observed were not significantly different enough to suggest one method of grazing over the other as being better for minimising sapling damage. Management requirements are more likely to dictate grazing regime. Overall, stock density is likely to have a more significant effect on damage than stocking system.     

Effect of natural disturbances on the abundance of Norway spruce (Picea abies (L.) Karst.) regeneration in nemoral forests of the southern boreal zone

The impact of natural disturbances on the canopy (trees ≥14 m high) and sapling stratum (>0.3 and ≤14 m high) composition was studied in nemoral old-growth forests located within the southern boreal zone in Central Russia (Central Forest Reserve, 32°29′–33°01′E, 56°26′–56°31′N). I hypothesized that the current disturbance regime does not allow the maintenance of current spruce abundance in the canopy, and, as a result, there is a continuous shift in the canopy composition towards a greater abundance of deciduous species. Three 300×20 m² transects were established to estimate the proportions of stand under non-closed unexpanded canopy gaps. Data on sapling composition of 49 canopy gaps were used to analyze pattern of gap refuting in these forests. Additionally, data from three forest inventories showed changes in canopy composition over a period from 1972 to 1990.

The current status of nemoral forests is characterized by the high proportion of stand area under treefall gaps (71%). The loss of spruce from the canopy caused by treefalls (53% of the total basal area of gap-makers) was slightly greater than its canopy abundance (45%). Canopy gaps of all sizes encouraged spruce regeneration which might be due to a decrease in sapling mortality and/or more active recruitment of spruce seedlings. After a gap was formed, the presence of spruce in sapling strata increased. However, within both small (<200 m² in size) and large (>200 m²) gaps, tall (>6 m) spruce saplings did not reach the level of its abundance in the tree canopy. In gaps, tall (>6 m) saplings of lime (Tilia cordata) and elm (Ulmus glabra) grew more quickly than those of spruce and maple. These data suggested a decrease in canopy spruce and an increase in deciduous species in the near future which supported the original hypothesis. Analysis of forest inventory records revealed similar changes in the canopy structure over the past two decades. However, the observed high proportion of stand area under gaps implies that for the next few decades large areas of nemoral communities will be occupied by relatively young stands. This may, in turn, decrease the frequency of large-scale treefalls revegetated mainly by deciduous saplings.     

闽楠天然次生林自然更新的影响因子研究

[目的]分析闽楠天然次生林自然更新与环境因子之间的关系,为其科学经营及保护提供参考依据。[方法]以江西省吉安市闽楠天然次生林为研究对象,通过标准地调查获取数据,运用多元数量化模型I建立闽楠幼树幼苗重要值与环境因子的关系模型,采用单因素方差分析单个环境因子对闽楠幼树幼苗更新的影响。[结果]研究表明:密度(闽楠下种母树株数、郁闭度、株数密度)、坡位、腐殖质层厚度、坡向、林下植被盖度、凋落物层厚度是影响闽楠天然次生林自然更新的主要因素,偏相关系数在0.325 7 0.715 7之间,t检验结果为极显著或显著;模型复相关系数为0.966,F检验结果为极显著(F=30.96~(**))。[结论]闽楠下种母树株数对其幼树幼苗的更新起着最主要的作用,而郁闭度与株数密度过高或过低、凋落物层越厚、腐殖质层越薄、林下植被盖度越大均不利于其自然更新,同时,半阴坡、下坡位条件下的闽楠幼树幼苗的更新好于其它坡向与坡位。为促进闽楠幼树幼苗的自然更新,林分中闽楠下种母树应保留200株·hm~(-2)以上,以及对郁闭度、林下植被盖度及凋落物层厚度等实施相应的调控措施。     

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.     

Removing aboveground vegetation facilitates survival but slows height growth of spruce saplings in a fenced, degraded sub-alpine forest in central Japan

The effects of removing aboveground vegetation on survival and growth of Hondo spruce (Picea jezoensis var. hondoensis) saplings were examined in a dense dwarf bamboo (Sasa nipponica) field inside a deer-proof fence built in a degraded sub-alpine forest on Mt Ohdaigahara, central Japan. All bamboo culms were removed within a radius of 1 m from each sapling in a plot of 0.6 ha (removal plot), whereas no culms were removed in two control plots of 0.1 ha. We measured the height, crown depth, and main shoot elongation over 1 year for all spruce saplings, culm height and cover ratios of dwarf bamboo growing around each spruce sapling, and the light levels above each sapling. Removal of dwarf bamboo regulated the culm height and bamboo coverage until the following summer such that light conditions improved. The mortality of saplings smaller than the average culm height was lower in the removal plot (0.89%) than in a control plot (5.9%). After bamboo removal, sapling height growth was reduced with increasing bamboo cover in the previous year. Complete removal of the aboveground portions of covering bamboo improved sapling survival while regulating elongation growth, possibly because of the sudden increase in light intensity. Thus, controlling the extent of dwarf bamboo removal might be necessary to facilitate the growth of spruce saplings to restore the forest.     

Light-related competitive effects of overstory trees on the understory conifer saplings in a subalpine forest

To clarify the mechanism by which overstory trees shade understory saplings, we investigated the relationships among light conditions of the saplings (measured as indirect site factor; ISF and direct site factor; DSF), the calculated competition effects of overstory trees on the saplings (W), and relative height growth rate of the saplings (RHGR). We calculated several W values in order to find a W value which can express the light conditions as appropriately as possible, and the results indicated that W explained only 21.9%–24.7% of the total variance of light conditions in the cases where W gave the best fit. In this study, W was calculated based on the basal areas of overstory trees. However, it is known that canopy structure also affects the light regimes in the forest understory, and this might yield the possible errors even within W representing the shading effects most adequately. Therefore, although W significantly represents the shading effect from overstory trees, a great proportion of the variance remained without being explained by W. RHGR was negatively correlated with W, and the W value which had the most adequate explanation of the shading effect also showed the best negative correlation with RHGR. This provides the evidence that the competitive effect of overstory trees on sapling growth is mediated by the shading effect, indicating that competition for light clearly exists within this forest. Such competition for light may closely relate to the well-known phenomenon of gap regeneration in subalpine forests in central Japan.     

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.

     

Site preference and occurrence patterns of <Emphasis Type="Italic">Picea jezoensis</Emphasis> and <Emphasis Type="Italic">Abies sachalinensis</Emphasis> on decayed logs in natural coniferous forests in Hokkaido,northern Japan

The objectives of this study were to investigate differences in the site preferences of seedlings of Picea jezoensis and Abies sachalinensis on decayed logs, and to examine the occurrence patterns of seedlings and saplings of the two species and whether they occur together or separately on logs. We characterized the habitats of 1–2-year-old seedlings of the two species on logs and examined the relationship of the two species on logs by growth stages in two plots. One plot had been disturbed about 50 years ago whereas the other had not for a long time. Although the thickness of moss and the litter layer in the habitats of 1–2-year-old seedlings were significantly different between the two species, the two species could occur together. In one study plot, seedlings and saplings of the two species occurred together. The initial occurrence pattern of the seedlings affected the occurrence patterns of the saplings on logs. The occurrence patterns of the seedlings and saplings of the two species on logs seemed to be affected by the abundance of seed trees. In the other study plot saplings of the two species occurred separately, but one species was not always competitively superior to the other species. Disturbance history affected the occurrence patterns of the saplings of the two species on decayed logs at the two study plots. Consequently, it is concluded that seed dispersal and the abundance of available logs, which are usually affected by disturbance, are significant factors in the natural regeneration of conifers in Hokkaido.     

Regeneration in experimental gaps of subalpine <Emphasis Type="Italic">Picea abies</Emphasis> forest in the Slovenian Alps

In 1998, we analysed regeneration success in four 20 year old medium-sized (600–800 m²) and four small experimental gaps (100–200 m²) in subalpine Norway spruce forest in Triglav National Park, Slovenia. We assessed the influence of site (depressions, slopes), gap size, and position within gap (centre, gap south, and north edge) on regeneration success and interactions between regeneration density, height, height increment, direct and diffuse light, ground vegetation, soil depth, thickness of humus horizons, presence of woody debris, and microrelief. Seedling density was higher in depressions, where more seedlings developed in central positions of gaps in both gap sizes, and in microsites with deeper soils, less competition from ground vegetation and convex microsites. On slopes, a higher seedling density was found in small gaps, and at the northern edge of gaps regardless of gap size. Here, seedling density was negatively influenced by diffuse radiation and positively by soil depth. The results suggest that regeneration should be initiated from medium to large gaps in depressions and small gaps on slopes. Development of established seedlings was enhanced by higher radiation levels on both sites, therefore favourable extension of the gaps would be lateral to northwards.     

Natural regeneration of beech (Fagus sylvatica L.) with respect to canopy density,soil moisture and soil carbon content

The purpose of the present study is to model the regeneration success using quantifiable growth factors. The hypothesis is that the regeneration success can be predicted from a model that incorporates canopy and ground flora leaf area indexes (LAI), soil water content and soil carbon content. In April 1992, 10 plots (1 m²) were established in each of 22 Danish beech stands with natural regeneration originating from the 1989 seed fall. The regeneration success was investigated until autumn 1994. In each of the 220 plots the light conditions, soil water supply and soil carbon content were quantified. The survival was high during the study period. From spring 1992 to autumn 1993 the mean number of saplings decreased from 58 m⁻² to 54 m⁻². The sapling number was reduced by increased canopy density but it was neither influenced by water supply nor by soil carbon content. The height growth variation was evident. After the fifth growing season (autumn 1994) the height of the tallest sam3pling per plot ranged from 7 to 120 cm. Generally, an increased canopy opening increased the potential for and variance of height growth. Increased soil water content caused a significant increase in the growth of the saplings; whereas, a significant reduction in growth occurred with increased soil carbon content. The shading from the competing ground flora had no significant effect. However, the contribution to the total sum of squares from the quantitative variables of the relevant statistical models was only 3%–13%. Therefore, the models were unable to predict regeneration success satisfactorily and it was not possible to establish the causal effects underlying silvicultural treatments and site. It is concluded that the beech saplings probably were so well established at the beginning of the study that they had become tolerant to poor growing conditions, such as the drought in 1992. This indicates that the growing conditions during the early phases of seedling establishment may be crucial for the regeneration success. For silviculturists, these conclusions stress the importance of creating favourable growth conditions prior to the first growing season. Future research should examine the early phases of seedling establishment by quantifying growth factors in more detail than was done in the present study.     

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.     

Predicting tree regeneration in <Emphasis Type="Italic">Picea abies</Emphasis> snag stands

A bark beetle (Ips typographus) infestation caused the death of almost all Norway spruce (Picea abies) trees in a mountain forest in the Swiss Alps. We developed a tree regeneration model, ‘RegSnag’ (=REGeneration in a SNAG stand), to project the future amount and height of tree regeneration in these snag stands. The model combines a height-class structured tree module with a microsite-based module of snag decay and ground-vegetation succession. Microsite-specific rates of germination, mortality and height growth were modelled for four tree species (Picea abies, Sorbus aucuparia, Acer pseudoplatanus and Betula pendula) in eight height classes (from seedlings to saplings 5 m tall) and on 26 microsite types (e.g. moss, grass). Model tests with independent field data from 8 years after the Picea die-back demonstrated that microsites had a considerable effect on the development of tree regeneration on both the montane and the subalpine level. With microsite-specific parameters, the height and frequency of Picea in each microsite could be simulated more accurately than without considering microsite effects (e.g. bias of 8 vs. 119 saplings ha⁻¹ on the montane level). Results of simulations 40 years into the future suggest that about 330–930 Picea saplings per ha out of those that germinated in 1994 and 1996 will reach a height of 5 m within 30–35 years after Picea die-back. This is due to differences in seed inflow and browsing intensities. Picea and not Betula or Sorbus trees will replace the current herbaceous vegetation in these snag stands.     

Snow gliding and loading under two different forest stands: a case study in the north-western Italian Alps

The presence of a thick snowpack could interfere with forest stability, especially on steep slopes with potential damages for young and old stands. The study of snow gliding in forests is rather comple...     

Weeds in <Emphasis Type="Italic">Eucalyptus globulus</Emphasis> subsp. <Emphasis Type="Italic">maidenii</Emphasis> (F. Muell) establishment: effects of competition on sapling growth and survivorship

The effect of herbaceous vegetation on growth and survival was assessed in planted eucalyptus saplings grown under four levels of weed cover. Seedlings of Eucalyptus globulus subsp. maidenii were planted with 0 (W0), 25 (W25), 50 (W50) and 100% (W100) weed cover. Weed species composition and aboveground biomass was determined. Soil water content was evaluated by the gravimetric method. Seedling leaf area, diameter and height were evaluated at planting and during the 3 months following establishment. Tree height, diameter and stem volume was estimated at 12, 24 and 36 months. First year tree survival was recorded. The ratio of cumulative stem growth under W100: cumulative stem growth under W0 was used to measure competitive performance. Regression analysis was used to determine competition thresholds. Weeds seriously threaten the growth of E. globulus subsp. maidenii. Early negative effects of competition on growth were evident as from the second month after establishment, and both seedlings and 1-year-old saplings were more affected than 2- and 3-year-old ones. A “minimum-response threshold” was determined near 500 kg/ha (corresponding to W25 cover). No clear effect was found between sapling survival and weed biomass.     

Predicting regeneration establishment in Norway spruce plantations using a multivariate multilevel model

This study predicts the regeneration establishment on 3-year-old Norway spruce (Picea abies (L.) Karst.) plantations in southern Finland using regeneration survey data. Regeneration establishment was described by seven response variables: number of planted spruces, natural Scots pines (Pinus sylvestris L.), natural spruces, natural seed-origin birches (Betula pubescens Ehrh. and B. pendula Roth.) and other broadleaves (i.e. sprout-origin birches and other broadleaves than birch), as well as height of crop-tree spruce and dominant height of broadleaves. Due to the multivariate (several responses for each plot) and multilevel (plot, stand, municipality, forest centre) structure, regeneration establishment was modelled by fitting a multivariate multilevel model with explanatory variables such as temperature sum, site fertility, soil quality and method of site preparation. In the model, the numbers of tree seedlings were modelled using over-dispersed Poisson distributed equations, and the tree heights were modelled using normally distributed linear equations. The estimated fixed and random parameters of the equations were logical, and there was no serious bias in predicting the regeneration establishment in the independent test data set. This modelling approach can be used to predict the regeneration establishment stochastically by taking into account the large unexplained variation in regeneration models.     

Factors influencing germination and establishment of Eucalyptus pauciflora near the alpine tree line

Factors affecting seed germination, seedling establishment and growth to reproductive maturity in Eucalyptus pauciflora Sieb. ex Spreng. at the tree line in southeastern Australia were investigated. Timing of germination was determined principally by temperature but could be strongly modified by soil water. Mortality of germinants was heaviest during the first growing season (dependent on soil water) and the first winter (dependent on the depth and duration of snow cover). Neither plant nor soil treatments with insecticide or fungicide had an overriding or consistent effect on germination or seedling establishment. The probability of reaching the sapling stage was greater for planted seedlings than for plants germinated on site, indicating the dependence of survival on accumulated reserves. There was no clear indication of a general decrease in survival with increasing altitude, because of an interplay between elevation and site factors including competing vegetation, snow depth and period of snow cover. Five-year-old saplings planted up to 200 m above the tree line produced reproductive structures and viable seed between the ages of 13 and 18 years. The relevance of the results to an understanding of the location of the present tree line is discussed.